Ontogeny of limb mass distribution in infant baboons (Papio cynocephalus)

Primates have more distally distributed limb muscle mass compared to most nonprimate mammals. The heavy distal limbs of primates are likely related to their strong manual and pedal grasping abilities, and interspecific differences in limb mass distributions among primates are correlated with the amount of time spent on arboreal supports. Within primate species, individuals at different developmental stages appear to differ in limb mass distribution patterns. For example infant macaques have more distally distributed limb mass at young ages. A shift from distal to proximal limb mass concentrations coincides with a shift from dependent travel (grasping their mother's hair) to independent locomotion. Because the functional demands placed on limbs may differ between taxa, understanding the ontogeny of limb mass distribution patterns is likely an essential element in interpreting the diversity of limb mass distribution patterns present in adult primates. This study examines changes in limb inertial properties during ontogeny in a longitudinal sample of infant baboons (Papio cynocephalus). The results of this study show that infant baboons undergo a transition from distal to proximal limb mass distribution patterns. This transition in limb mass distribution coincides with the transition from dependent to independent locomotion during infant development. Compared to more arboreal macaques, infant baboons undergo a faster transition to more proximal limb mass distribution patterns. These results suggest that functional demands placed on the limbs during ontogeny have a strong impact on the development of limb mass distribution patterns.     

Limb growth in captive Galago senegalensis: getting in shape to be an adult

Since primate infants are not simply miniature adults, adult shape results from differential growth patterns of individual body segments. Initially an infant relies on its mother for transportation, and later begins independent locomotion. Skeletal growth patterns must meet the functional demands of independent locomotion. In this study we sought to determine whether Galago senegalensis braccatus follow the general primate pattern of decreasing intermembral index (IMI) throughout ontogeny. We also asked whether ontogenetic attainment of adult limb proportions coincides with attainment of independent locomotion, i.e., do infants reach adult limb proportions near the time they begin independent locomotion (approximately 7 weeks of age)? Mixed‐longitudinal data were taken from a sample of 10 captive‐born Galago senegalensis. Linear lengths of the trunk, arm, forearm, thigh, and leg were measured in the animals from birth until they were approximately 500 days old. The IMI and the ratio of each limb segment to both trunk length and the cube root of body mass were calculated. The results of a Mann‐Whitney Wilcoxon rank‐sum test for unmatched samples indicate that G. senegalensis do exhibit the primate pattern of decreasing IMI throughout ontogeny, and that the IMIs of infants at the time of initial locomotor independence are significantly higher than those of adult IMIs. Some (but not all) measures of relative limb lengths differed between neonates or 7‐week‐old infants and adults. Therefore, the hypothesis that infants acquire adult limb proportions by the time they begin independent locomotion is not supported by this study. The current results indicate that ontogenetic shape changes in galagos are a complex process and apparently cannot be explained by simple initial locomotor competency. Am. J. Primatol. 69:103–111, 2007. © 2006 Wiley‐Liss, Inc.     

Catabolic capacity of the muscles of shorebird chicks: maturation of function in relation to body size

