Locomotor Pattern Research Articles

Morphine with or without Acepromazine in Horses: A Kinematic Evaluation.

Simple SummaryMorphine is an opioid agonist drug and produces a significant analgesic effect in horses but besides the evidenced analgesic effect, the use of morphine is not routine due to the potential excitatory effects described in the literature. To minimize these effects, neuroleptanalgesia, or the combination of opioids and sedative drugs, is encouraged. Our aim was to describe changes occurring in the locomotor pattern after co-administration of a tranquilizer, acepromazine, and morphine in horses. Six mature horses were used and received four different treatments with saline solution, morphine, acepromazine, or a combination of morphine and acepromazine. A three-dimensional accelerometric device was used to collect data and objectivize those findings moreover the sedative effect of the treatments was also measured. Significant differences were observed when comparing all the treatments in the majority of accelerometric variables, except the regularity of the pattern, some energetic parameters, and tranquilization. An evident counteraction of the effects caused by both morphine and acepromazine was observed. Due to these effects, the possibility of adding acepromazine to an additional analgesic treatment with morphine in the clinical setting ensures the absence of the supplemental instability caused by other sedatives and minimizes the potential opioid excitatory effects.The objective was to demonstrate walking locomotor pattern alterations after co-administration of acepromazine and morphine in horses. Six mature horses receiving four different treatments were used. Treatments consisted of a single dose of saline solution, 0.2 mg/kg bwt of morphine hydrochloride, 0.02 mg/kg bwt of acepromazine maleate, and a combination of 0.2 mg/kg bwt of morphine hydrochloride with 0.02 mg/kg bwt of acepromazine maleate. A three-dimensional accelerometric device was used to collect data. Walking tests were performed 10 min prior to injection, and then at 5, 10, 15, and 20 min after the injection, and then every 10 min for 3 h. Eight variables were calculated including stride kinematic, coordination, and energetic parameters; moreover ground-to-lip distance (GLD), as a tranquilization parameter, was also measured. A significant interaction was observed in all the variables studied but regularity, mediolateral power, the propulsive part of the power, and the GLD. An evident counteraction of the effects caused by both, opioids and phenothiazines, in the gait pattern was observed. The co-administration of acepromazine and morphine could allow a safe opiate administration while minimizing the possible central nervous system (CNS) excitation and reducing potential locomotor adverse effects.

The role of intraspinal sensory neurons in the control of quadrupedal locomotion

From swimming to walking and flying, animals have evolved specific locomotor strategies to thrive in different habitats. All types of locomotion depend on the integration of motor commands and sensory information to generate precisely coordinated movements. Cerebrospinal-fluid-contacting neurons (CSF-cN) constitute a vertebrate sensory system that monitors CSF composition and flow. In fish, CSF-cN modulate swimming activity in response to changes in pH and bending of the spinal cord; however, their role in mammals remains unknown. We used mouse genetics to study their function in quadrupedal locomotion. We found that CSF-cN are directly integrated into spinal motor circuits. The perturbation of CSF-cN function does not affect general motor activity nor the generation of locomotor rhythm and pattern but results in specific defects in skilled movements. These results identify a role for mouse CSF-cN in adaptive motor control and indicate that this sensory system evolved a novel function to accommodate the biomechanical requirements of limb-based locomotion.

Evolutionary Origins of Mammalian Axial Function Revealed through Digital Bending Experiments

