Elastic extension of leg tendons in the locomotion of horses (Equus caballus)

Several of the distal leg muscles of horses have such extremely short muscle fibres that their changes of length in locomotion must be due almost entirely to elastic extension of their tendons. Films of a horse have been analysed to determine these extensions, using data obtained by experiments on dissected legs. The tendons investigated experience peak strains of 3–6% in walking, 3–7% in trotting and 4–9% in galloping. These strains occur while the foot is on the ground.     

Why Mammals Gallop

SYNOPSIS: Most mammals use symmetrical gaits (such as the trot)at moderate speeds but change to asymmetrical gaits (gallops)at high speeds. A mathematical model of quadrupedal gaits failedto show any advantage in this change: it seemed to show that,even at high speeds, there was always a symmetrical gait thatwas at least as economical as galloping. That model treatedthe back as rigid, but another model seemed to show that backmovements such as occur in galloping could only increase theenergy cost. However, metabolic measurements on horses showedthat galloping is more economical than trotting at high speeds.The explanation seems to be that kinetic energy fluctuations,due to backward and forward swinging of the legs, become verylarge at high speeds. Galloping makes it possible for kineticenergy associated with leg movements to be stored briefly asstrain energy in elastic structures in the back, and returnedin an elastic recoil. The most important of the strain energystores in the back, that have been discovered so far, is theaponeurosis of the longissimus muscle.     

Fast locomotion in caterpillars

The maximum forward crawling speeds of caterpillars are limited by the hydraulic design of the body and the peristaltic mode of operation of the segmental muscles. High speed locomotory manoeuvers can be achieved by reversing the direction of the normal peristaltic wave (from posterior-anterior to anterior-posterior) although the penalty is a dramatically reduced duty factor of the legs and potential instability. This study describes the suite of reverse gaits available to caterpillars, from reverse walking (the kinematic inverse of normal forward walking), through to reverse galloping (in which all the legs save the claspers are wrenched free of the ground with each step) to recoil-and-roll, a unique form of locomotion in which the insect free-wheels backwards at high-speed. These reverse forms of locomotion are produced primarily in response to threat, involve bilateral activation of the intersegmental muscles and are relatively simple in terms of neural control. The ecological roles of high-speed locomotion are considered in the light of potential predators and the normal habitat and terrain.     

Mutual influences of upper and lower extremities during cyclic movements

The possibility for the activation of muscles in a passive arm during its cyclic movements imposed by active movements of the contralateral arm or by an experimenter and the effect that the movements of lower extremities have on the activity of the arm muscles have been studied. In addition, the activity of the leg muscles was studied as dependent on the motor task performed by the arms. Ten healthy subjects performed antiphase arm movements with and without stepping-like movements of both legs in the supine position. The experiment was performed under three conditions for the arm movements: (1) both arms performed active movements; (2) one arm performed active movements, and the contralateral arm, being entirely passive, was forced to participate in movements; (3) the movement of the passive arm was caused by an experimenter. Under condition (2), additional loadings of 30 and 60 N were applied to the active arm. Under all conditions, the arm movements were performed with and without leg movements. The possibility for the activation of muscles in the arm performing passive movements has been demonstrated. To a large extent, this is possible due to an increase in the afferent inflow from the muscles of the contralateral arm. The electrical activity was modulated during cyclic arm movements and depended on the level of loading of the active arm. During the combined active movements of the arms and legs, the reduction in the activity of the flexor muscles of the shoulder and forearm was observed. In the case of passive stepping-like movements, the concomitant arm movements increased the magnitude of electromyographic bursts in most of the examined leg muscles. During active leg movements, a similar increase in electromyographic bursts was observed only in the m. biceps femoris (BF) and the anterior tibial muscle. An increase in the loading of one arm caused a significant increase in the EMG activity in most examined muscles of the legs. The data obtained provide additional proof for the existence of a functionally significant neuronal interaction between the arms, as well as between the upper and lower extremities, which is probably due to intraspinal neuronal connections.     