Newly hatched precocial chicks of arctic shorebirds are able to walk and regulate their body temperatures to a limited extent. Yet, they must also grow rapidly to achieve independence before the end of the short arctic growing season. A rapid growth rate may conflict with development of mature function, and because of the allometric scaling of thermal relationships, this trade-off might be resolved differently in large and small species. We assessed growth (mass) and functional maturity (catabolic enzyme activity) in leg and pectoral muscles of chicks aged 1-16 d and adults of two scolopacid shorebirds, the smaller dunlin (Calidris alpina: neonate mass 8 g, adult mass 50 g) and larger whimbrel (Numenius phaeopus; neonate mass 34 g, adult mass 380 g). Enzyme activity indicates maximum catabolic capacity, which is one aspect of the development of functional maturity of muscle. The growth rate-maturity hypothesis predicts that the development of catabolic capacity should be delayed in faster-growing muscle masses. Leg muscles of both species were a larger proportion of adult size at hatching and grew faster than pectoral muscles. Pectoral muscles grew more rapidly in the dunlin than in the whimbrel, whereas leg muscles grew more rapidly in the whimbrel. In both species and in both leg and pectoral muscles, enzyme activities generally increased with age, suggesting increasing functional maturity. Levels of citrate synthase activity were similar to those reported for other species, but l-3-hydroxyacyl-CoA-dehydrogenase and pyruvate kinase (PK) activities were comparatively high. Catabolic capacities of leg muscles were initially high compared to those of pectoral muscles, but with the exception of glycolytic (PK) capacities, these subsequently increased only modestly or even decreased as chicks grew. The earlier functional maturity of the more rapidly growing leg muscles, as well as the generally higher functional maturity in muscles of the more rapidly growing dunlin chicks, contradicts the growth rate-maturity function trade-off and suggests that birds have considerable latitude to modify this relationship. Whimbrel chicks, apparently, can rely on allometric scaling of power requirements for locomotion and the thermal inertia of their larger mass to reduce demands on their muscles, whereas dunlin chicks require muscles with higher metabolic capacity from an earlier age. Thus, larger and smaller species may adopt different strategies of growth and tissue maturation.     

Muscle architectural properties in the common marmoset (<Emphasis Type="Italic">Callithrix jacchus</Emphasis>)

The common marmoset, Callithrix jacchus, is a small New World monkey that has recently gained attention as an important experimental animal model in the field of neuroscience as well in rehabilitative and regenerative medicine. This attention reflects the closer phylogenetic relationship between humans and common marmosets compared to that between humans and other experimental animals. When studying the neuronal mechanism behind various types of neurological motor disorders using the common marmoset, possible differences in muscle parameters (e.g., the force-generating capacity of each of the muscles) between the common marmoset and other animals must be taken into account to permit accurate interpretation of observed motor behavior. Differences in the muscle architectural properties are expected to affect biomechanics, and hence to affect neuronal control of body movements. Therefore, we dissected the forelimbs and hind limbs of two common marmosets, including systematic analysis of the muscle mass, fascicle length, and physiological cross-sectional area (PCSA). Comparisons of the mass fractions and PCSA fractions of the forelimb and hind limb musculature among the common marmoset, human, Japanese macaque, and domestic cat demonstrated that the overall muscle architectural properties of the forelimbs and hind limbs in the common marmoset are very similar to those of the Japanese macaque, a typical quadrupedal primate. However, muscle architectural properties of the common marmoset differ from those of the domestic cat, which has relatively larger hamstrings and pedal digital flexor muscles. Compared to humans, the common marmoset exhibits relatively smaller shoulder protractor, retractor, and abductor muscles and larger elbow extensor and rotator-cuff muscles in the forelimb, and smaller plantarflexor muscles in the hind limb. These differences in the muscle architectural properties must be taken into account when interpreting motor behaviors such as locomotion and arm-reaching movements in the common marmoset.     

Architectural and histochemical diversity within the quadriceps femoris of the brown lemur (Lemur fulvus)