The origin of mammals from non‐mammalian synapsids represents an iconic locomotor transition, often described as shift from reptile‐like sprawling limbs and extensive lateral movements of the backbone to upright limbs and sagittal backbone movements that enhance mammal specific locomotor behaviors. However, recent research in comparative anatomy has called into question the lateral‐to‐sagittal functional shift, instead suggesting that vertebral morphology in basal synapsids implies a distinct functional regime not observed in extant sprawling reptiles and represents a unique ancestral condition from which mammals evolved. To investigate this idea further, we used Autobend,a novel, experimentally validated technique for estimating vertebral osteological range of motion (oROM) from skeletons using digital modeling. We applied Autobendto seven extant mammal and reptile species and 11 exceptionally preserved non‐mammalian synapsids to estimate vertebral oROM and intervertebral joint stiffness. Results revealed a clear distinction between extant mammals and reptiles in oROM, with reptiles emphasizing lateral bending and mammals sagittal bending as expected. While most extant taxa exhibited relatively similar levels of stiffness in lateral and sagittal directions, extant lizards and salamanders displayed much more compliance in lateral bending and lower stiffness overall, in keeping with their observed axial kinematics. Conversely, most non‐mammalian synapsids displayed an intermediate condition, with stiffer intervertebral joints than extant lizards, crocodiles, and therian mammals, but similar to the pattern recovered for the modern tuatara. The mammalian condition of sagittal mobility accompanied by axial twisting in the anterior column is first observed in the crownward tritylodontid cynodont Kayentatherium. Together, these results support previous morphology‐based assertions of a distinctive ancestral vertebral function for synapsids characterized by high stiffness without the specialization for lateral bending of extant lizards or the sagittal bending and twisting observed in mammals today. This indicates a more limited role for the axial skeleton in terrestrial locomotion relative to extant groups, and a greater focus on body support than propulsion during the initial stages of synapsid evolution. This stage was followed by the evolution of mammalian‐type mobility patterns in cynodonts, and a secondary reduction in axial stiffness in certain therian clades. Given the strong association between sagittal bending and asymmetric gaits in mammals, these results provide an important first step in reconstructing the evolution of synapsid locomotor patterns.

A Template Model Explains Jerboa Gait Transitions Across a Broad Range of Speeds.

For cursorial animals that maintain high speeds for extended durations of locomotion, transitions between footfall patterns (gaits) predictably occur at distinct speed ranges. How do transitions among gaits occur for non-cursorial animals? Jerboas (Jaculus) are bipedal hopping rodents that frequently transition between gaits throughout their entire speed range. It has been hypothesized that these non-cursorial bipedal gait transitions are likely to enhance their maneuverability and predator evasion ability. However, it is difficult to use the underlying dynamics of these locomotion patterns to predict gait transitions due to the large number of degrees of freedom expressed by the animals. To this end, we used empirical jerboa kinematics and dynamics to develop a unified spring Loaded Inverted Pendulum model with defined passive swing leg motions. To find periodic solutions of this model, we formulated the gait search as a boundary value problem and described an asymmetrical running gait exhibited by the jerboas that emerged from the numerical search. To understand how jerboas change from one gait to another, we employed an optimization approach and used the proposed model to reproduce observed patterns of jerboa gait transitions. We then ran a detailed numerical study of the structure of gait patterns using a continuation approach in which transitions are represented by bifurcations. We found two primary mechanisms to increase the range of speeds at which gait transitions can occur. Coupled changes in the neutral leg swing angle alter leg dynamics. This mechanism generates changes in gait features (e.g., touchdown leg angle and timings of gait events) that have previously been shown to induce gait transitions. This mechanism slightly alters the speeds at which existing gait transitions occur. The model can also uncouple the left and right neutral leg swing angle, which generates asymmetries between left and right leg dynamics. New gait transitions emerge from uncoupled models across a broad range of speeds. In both the experimental observations and in the model, the majority of the gait transitions involve the skipping and asymmetrical running gaits generated by the uncoupled neutral leg swing angle mechanism. This simulated jerboa model is capable of systematically reproducing all biologically relevant gait transitions at a broad range of speeds.

Locomotor Pattern and Force Generation Modulated by Ionic Channels: A Computational Study of Spinal Networks Underlying Locomotion.

