Martin S. Fischer

Dogs are an interesting object of investigation because of the wide range of body size, body mass, and physique. In the last several years, the number of clinical and biomechanical studies on dog locomotion has increased. However, the relationship between body structure and joint load during locomotion, as well as between joint load and degenerative diseases of the locomotor system (e.g. dysplasia), are not sufficiently understood. In vivo measurements/records of joint forces and loads or deep/small muscles are complex, invasive, and sometimes ethically questionable. The use of detailed musculoskeletal models may help in filling that knowledge gap. We describe here the methods we used to create a detailed musculoskeletal model with 84 degrees of freedom and 134 muscles. Our model has three key-features: Three-dimensionality, scalability, and modularity. We tested the validity of the model by identifying forelimb muscle synergies of a beagle at walk. We used inverse dynamics and static optimization to estimate muscle activations based on experimental data. We identified three muscle synergy groups by using hierarchical clustering. Predicted activation patterns exhibited good agreement with experimental data for most of the forelimb muscles. We expect that our model will speed up the analysis of how body size, physique, agility, and disease influence joint neuronal control and loading in dog locomotion.

This work demonstrates a neuromechanical model of rat hindlimb locomotion undergoing nominal walking with perturbations. In the animal, two types of responses to perturbations are observed: resetting and non-resetting deletions. This suggests that the animal locomotor system contains a memory-like organization. To model this phenomenon, we built a synthetic nervous system that uses separate rhythm generator and pattern formation layers to activate antagonistic muscle pairs about each joint in the sagittal plane. Our model replicates the resetting and non-resetting deletions observed in the animal. In addition, in the intact (i.e., fully afferented) rat walking simulation, we observe slower recovery after perturbation, which is different from the deafferented animal experiment. These results demonstrate that our model is a biologically feasible description of some of the neural circuits in the mammalian spinal cord that control locomotion, and the difference between our simulation and fictive motion shows the importance of sensory feedback on motor output. This model also demonstrates how the pattern formation network can activate muscle synergies in a coordinated way to produce stable walking, which motivates the use of more complex synergies activating more muscles in the legs for three-dimensional limb motion.

The first high-precision 3D in vivo hindlimb kinematic data to be recorded in normal dogs of four different breeds (Beagle, French bulldog, Malinois, Whippet) using biplanar, high-frequency fluoroscopy combined with a 3D optoelectric system followed by a markerless XROMM analysis (Scientific Rotoscoping, SR or 3D-2D registration process) reveal a) 3D hindlimb kinematics to an unprecedented degree of precision and b) substantial limitations to the use of skin marker-based data. We expected hindlimb kinematics to differ in relation to body shape. But, a comparison of the four breeds sets the French bulldog aside from the others in terms of trajectories in the frontal plane (abduction/adduction) and long axis rotation of the femur. French bulldogs translate extensive femoral long axis rotation (>30°) into a strong lateral displacement and rotations about the craniocaudal (roll) and the distal-proximal (yaw) axes of the pelvis in order to compensate for a highly abducted hindlimb position from the beginning of stance. We assume that breeds which exhibit unusual kinematics, especially high femoral abduction, might be susceptible to a higher long-term loading of the cruciate ligaments.

The increase in locomotor and metabolic performance during mammalian evolution was accompanied by the limitation of the number of cervical vertebrae to only seven. In turn, nuchal muscles underwent a reorganization while forelimb muscles expanded into the neck region. As variation in the cervical spine is low, the variation in the arrangement of the neck muscles and their attachment sites (i.e., the variability of the neck’s musculoskeletal organization) is thus proposed to be an important source of neck disparity across mammals. Anatomical network analysis provides a novel framework to study the organization of the anatomical arrangement, or connectivity pattern, of the bones and muscles that constitute the mammalian neck in an evolutionary context. Neck organization in mammals is characterized by a combination of conserved and highly variable network properties. We uncovered a conserved regionalization of the musculoskeletal organization of the neck into upper, mid and lower cervical modules. In contrast, there is a varying degree of complexity or specialization and of the integration of the pectoral elements. The musculoskeletal organization of the monotreme neck is distinctively different from that of therian mammals. Our findings reveal that the limited number of vertebrae in the mammalian neck does not result in a low musculoskeletal disparity when examined in an evolutionary context. However, this disparity evolved late in mammalian history in parallel with the radiation of certain lineages (e.g., cetartiodactyls, xenarthrans). Disparity is further facilitated by the enhanced incorporation of forelimb muscles into the neck and their variability in attachment sites.