Physiologically related features of muscle morphology are considered with regard to functional adaptation for locomotor and postural behavior in the brown lemur (Lemur fulvus). Reduced physiological cross-sectional area, estimated maximum excursion of the tendon of insertion, length of tendon per muscle fasciculus, and areal fiber type composition were examined in the quadriceps femoris in order to assess the extent of a "division of labor" among four apparent synergists. Each of these four muscles in this prosimian primate displays a distinguishing constellation of morphological features that implies functional specialization during posture and normal locomotion (walk/run, galloping, leaping). Vastus medialis is best suited for rapid whole muscle recruitment and may be reserved for relatively vigorous activities such as galloping and leaping (e.g., small cross-sectional area per mass, long excursion, predominance of fast-low oxidative fibers, relatively little tendon per fasciculus). In theory, rectus femoris could be employed isometrically in order to store elastic strain energy during all phasic activities (e.g., large cross-sectional area per mass, short excursion, predominance of fast-high oxidative fibers, large amount of tendon per fasciculus). Vastus intermedius exhibits an overall morphology indicative of a typical postural muscle (e.g., substantial cross-sectional area, short excursion, predominance of slow-high oxidative fibers, large amount of tendon per fasciculus). The construction of vastus lateralis reflects an adaptation for high force, relatively high velocity, and resistance to fatigue (e.g., large cross-sectional area, long excursion, most heterogeneous distribution of fiber types, large amount of tendon per fasciculus); this muscle is probably the primary contributor to a wide range of locomotor behaviors in lemurs. Marked dramatic architectural disparity among the four bellies, coupled with relative overall fiber type heterogeneity, suggests the potential for exceptional flexibility in muscle recruitment within this mass. One interpretation of this relatively complex neuromuscular organization in the brown lemur is that it represents an adaptation for the exploitation of a three-dimensional arboreal environment by rapid quadrupedalism and leaping among irregular and spatially disordered substrates.     

A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth

The large theropod dinosaur Tyrannosaurus rex underwent remarkable changes during its growth from <10 kg hatchlings to >6000 kg adults in <20 years. These changes raise fascinating questions about the morphological transformations involved, peak growth rates, and scaling of limb muscle sizes as well as the body's centre of mass that could have influenced ontogenetic changes of locomotion in T. rex. Here we address these questions using three-dimensionally scanned computer models of four large, well-preserved fossil specimens as well as a putative juvenile individual. Furthermore we quantify the variations of estimated body mass, centre of mass and segment dimensions, to characterize inaccuracies in our reconstructions. These inaccuracies include not only subjectivity but also incomplete preservation and inconsistent articulations of museum skeletons. Although those problems cause ambiguity, we conclude that adult T. rex had body masses around 6000-8000 kg, with the largest known specimen ("Sue") perhaps ～9500 kg. Our results show that during T. rex ontogeny, the torso became longer and heavier whereas the limbs became proportionately shorter and lighter. Our estimates of peak growth rates are about twice as rapid as previous ones but generally support previous methods, despite biases caused by the usage of scale models and equations that underestimate body masses. We tentatively infer that the hindlimb extensor muscles masses, including the large tail muscle M. caudofemoralis longus, may have decreased in their relative size as the centre of mass shifted craniodorsally during T. rex ontogeny. Such ontogenetic changes would have worsened any relative or absolute decline of maximal locomotor performance. Regardless, T. rex probably had hip and thigh muscles relatively larger than any extant animal's. Overall, the limb "antigravity" muscles may have been as large as or even larger than those of ratite birds, which themselves have the most muscular limbs of any living animal.     

The skeleton of early Eocene Cantius, oldest lemuriform primate

A recently discovered partial skeleton of the adapid Cantius trigonodus from the early Eocene Willwood Formation of the Bighorn Basin, Wyoming, documents substantial new information about the anatomy of the oldest lemuriform primates. It is very similar in all features to its descendant, middle Eocene Notharctus, and both exhibit numerous resemblances to certain extant Malagasy lemurs, particularly Lepilemur, Propithecus, Lemur, and Hapalemur griseus. Like these forms, Cantius had relatively long hind limbs and short forelimbs. Forelimb traits (prominent brachialis flange of the humerus, well-developed olecranon process of the ulna, and strong shafts of the ulna and radius) suggest active use of the forelimbs in progression. Specializations in the hind limb (e.g., expanded articular surface of the femoral head, narrow and elevated patellar trochlea and prominent lateral trochlear ridge, posteriorly oriented femoral and tibial condyles, narrow and elongate talus, and hallucal metatarsal with prominent peroneal tubercle) indicate capabilities for leaping and for powerful grasping with an opposable hallux. Cantius was presumably primitive in having a relatively long ischium and much more distal inferior tibial tuberosity than most extant lemurs--traits suggesting that powerful extension of the thigh and flexion at the knee were important in its locomotion and posture. We interpret Cantius as an active arboreal quadruped with a propensity for leaping. The existence of this skeletal structure in one of the oldest primates of modern aspect suggests that it represents the primitive lemuriform morphology.     