Locomotion is a fundamental movement in vertebrates produced by spinal networks known as central pattern generators (CPG). During fictive locomotion cat lumbar motoneurons (MNs) exhibit changes in membrane properties, including hyperpolarization of voltage threshold, reduction of afterhyperpolarization and input resistance, and amplification of nonlinear membrane properties. Both modeling and electrophysiological studies suggest that these changes can be produced by upregulating voltage-gated sodium channel (VGSC), persistent sodium (NaP), or L-type calcium channel (LTCC) or downregulating delayed-rectifier potassium (K(DR)) or calcium-dependent potassium channel (KCa) in spinal MNs. Further studies implicate that these channel modulations increase motor output and facilitate MN recruitment. However, it remains unknown how the channel modulation of CPG networks or MN pools affects the rhythmic generation of locomotion and force production of skeletal muscle during locomotion. In order to investigate this issue, we built a two-level CPG model composed of excitatory interneuron pools (Exc-INs), coupled reciprocally with inhibitory interneuron pools (Inh-INs), and projected to the flexor-extensor MN pools innervating skeletal muscles. Each pool consisted of 100 neurons with membrane properties based on cat spinal neurons. VGSC, K(DR), NaP, KCa, LTCC, and H-current channels were included in the model. Simulation results showed that (1) upregulating VGSC, NaP, or LTCC or downregulating KCa in MNs increased discharge rate and recruitment of MNs, thus facilitating locomotor pattern formation, increased amplitude of electroneurogram (ENG) bursting, and enhanced force generation of skeletal muscles. (2) The same channel modulation in Exc-INs increased the firing frequency of the Exc-INs, facilitated rhythmic generation, and increased flexor-extensor durations of step cycles. (3) Contrarily, downregulation of NaP or LTCC in MNs or Exc-INs or both CPG (Exc-INs and Inh-INs) and MNs disrupted locomotor pattern and reduced or even blocked the ENG bursting of MNs and force generation of skeletal muscles. (4) Pharmacological experiments showed that bath application of 25 μM nimodipine or 2 μM riluzole completely blocked fictive locomotion in isolated rat spinal cord, consistent with simulation results. We concluded that upregulation of VGSC, NaP, or LTCC or downregulation of KCa facilitated rhythmic generation and force production during walking, with NaP and LTCC playing an essential role.

Influence of Step Frequency on the Dynamic Characteristics of Ventilation and Gas Exchange During Sinusoidal Walking in humans

We tested the hypothesis that restricting either step frequency (SF) or stride length (SL) causes a decrease in ventilatory response with limited breath frequency during sinusoidal walking. In this study, 13 healthy male and female volunteers (mean ± SD; age: 21.5 ± 1.8 years, height: 168 ± 7 cm, weight: 61.5 ± 8.3 kg) participated. The walking speed was sinusoidally changed between 50 and 100 m⋅min–1 with periods from 10 to 1 min. Using a customized sound system, we fixed the SF at 120 steps⋅min–1 with SL variation (0.83–0.41 m) (SFfix) or fixed the SL at 0.7 m with SF variation (143–71 steps⋅min–1) (SLfix) during the subjects’ sinusoidal walking. Both the subjects’ preferred locomotion pattern without a sound system (Free) and the unprompted spontaneous locomotor pattern for each subject (Free) served as the control condition. We measured breath-by-breath ventilation [tidal volume (VT) and breathing frequency (Bf)] and gas exchange [CO2 output (CO2), O2 uptake (O2)]. The amplitude (Amp) and the phase shift (PS) of the fundamental component of the ventilatory and gas exchange variables were calculated. The results revealed that the SFfix condition decreased the Amp of the Bf response compared with SLfix and Free conditions. Notably, the Amp of the Bf response under SFfix was reduced by less than one breath at the periods of 5 and 10 min. In contrast, the SLfix condition resulted in larger Amps of Bf and E responses as well as Free. We thus speculate that the steeper slope of the E-CO2 relationship observed under the SLfix might be attributable to the central feed-forward command or upward information from afferent neural activity by sinusoidal locomotive cadence. The PSs of the E, O2, and CO2 responses were unaffected by any locomotion patterns. Such a sinusoidal wave manipulation of locomotion variables may offer new insights into the dynamics of exercise hyperpnea.

Target Constraints Influence Locomotion Pattern to the First Hurdle.

The study examined to what extent the manipulation of hurdle height (0.76-m hurdle, low hurdle 0.50m, and white stripe) would affect visual regulation strategies and kinematic reorganization when approaching the first hurdle. In addition, the impact of constraints as a training tool in terms of creating movement patterns functional for and representative of competitive movement models was assessed. The approach phase to the first hurdle of 13 physical education students with no previous experience in hurdling was video recorded and analyzed. Emergence of different footfall variability curves and movement coordination patterns suggests that participants interact differently with features of the performance context. Contrary to the white stripe, the hurdle height required participants to initiate regulation and distribute adjustments over a larger number of steps, and afforded the preparation for takeoff in order to clear the hurdle. In task design, manipulation of task constraints should offer valuable information regarding the dynamics of movement.

A Markerless Pose Estimator Applicable to Limbless Animals.

