Emanuel Andrada

To understand how small animals cope with complex, unstructured, and unpredictable substrates, we analyzed the kinetics of rats (n = 10) moving at a fast and at a moderate trot over an unperturbed substrate and a substrate subjected to active horizontal perturbations. Perturbations were active single forwards or backwards displacements of an instrumented platform by amplitudes of 5mm or 10mm in 0.05s. Single leg ground reaction forces (SLGRF) were collected for unperturbed and perturbed locomotion (hindlimbs: 50/102, forelimbs: 45/130, respectively). When negotiating horizontal perturbations, rats displayed gait resetting (braking, accelerating) and non-resetting behaviors. Feedforward strategies differed between the fore- and hindlimbs. In circa 60% of the perturbed trials, forelimbs started the step in acceleration mode, while hindlimbs began the stance mostly in non-resetting mode (∼45%). In about 50% of all perturbed steps, the impulse provided by the SLGRF displayed a change in behavior according to the expected response to the perturbation. The remaining 50% retained the feedforward strategy. Still, most perturbed trials displayed changes in SLGRF patterns that indicated passive and active reactions to platform shifts. Our results indicate that rats’ sensorimotor control system tunes fore- and hindlimbs differently in expectation of a perturbation. In addition, the tendon-muscle systems of the limbs are recruited to prevent leg collapse at the beginning and end of the stance. At lower speeds, spinal and/or higher center commands have enough time to re-adapt limb behavior. At higher speeds of locomotion, rats rely more on their limbs’ intrinsic stability and on feedforward control.

Macaques trained to perform bipedally used running gaits across a wide range of speeds. At higher speeds they preferred unilateral skipping (galloping). The same asymmetric stepping pattern was used while hurdling across two low obstacles placed at the distance of a stride within our experimental track. In bipedal macaques during skipping, we expected a differential use of the trailing and leading legs. The present study investigated global properties of the effective and virtual leg, the location of the virtual pivot point (VPP), and the energetics of the center of mass (CoM), with the aim of clarifying the differential leg operation during skipping in bipedal macaques. When skipping, macaques displayed minor double support and aerial phases during one stride. Asymmetric leg use was indicated by differences in leg kinematics. Axial damping and tangential leg work did not influence the indifferent peak ground reaction forces and impulses, but resulted in a lift of the CoM during contact of the leading leg. The aerial phase was largely due to the use of the double support. Hurdling amplified the differential leg operation. Here, higher ground reaction forces combined with increased double support provided the vertical impulse to overcome the hurdles. Following CoM dynamics during a stride, skipping and hurdling represented bouncing gaits. The elevation of the VPP of bipedal macaques resembled that of human walking and running in the trailing and leading phases, respectively. Because of anatomical restrictions, macaque unilateral skipping differs from that of humans, and may represent an intermediate gait between grounded and aerial running.

When locomoting bipedally at higher speeds, macaques preferred unilateral skipping (galloping). The same skipping pattern was maintained while hurdling across two low obstacles at the distance of a stride within our experimental track. The present study investigated leg and trunk joint rotations and leg joint moments, with the aim of clarifying the differential leg and trunk operation during skipping in bipedal macaques. Especially at the hip, the range of joint rotation and extension at lift off was larger in the leading than in the trailing leg. The flexing knee absorbed energy and the extending ankle generated work during each step. The trunk showed only minor deviations from symmetry. Hurdling amplified the differences and notably resulted in a quasi‐elastic use of the leading knee and in an asymmetric operation of the trunk.

Macaques trained to perform bipedally used running gaits across a wide range of speed. At higher speeds they preferred unilateral skipping (galloping). The same asymmetric stepping pattern was used while hurdling across two low obstacles placed at the distance of a stride within our experimental track. In bipedal macaques during skipping, we expected a differential use of the trailing and leading legs. The present study investigated global properties of the effective and virtual leg, the location of the virtual pivot point (VPP), and the energetics of the center of mass (CoM), with the aim of clarifying the differential leg operation during skipping in bipedal macaques. Macaques skipping displayed minor double support and aerial phases during one stride. Asymmetric leg use indicated by differences in leg kinematics. Axial damping and tangential leg work did not influence the indifferent peak ground reaction forces and impulses, but resulted in a lift of the CoM during contact of the leading leg. The aerial phase was largely due to the use of the double support. Hurdling amplified the differences. Here, higher ground reaction forces combined with increased double support provided the vertical impulse to overcome the hurdles. Following CoM dynamics during a stride skipping and hurdling represented bouncing gaits. The elevation of the VPP of bipedal macaques resembled that of human walking and running in the trailing and leading phases, respectively. Due to anatomical restrictions, macaque unilateral skipping differs from that of humans, and may represent an intermediate gait between grounded and aerial running.