Takeoff and landing forces of leaping strepsirhine primates.

Knowledge of the forces animals generate and are exposed to during locomotion is an important prerequisite for understanding the musculoskeletal correlates of locomotor modes. We recorded takeoff and landing forces for 14 animals representing seven species of strepsirhine primates with a compliant force pole. Our sample included both specialized vertical clingers and leapers and more generalized species. Takeoff forces are higher than landing forces. Peak forces during acceleration for takeoff ranged from 6 to 12 times body weight, and the peak impact forces at landing are between 5 and 9 times body weight. There is a size-related trend in peak force magnitudes. Both takeoff and landing forces decrease with increasing body size in our sample of animals from 1 kg to over 5 kg. Peak forces increase with distance leapt. The distance effect is less clear, probably due to the narrow range of distances represented in our sample. A comparison of subadult and adult animals of two species of sifakas reveals a tendency for the young animals to exert relatively higher peak forces in comparison to their adult conspecifics. Finally, Lemur catta and Eulemur rubriventer, the "generalists" in our sample, tend to generate higher forces for equal tasks than the specialized vertical clingers and leapers (i.e., the indriids and Hapalemur).A broad-scale comparison of peak leaping forces and peak forces for quadrupedal and bipedal walking and running shows that leaping at small body size is associated with exceptionally high forces. Whereas relative forces (i.e., forces divided by body weight) decrease with increasing body mass for leaping, forces for walking and running do not change much with size. Leaping forces in our sample scale to (mass)(-1/3), which is consistent with expectations derived from geometric similarity models. Forces associated with other locomotor activities do not appear to follow this pattern. The very high forces found in strepsirhine leapers do not seem to be matched by bone robusticity beyond that documented for quadrupedal species.     

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.     

Interlimb coordination and gait in the brown lemur (Lemur fulvus) and the talapoin monkey (Miopithecus talapoin)

Patterns of interlimb coordination based on telemetered electromyography of extensor muscles are described for the brown lemur (Lemur fulvus) and the talapoin monkey (Miopithecus talapoin) in order to address the issue of possible motor programs for quadrupedal stepping in primates. Differences in modal patterns of ipsilateral limb coupling (phase intervals) between walking and galloping indicate that gait-specific programs do exist in primates, especially for symmetrical gaits. These preferred patterns distinguish primates from most other mammals (e.g., the domestic cat), but do not rule out the possibility of subtle differences among primates in species-specific mechanisms of neural control. Variability about the preferred modes is better interpreted as an expression of the flexibility or facultative capabilities of the neural mechanisms controlling locomotion than as “errors” in the motor program.     

Evolution of the axial system in craniates: morphology and function of the perivertebral musculature

The axial musculoskeletal system represents the plesiomorphic locomotor engine of the vertebrate body, playing a central role in locomotion. In craniates, the evolution of the postcranial skeleton is characterized by two major transformations. First, the axial skeleton became increasingly functionally and morphologically regionalized. Second, the axial-based locomotion plesiomorphic for craniates became progressively appendage-based with the evolution of extremities in tetrapods. These changes, together with the transition to land, caused increased complexity in the planes in which axial movements occur and moments act on the body and were accompanied by profound changes in axial muscle function. To increase our understanding of the evolutionary transformations of the structure and function of the perivertebral musculature, this review integrates recent anatomical and physiological data (e.g., muscle fiber types, activation patterns) with gross-anatomical and kinematic findings for pivotal craniate taxa. This information is mapped onto a phylogenetic hypothesis to infer the putative character set of the last common ancestor of the respective taxa and to conjecture patterns of locomotor and muscular evolution. The increasing anatomical and functional complexity in the muscular arrangement during craniate evolution is associated with changes in fiber angulation and fiber-type distribution, i.e., increasing obliqueness in fiber orientation and segregation of fatigue-resistant fibers in deeper muscle regions. The loss of superficial fatigue-resistant fibers may be related to the profound gross anatomical reorganization of the axial musculature during the tetrapod evolution. The plesiomorphic function of the axial musculature -mobilization- is retained in all craniates. Along with the evolution of limbs and the subsequent transition to land, axial muscles additionally function to globally stabilize the trunk against inertial and extrinsic limb muscle forces as well as gravitational forces. Associated with the evolution of sagittal mobility and a parasagittal limb posture, axial muscles in mammals also stabilize the trunk against sagittal components of extrinsic limb muscle action as well as the inertia of the body's center of mass. Thus, the axial system is central to the static and dynamic control of the body posture in all craniates and, in gnathostomes, additionally provides the foundation for the mechanical work of the appendicular system.     