The analysis of kinematics, locomotion, and spatial tasks relies on the accurate detection of animal positions and pose. Pose and position can be assessed with video analysis programs, the “trackers.” Most available trackers represent animals as single points in space (no pose information available) or use markers to build a skeletal representation of pose. Markers are either physical objects attached to the body (white balls, stickers, or paint) or they are defined in silico using recognizable body structures (e.g., joints, limbs, color patterns). Physical markers often cannot be used if the animals are small, lack prominent body structures on which the markers can be placed, or live in environments such as aquatic ones that might detach the marker. Here, we introduce a marker-free pose-estimator (LACE Limbless Animal traCkEr) that builds the pose of the animal de novo from its contour. LACE detects the contour of the animal and derives the body mid-line, building a pseudo-skeleton by defining vertices and edges. By applying LACE to analyse the pose of larval Drosophila melanogaster and adult zebrafish, we illustrate that LACE allows to quantify, for example, genetic alterations of peristaltic movements and gender-specific locomotion patterns that are associated with different body shapes. As illustrated by these examples, LACE provides a versatile method for assessing position, pose and movement patterns, even in animals without limbs.

A Fully-Immersive Virtual Reality Setup to Study Gait Modulation.

Objective: Gait adaptation to environmental challenges is fundamental for independent and safe community ambulation. The possibility of precisely studying gait modulation using standardized protocols of gait analysis closely resembling everyday life scenarios is still an unmet need.Methods: We have developed a fully-immersive virtual reality (VR) environment where subjects have to adjust their walking pattern to avoid collision with a virtual agent (VA) crossing their gait trajectory. We collected kinematic data of 12 healthy young subjects walking in real world (RW) and in the VR environment, both with (VR/A+) and without (VR/A-) the VA perturbation. The VR environment closely resembled the RW scenario of the gait laboratory. To ensure standardization of the obstacle presentation the starting time speed and trajectory of the VA were defined using the kinematics of the participant as detected online during each walking trial.Results: We did not observe kinematic differences between walking in RW and VR/A-, suggesting that our VR environment per se might not induce significant changes in the locomotor pattern. When facing the VA all subjects consistently reduced stride length and velocity while increasing stride duration. Trunk inclination and mediolateral trajectory deviation also facilitated avoidance of the obstacle.Conclusions: This proof-of-concept study shows that our VR/A+ paradigm effectively induced a timely gait modulation in a standardized immersive and realistic scenario. This protocol could be a powerful research tool to study gait modulation and its derangements in relation to aging and clinical conditions.

Morphological correlates of distal fibular morphology with locomotion in great apes, humans, and Australopithecus afarensis.

ObjectivesRecent studies highlighted the importance of the fibula to further our understanding of locomotor adaptations in fossil hominins. In this study, we present a three‐dimensional geometric morphometric (3D‐GM) investigation of the distal fibula in extant hominids and Australopithecus afarensis with the aim of pointing out morphological correlations to arboreal behavior.MethodsThree‐dimensional surface meshes of the distal fibula were obtained using computer tomography for 40 extant hominid specimens and laser scanner for five A. afarensis specimens. Distal fibula morphology was quantified positioning 11 fixed landmarks, 40 curve semilandmarks, and 20 surface landmarks on each specimen. A generalized Procrustes analysis (GPA) was carried out on all landmark coordinates followed by Procrustes ANOVA. Principal component analysis (PCA) was performed on the GPA‐aligned shape coordinates. Kruskal‐Wallis tests and Mann–Whitney test were performed on scores along PCs.ResultsGreat apes are characterized by a shorter subcutaneous triangular surface (STS), more downward facing fibulotalar articular facets, more anteriorly facing lateral malleolus and wider/deeper malleolar fossa than humans. Within great apes, orangutans are characterized by more medially facing fibulotalar articular facets. Australopithecus afarensis shows a unique distal fibular morphology with several traits that are generally associated more to arboreality and less to bipedalism such as a short STS, a more anteriorly facing, laterally pointing malleolus and deeper and larger malleolar fossa.ConclusionsThe distal fibula morphology is indicative of locomotor patterns within extant hominids. The 3D‐GM method presented here can be successfully used to further our understanding of arboreal adaptations in fossil hominins.

The evolution of asymmetrical gaits in gnathostome vertebrates.