Dogs (Canis familiaris) prefer the walk at lower speeds and the more economical trot at speeds ranging from 0.5 Fr up to 3 Fr. Important works have helped to understand these gaits at the levels of the center of mass, joint mechanics, and muscular control. However, less is known about the global dynamics for limbs and if these are gait or breed-specific. For walk and trot, we analyzed dogs’ global dynamics, based on motion capture and single leg kinetic data, recorded from treadmill locomotion of French Bulldog (N= 4), Whippet (N= 5), Malinois (N= 4), and Beagle (N= 5). Dogs’ pelvic and thoracic axial leg functions combined compliance with leg lengthening. Thoracic limbs were stiffer than the pelvic limbs and absorbed energy in the scapulothoracic joint. Dogs’ ground reaction forces (GRF) formed two virtual pivot points (VPP) during walk and trot each. One emerged for the thoracic (fore) limbs (VPP_TL) and is roughly located above and caudally to the scapulothoracic joint. The second is located roughly above and cranially to the hip joint (VPP_PL). The positions of VPPs and the patterns of the limbs’ axial and tangential projections of the GRF were gaits but not always breeds-related. When they existed, breed-related changes were mainly exposed by the French Bulldog. During trot, positions of the VPPs tended to be closer to the hip joint or the scapulothoracic joint, and variability between and within breeds lessened compared to walk. In some dogs, VPP_PLwas located below the pelvis during trot. Further analyses revealed that leg length and not breed may better explain differences in the vertical position of VPP_TLor the horizontal position of VPP_PL. The vertical position of VPP_PLwas only influenced by gait, while the horizontal position of VPP_TLwas not breed or gait-related. Accordingly, torque profiles in the scapulothoracic joint were likely between breeds while hip torque profiles were size-related. In dogs, gait and leg length are likely the main VPPs positions’ predictors. Thus, variations of VPP positions may follow a reduction of limb work. Stability issues need to be addressed in further studies.

Dogs ( Canis familiaris ) prefer the walk at lower speeds and the more economical trot at speeds ranging from 0.5 Fr up to 3 Fr. Important works were carried out to understand these gaits at the levels of center of mass, joint mechanics, and muscular control. However, less is known about the global control goals for limbs and overall locomotion, and of whether these global control goals are gait or breed specific. For walk and trot, we analyzed dog global dynamics based on motion capture and single leg kinetic data recorded from treadmill locomotion of French Bulldog (N = 4), Whippet (N = 5), Malinois (N = 4) and Beagle (N = 5). Dogs displayed two virtual pivot points (VPP) during walk and trot each. One resembles control of both thoracic (fore) limbs and is roughly located above and caudally to the scapular pivot, while the second is located roughly above and cranially to the hip and mirrors the control of the pelvic (hind-) limbs. The positions of VPPs and the patterns of the legs‘ axial and tangential functions were gait and breed related. However, breed related changes were mainly exposed by the French Bulldog. The position of VPPs relative to the proximal pivots explains the propulsive and breaking forces observed in quadrupedal locomotion and may help to reduce limb work. In combination with former work, from the present study the VPP template emerges as the expression of a simple and general global control rule for both bipeds and quadrupeds.

Small cursorial birds display remarkable walking skills and can negotiate complex and unstructured terrains with ease. The neuromechanical control strategies necessary to adapt to these challenging terrains are still not well understood. Here, we analyzed the 2D- and 3D pelvic and leg kinematic strategies employed by the common quail to negotiate visible steps (upwards and downwards) of about 10%, and 50% of their leg length. We used biplanar fluoroscopy to accurately describe joint positions in three dimensions and performed semi-automatic landmark localization using deep learning. Quails negotiated the vertical obstacles without major problems and rapidly regained steady-state locomotion. When coping with step upwards, the quail mostly adapted the trailing limb to permit the leading leg to step on the elevated substrate similarly as it did during level locomotion. When negotiated steps downwards, both legs showed significant adaptations. For those small and moderate step heights that did not induce aerial running, the quail kept the kinematic pattern of the distal joints largely unchanged during uneven locomotion, and most changes occurred in proximal joints. The hip regulated leg length, while the distal joints maintained the spring-damped limb patterns. However, to negotiate the largest visible steps, more dramatic kinematic alterations were observed. There all joints contributed to leg lengthening/shortening in the trailing leg, and both the trailing and leading legs stepped more vertically and less abducted. In addition, locomotion speed was decreased. We hypothesize a shift from a dynamic walking program to more goal-directed motions that might be focused on maximizing safety.