Maternal encouragement in nonhuman primates and the question of animal teaching

Most putative cases of teaching in nonhuman animals involve parent-offspring interactions. The interpretation of these cases, particularly with regard to the cognitive processes involved, is controversial. Qualitative and quantitative observations made in nonhuman primates suggest that, in some species, mothers encourage their infants’ independent locomotion and that encouragement can be considered a form of instruction. In macaques, experience in raising previous offspring accounts in part for variability between mothers in propensity to encourage infant motor skills. Parsimony suggests that the cognitive mechanisms underlying maternal encouragement of infant locomotion in primates as well as some other putative cases of animal teaching may involve first-order intentionality (i.e., goal-directed behavior) and not higher cognitive processes such as attribution of knowledge/ignorance or perspective-taking. Encouragement of infant independent locomotion early in life may have benefits to mothers later on, in terms of reduction of costs of infant carrying, earlier infant weaning, and increased probability of reproduction in the mating season. The elementary forms of teaching observed in nonhuman primates may have played an important role in the origin and evolution of human culture.     

Spatial and temporal patterns of muscle cleavage in the chick thigh and their value as criteria for homology

Regions of lower cell density, called cleavage zones, emerge within the dorsal and ventral muscle masses in the vertebrate limb to separate distinct muscles. In the chick thigh, the stereotyped patterns of separation have been broadly outlined, but differences in interpretation exist because no criteria for separation have been defined, and the tissues of the limb are indistinct early in development. We have examined the cleavage process using modern applications of light microscopy and immunocytochemistry to completely detail the spatial and temporal progression of cleavage in stage 27-32 embryos. We find that each muscle has a complex but characteristic pattern of separation along the proximodistal axis. The complex pattern of separation is not related to the positions of muscles within the thigh, locations of blood vessels, activity patterns of muscles, or innervation patterns. The initial separation patterns are more straightforward than later separations and may be of value in determining the phylogenetic history of limb muscles since the same patterns are common to many tetrapods. Our detailed documentation clarifies the ontogeny of the thigh musculature and reveals more complex separation patterns between muscles than previously described.     

Transneuronal induction of muscle atrophy in grasshoppers

Autotomy is a process in grasshoppers whereby one or both hindlimbs can be shed to escape a predator or can be abandoned if damaged. It occurs between the trochanter and the femur (second and third leg segments) and once lost, the legs never regenerate. Autotomy severs branches of the leg nerve (N5) but damages no muscles since none span the autotomy plane. We find, however, that undamaged muscles intrinsic to the thorax of grasshoppers, Barytettix psolus, atrophy to less than 15% of their normal mass after autotomy of a hindlimb. These muscles operate the coxa and trochanter (first and second leg segments) and are innervated by branches of nerves 3 and 4; nerve branches that are not damaged by autotomy. Atrophy is localized to the side and body segment where autotomy occurs. Atrophy is evident 7-10 days after loss of a limb, is complete by about 30 days, and follows a similar time course whether induced in young adult, or sexually mature grasshoppers. During autotomy, leg nerve 5 is served distal to the trochanter, the thoracic muscles lose their normal static and dynamic load, and these muscles are subsequently no longer used to support the weight of the insect during posture and locomotion. Experimental loading and unloading of the affected muscles, and cutting of nerves indicated that it is the severing of leg nerve 5 during autotomy that transneuronally induces muscle atrophy.     