The difficulty of quantifying asymmetrical limb movements, compared with symmetrical gaits, has resulted in a dearth of information concerning the mechanics and adaptive benefits of these locomotor patterns. Further, no study has explored the evolutionary history of asymmetrical gaits using phylogenetic comparative techniques. Most foundational work suggests that symmetrical gaits are an ancestral feature and asymmetrical gaits are a more derived feature of mammals, some crocodilians, some turtles, anurans and some fish species. In this study, we searched the literature for evidence of the use of asymmetrical gaits across extant gnathostomes, and from this sample (n=308 species) modeled the evolution of asymmetrical gaits assuming four different scenarios. Our analysis shows strongest support for an evolutionary model where asymmetrical gaits are ancestral for gnathostomes during benthic walking and could be both lost and gained during subsequent gnathostome evolution. We were unable to reconstruct the presence/absence of asymmetrical gaits at the tetrapod, amniote, turtle and crocodilian nodes with certainty. The ability to adopt asymmetrical gaits was likely ancestral for Mammalia but was probably not ancestral for Amphibia and Lepidosauria. The absence of asymmetrical gaits in certain lineages may be attributable to neuromuscular and/or anatomical constraints and/or generally slow movement not associated with these gaits. This finding adds to the growing body of work showing the early gnathostomes and tetrapods may have used a diversity of gaits, including asymmetrical patterns of limb cycling.

A serotonergic circuit regulates aversive associative learning under mitochondrial stress in C. elegans

SignificancePhysiological stress triggers avoidance behavior, allowing the animals to stay away from potential threats and optimize their chance of survival. Mitochondrial disruption, a common physiological stress in diverse species, induces the nematode Caenorhabditis elegans to avoid non-pathogenic bacteria through a serotonergic neuronal circuit. We find that distinct neurons, communicated through serotonin and a specific serotonin receptor, are required for the formation and retrieval of this learned aversive behavior. This learned avoidance behavior is associated with increased serotonin synthesis, altered neuronal response property, and reprogramming of locomotion patterns. The circuit and neuromodulatory mechanisms described here offer important insights for stress-induced avoidance behavior.

Performance of Impedance Control-Based Strategies in Power-Assisted Wheelchairs: A Predictive Simulation Study.

Manual wheelchair propulsion is known to be inefficient and causes upper extremity pain, fatigue, and injury. Power-assisted wheelchairs can mitigate these effects through motors that reduce users' effort and load during propulsion. Among the different control strategies proposed to govern the user-wheelchair interaction, impedance control-based ones appear to be the most natural and effective. It can change the apparent dynamical properties of the wheelchair, particularly mass and friction, and automatically compensate for external disturbances such as terrain conditions. This study investigates the advantages and disadvantages of this control strategy employing predictive simulations of locomotion with power-assisted wheelchairs in different scenarios. The simulations are generated using a biomechanically realistic model of the upper extremities and their interaction with the power-assisted wheelchair by solving an optimal control problem. Investigated scenarios include steady-state locomotion vs. a transient maneuver starting from rest, movement on a ramp vs. a level surface, and different choices of reference model parameters. The results reveal that the investigated impedance control-based strategy can effectively reproduce the reference model and reduce the user's effort, with a more significant effect of inertia in the transient maneuver and of friction in steady-state locomotion. However, the simulations also show that imposing a first-order, linear reference model with constant parameters can produce disadvantageous locomotion patterns, particularly in the recovery phase, leading to unnecessary energy dissipation and consequent increase in energy consumption from the batteries. These observations indicate there is room for improvement, for instance, by exploring energy regeneration in the recovery phase or by switching reference model nature or parameters along the cycle. To the best of our knowledge, this is the first investigation of impedance control-based strategies for power-assisted wheelchairs using predictive simulations and a realistic, nonlinear model of the user-wheelchair system.

Does a suspensory lifestyle result in increased tensile strength? Organ-level material properties of sloth limb bones.