Recently, algorithms were developed to track radiopaque markers in the heart fully automated. However, the methodology did not allow to assign the exact anatomical location to each marker. In this case study we describe the steps from the generation of three-dimensional marker coordinates to quantitative data analyses in an in vivo ovine model. In one adult sheep, twenty silver balls were sutured to the right side of the heart: 10 to the tricuspid annulus, one to the anterior tricuspid leaflet and nine to the epicardial surface of the right ventricle. In addition, 13 cylindrical tantalum markers were implanted into the left ventricle. Data were acquired with a biplanar X-ray acquisition system (Neurostar R, Siemens AG, 500 Hz). Radiopaque marker coordinates were determined fully automated using novel tracking algorithms. The anatomical marker locations were identified using a 3-dimensional model of a single frame containing all tracked markers. First, cylindrical markers were manually separated from spherical markers, thus allowing to distinguish right from left heart markers. The fast moving leaflet marker was identified by using video loops constructed of all recorded frames. Rotation of the 3-dimensional model allowed the identification of the precise anatomical position for each marker. Data sets were then analyzed quantitatively using customized software. The method presented in this case study allowed quantitative data analyses of radiopaque cardiac markers that were tracked fully automated with high temporal resolution. However, marker identification still requires substantial manual work. Future improvements including the implication of marker identification algorithms and data analysis software could allow almost real-time quantitative analyses of distinct cardiac structures with high temporal and spatial resolution.

Theria (marsupials and placental mammals) are characterized by a highly mobile pectoral girdle in which the scapula has been shown to be an important propulsive element during locomotion. Shoulder function and kinematics are highly conservative during locomotion within quadrupedal therian mammals. In order to gain insight into the functional morphology and evolution of the pectoral girdle of the two-toed sloth we here analyze the anatomy and the three-dimensional (3D) pattern of shoulder kinematics during quadrupedal suspensory ('upside-down') locomotion.

Talking about legged locomotion often evokes the idea that animals using such devices are perfectly adapted to this kind of motion and should be copied by robotics. The aim of this contribution is to show that the evolution of legs comes late in phylogeny, be it in arthropods or vertebrates. Neural control of legs in vertebrates has to deal with conservative arrangements ‘invented’ for axial locomotion of metameric organisms. The structure of this paper is to show the importance of axial driven propulsion in vertebrates without legs, with legs and only at the end how limbs move the body in eutherian mammals.

The evolution of therian mammals is to a large degree marked by changes in their motion systems. One of the decisive transitions has been from the sprawled, bi‐segmented to the parasagittal, tri‐segmented limb. Here, we review aspects of the tri‐segmented limb in locomotion which have been elucidated in our research groups in the last 10 years. First, we report the kinematics of the tri‐segmented therian limb from mouse to elephant in order to explore general principles of the therian limb configuration and locomotion. Torques will be reported from a previous paper (Witte et al., 2002. J Exp Biol 205:1339–1353) for a better understanding of the anti‐gravity work of all limb joints. The stability of a limb in z‐configuration will be explained and its advantage with respect to other potential solutions from modeling will be discussed. Finally, we describe how the emerging concept of self‐stability can be explained for a tri‐segmented leg template and how it affects the design of the musculoskeletal system and the operation of legs during locomotion. While locomotion has been considered as mainly a control problem in various disciplines, we stress the necessity to reduce control as much as possible. Central control can be cheap if the limbs are “intelligent” by means of their design. Gravity‐induced movements and self‐stability seem to be energy‐saving mechanisms. J. Exp. Zool. 305A, 2006.