Muscle development in the grasshopper embryo. I. Muscles, nerves, and apodemes in the metathoracic leg

Much is known about the development of nerve pathways in the metathoracic limb bud of the grasshopper embryo. In this series of three papers, we report on the development of muscles in the same embryonic appendage. In a fourth paper (E. E. Ball, R. K. Ho, and C. S. Goodman, 1985, J. Neurosci, in press) we examine the development of specific neuromuscular connections for one of these muscles (coxal muscle 133a). In this first paper, we present an overview of the development of muscles, nerves, and apodemes (tendons). We previously reported on a class of large mesodermal cells, called muscle pioneers (MPs), that arises early in development and appears to act as a scaffold for developing muscles and guidance cue for motoneuron growth cones (R. K. Ho, E. E. Ball, and C. S. Goodman, 1983, Nature (London) 301, 66-69). We have used the I-5 monoclonal antibody (which specifically labels the MPs as well as the nerve pathways), HRP immunocytochemistry, and Normarski optics to visualize muscle, nerve, and apodeme development in the embryonic metathoracic limb bud from 27.5% (before the appearance of the MPs) to 55% (after the muscles have attained their basic adult pattern). Cell fusions, cell migration, and cell death all appear to play important roles in the development of MPs. The patterns of muscle development vary greatly, ranging from (i) single MPs for simple muscles (which in the adult have only one bundle of muscle fibers, e.g., coxal muscle 133a), to (ii) arrays of MPs for complex muscles [which in the adult have many bundles of muscle fibers each with separate sites of insertion, e.g., the extensor tibiae (ETi) and flexor tibiae (FlTi) muscles in the femur].     

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.     

Modular neuromuscular control of human locomotion by central pattern generator

The central pattern generators (CPG) in the spinal cord are thought to be responsible for producing the rhythmic motor patterns during rhythmic activities. For locomotor tasks, this involves much complexity, due to a redundant system of muscle actuators with a large number of highly nonlinear muscles. This study proposes a reduced neural control strategy for the CPG, based on modular organization of the co-active muscles, i.e., muscle synergies. Four synergies were extracted from the EMG data of the major leg muscles of two subjects, during two gait trials each, using non-negative matrix factorization algorithm. A Matsuoka׳s four-neuron CPG model with mutual inhibition, was utilized to generate the rhythmic activation patterns of the muscle synergies, using the hip flexion angle and foot contact force information from the sensory afferents as inputs. The model parameters were tuned using the experimental data of one gait trial, which resulted in a good fitting accuracy (RMSEs between 0.0491 and 0.1399) between the simulation and experimental synergy activations. The model׳s performance was then assessed by comparing its predictions for the activation patterns of the individual leg muscles during locomotion with the relevant EMG data. Results indicated that the characteristic features of the complex activation patterns of the muscles were well reproduced by the model for different gait trials and subjects. In general, the CPG- and muscle synergy-based model was promising in view of its simple architecture, yet extensive potentials for neuromuscular control, e.g., resolving redundancies, distributed and fast control, and modulation of locomotion by simple control signals.     

Density of muscle spindle profiles in the intrinsic forelimb muscles of the dog

The concept of parallel muscle combinations, in which spindle density is significantly higher in small muscles compared to their larger counterparts in large-small muscle combinations acting across a joint, is supported by the results of this study regardless of the joint. Analysis of the canine data as well as previously published guinea pig forelimb and human pelvic limb data revealed no significant difference in spindle density between antigravity and non-antigravity muscles. Furthermore, a gradual increase in spindle density from proximal to distal on the limb was not found, although spindle density was significantly higher in the intrinsic manus or pes muscles compared to more proximal limb muscles in all three species. The significant differences in spindle densities in parallel muscle combinations and in manus/pes versus proximal muscles are discussed relative to their possible role in the control of locomotion.