The material composition of vertebrate connective tissue is highly conserved across taxa. Existing data suggest that the compressive and tensile strength of limb bones are very similar despite marked variation in limb posture and locomotor patterns. However, the material properties of limb bone tissue from suspensory taxa have not been formally evaluated. Sloths are nearly obligatory in their use of below-branch suspensory locomotion and posture, thus placing their limb bones and associated soft tissue structures under routine tensile loading. It is possible that sloth limb bones are modified for enhanced tensile strength, perhaps at the expense of compressive strength. Forelimb and hindlimb bones of two-toed (Choloepus hoffmanni) and three-toed (Bradypus variegatus) sloths were tested in compression and bending to evaluate this hypothesis. Strength and elastic (Young's) modulus were similarly lower in sloth limb bones during both compression and bending, as compared with pronograde taxa. Ratios of peak bending strength to compressive strength additionally were elevated (sloths: 1.4-1.7; upright taxa: 0.6-1.2) for sloth limb bones. Overall, the material properties measured from the limb bones of tree sloths support our hypothesis of predicted function in a tensile limb system. Future studies should aim to directly test bones in tension to confirm indications of elevated axial tensile strength. Nevertheless, the results herein expand understanding of functional adaptation in mammalian tissue for a range of locomotor/postural behaviors that were previously unexplored.

Kinematic Modeling at the Ant Scale: Propagation of Model Parameter Uncertainties.

Quadrupeds and hexapods are known by their ability to adapt their locomotive patterns to their functions in the environment. Computational modeling of animal movement can help to better understand the emergence of locomotive patterns and their body dynamics. Although considerable progress has been made in this subject in recent years, the strengths and limitations of kinematic simulations at the scale of small moving animals are not well understood. In response to this, this work evaluated the effects of modeling uncertainties on kinematic simulations at small scale. In order to do so, a multibody model of a Messor barbarus ant was developed. The model was built from 3D scans coming from X-ray micro-computed tomography. Joint geometrical parameters were estimated from the articular surfaces of the exoskeleton. Kinematic data of a free walking ant was acquired using high-speed synchronized video cameras. Spatial coordinates of 49 virtual markers were used to run inverse kinematics simulations using the OpenSim software. The sensitivity of the model’s predictions to joint geometrical parameters and marker position uncertainties was evaluated by means of two Monte Carlo simulations. The developed model was four times more sensitive to perturbations on marker position than those of the joint geometrical parameters. These results are of interest for locomotion studies of small quadrupeds, octopods, and other multi-legged animals.

Context-Specificity of Locomotor Learning Is Developed during Childhood.

Humans can perform complex movements with speed and agility in the face of constantly changing task demands. To accomplish this, motor plans are adapted to account for errors in our movements because of changes in our body (e.g., growth or injury) or in the environment (e.g., walking on sand vs ice). It has been suggested that adaptation that occurs in response to changes in the state of our body will generalize across different movement contexts and environments, whereas adaptation that occurs with alterations in the external environment will be context-specific. Here, we asked whether the ability to form generalizable versus context-specific motor memories develops during childhood. We performed a cross-sectional study of context-specific locomotor adaptation in 35 children (3–18 years old) and 7 adults (19–31 years old). Subjects first adapted their gait and learned a new walking pattern on a split-belt treadmill, which has two belts that move each leg at a different speed. Then, subjects walked overground to assess the generalization of the adapted walking pattern across different environments. Our results show that the generalization of treadmill after-effects to overground walking decreases as subjects’ age increases, indicating that age and experience are critical factors regulating the specificity of motor learning. Our results suggest that although basic locomotor patterns are established by two years of age, brain networks required for context-specific locomotor learning are still being developed throughout youth.

Preserved intersegmental coordination during locomotion after cervical spinal cord injury in common marmosets

It is known that primates including human regain some locomotor function after a partial spinal cord injury, but the locomotor pattern is different from before the injury. Although these observations have many implications for improving rehabilitative strategies, these mechanisms are not well understood. In this study, we used a common marmoset hemisection SCI model to examine temporal changes in locomotor pattern, in particular, intersegmental coordination of left hindlimb. Marmoset showed loss of detectable function in the left forelimb and hindlimb after left unilateral hemisection of cervical spinal cord. At two weeks after injury, weight-bearing of the left forelimb during locomotion was limited, but the left hindlimb was able to plantar step. Then marmosets showed gradual recovery in walking ability, but kinematics analysis showed differences in the endpoint trajectory and joint angle movement. Furthermore, intersegmental coordination in left hindlimb represented by planar covariation was preserved over time after the injury. Previous studies have reported that planar covariance is disrupted in patients with stroke or SCI, and that improvement in planarity correlates with recovery in walking ability after rehabilitation. In this study, quadrupedal marmosets were able to walk without loss of balance even after SCI; the different balance needs of bipedal and quadrupedal walkers may lead to differences in planar covariation. Our results show that planar covariation was preserved at all time points after the cervical unilateral hemisection.

State- and Condition-Dependent Modulation of the Hindlimb Locomotor Pattern in Intact and Spinal Cats Across Speeds.

Locomotion after complete spinal cord injury (spinal transection) in animal models is usually evaluated in a hindlimb-only condition with the forelimbs suspended or placed on a stationary platform and compared with quadrupedal locomotion in the intact state. However, because of the quadrupedal nature of movement in these animals, the forelimbs play an important role in modulating the hindlimb pattern. This raises the question: whether changes in the hindlimb pattern after spinal transection are due to the state of the system (intact versus spinal) or because the locomotion is hindlimb-only. We collected kinematic and electromyographic data during locomotion at seven treadmill speeds before and after spinal transection in nine adult cats during quadrupedal and hindlimb-only locomotion in the intact state and hindlimb-only locomotion in the spinal state. We attribute some changes in the hindlimb pattern to the spinal state, such as convergence in stance and swing durations at high speed, improper coordination of ankle and hip joints, a switch in the timing of knee flexor and hip flexor bursts, modulation of burst durations with speed, and incidence of bi-phasic bursts in some muscles. Alternatively, some changes relate to the hindlimb-only nature of the locomotion, such as paw placement relative to the hip at contact, magnitude of knee and ankle yield, burst durations of some muscles and their timing. Overall, we show greater similarity in spatiotemporal and EMG variables between the two hindlimb-only conditions, suggesting that the more appropriate pre-spinal control is hindlimb-only rather than quadrupedal locomotion.

Heterozygous Dcc Mutant Mice Have a Subtle Locomotor Phenotype.

Axon guidance receptors such as deleted in colorectal cancer (DCC) contribute to the normal formation of neural circuits, and their mutations can be associated with neural defects. In humans, heterozygous mutations in DCC have been linked to congenital mirror movements, which are involuntary movements on one side of the body that mirror voluntary movements of the opposite side. In mice, obvious hopping phenotypes have been reported for bi-allelic Dcc mutations, while heterozygous mutants have not been closely examined. We hypothesized that a detailed characterization of Dcc heterozygous mice may reveal impaired corticospinal and spinal functions. Anterograde tracing of the Dcc+/− motor cortex revealed a normally projecting corticospinal tract, intracortical microstimulation (ICMS) evoked normal contralateral motor responses, and behavioral tests showed normal skilled forelimb coordination. Gait analyses also showed a normal locomotor pattern and rhythm in adult Dcc+/− mice during treadmill locomotion, except for a decreased occurrence of out-of-phase walk and an increased duty cycle of the stance phase at slow walking speed. Neonatal isolated Dcc+/− spinal cords had normal left-right and flexor-extensor coupling, along with normal locomotor pattern and rhythm, except for an increase in the flexor-related motoneuronal output. Although Dcc+/− mice do not exhibit any obvious bilateral impairments like those in humans, they exhibit subtle motor deficits during neonatal and adult locomotion.

Comparison between the kinematics for kangaroo rat hopping on a solid versus sand surface.

In their natural habitats, animals move on a variety of substrates, ranging from solid surfaces to those that yield and flow (e.g. sand). These substrates impose different mechanical demands on the musculoskeletal system and may therefore elicit different locomotion patterns. The goal of this study is to compare bipedal hopping by desert kangaroo rats (Dipodomys deserti) on a solid versus granular substrate under speed-controlled conditions. To accomplish this goal, we developed a rotary treadmill, which is able to have different substrates or uneven surfaces. We video recorded six kangaroo rats hopping on a solid surface versus sand at the same speed (1.8 m s−1) and quantified the differences in the hopping kinematics between the two substrates. We found no significant differences in the hop period, hop length or duty cycle, showing that the gross kinematics on the two substrates were similar. This similarity was surprising given that sand is a substrate that absorbs mechanical energy. Measurements of the penetration resistance of the sand showed that the combination of the sand properties, toe-print area and kangaroo rat weight was probably the reason for the similarity.