Comparative analysis of the intrinsic leg musculature of the American cockroach,Periplaneta americana (L.)

The American cockroach has a total of 368 muscles inserting on the post-coxal segments of its legs. By using a narrow morphological definition for delimiting individual muscles, it is shown (i) that the protrochanteral musculatures (23 muscles/leg) differ from the essentially identical meso- and metatrochanteral musculatures (24 and 26 muscles/leg) in number and disposition of extensors and in having a completely different flexor composition, and (ii) that the musculatures of the more distal segments of the legs are completely serially homologous, there being 2 muscles for moving each femur, 23 for each tibia, 7 for each first tarsomere, and 5 for each of the paired pretarsal claws. In all six legs, the trochanteral and tibial musculatures each contain single slender muscles that may be acting proprioceptively to measure the angular displacements between, respectively, the coxas and trochanters, and the femurs and tibias. Neurological and phylogenetic considerations are used to demonstrate why a narrow morphological definition should be employed, and why the widely used functional definition of Snodgrass ('35) is not only fallacious on evolutionary grounds, but also leads to making erroneous conclusions regarding the manner in which insect musculature is controlled by the insect central nervous system. Finally, it is hypothesized that the physiological limitations imposed by having an open circulatory system and the problems inherent in the neural control of large muscles may have been major evolutionary factors in forcing insects to use many slender muscles to control their body movements.     

Three-dimensional knee joint contact forces during walking in unilateral transtibial amputees

Individuals with unilateral transtibial amputations have greater prevalence of osteoarthritis in the intact knee joint relative to the residual leg and non-amputees, but the cause of this greater prevalence is unclear. The purpose of this study was to compare knee joint contact forces and the muscles contributing to these forces between amputees and non-amputees during walking using forward dynamics simulations. We predicted that the intact knee contact forces would be higher than those of the residual leg and non-amputees. In the axial and mediolateral directions, the intact and non-amputee legs had greater peak tibio-femoral contact forces and impulses relative to the residual leg. The peak axial contact force was greater in the intact leg relative to the non-amputee leg, but the stance phase impulse was greater in the non-amputee leg. The vasti and hamstrings muscles in early stance and gastrocnemius in late stance were the largest contributors to the joint contact forces in the non-amputee and intact legs. Through dynamic coupling, the soleus and gluteus medius also had large contributions, even though they do not span the knee joint. In the residual leg, the prosthesis had large contributions to the joint forces, similar to the soleus in the intact and non-amputee legs. These results identify the muscles that contribute to knee joint contact forces during transtibial amputee walking and suggest that the peak knee contact forces may be more important than the knee contact impulses in explaining the high prevalence of intact leg osteoarthritis.     

The influence of increasing steady-state walking speed on muscle activity in below-knee amputees

The goal of this study was to identify changes in muscle activity in below-knee amputees in response to increasing steady-state walking speeds. Bilateral electromyographic (EMG) data were collected from 14 amputee and 10 non-amputee subjects during four overground walking speeds from eight intact leg and five residual leg muscles. Using integrated EMG measures, we tested three hypotheses for each muscle: (1) there would be no difference in muscle activity between the residual and intact legs, (2) there would be no difference in muscle activity between the intact leg and non-amputee legs, and (3) muscle activity in the residual and intact legs would increase with speed. Most amputee EMG patterns were similar between legs and increased in magnitude with speed. Differences occurred in the residual leg biceps femoris long head, vastus lateralis and rectus femoris, which increased in magnitude during braking compared to the intact leg. These adaptations were consistent with the need for additional body support and forward propulsion in the absence of the plantar flexors. With the exception of the intact leg gluteus medius, all intact leg muscles exhibited similar EMG patterns compared to the control leg. Finally, the residual, intact and control leg EMG all had a significant speed effect that increased with speed with the exception of the gluteus medius.     

The legs of ostriches (Struthio) and moas (Pachyornis)

Ostriches were filmed running at maximum speed, and forces on the feet were calculated. Measurements were made of the principal structures in the legs of an ostrich. Hence peak stresses in muscles, tendons and bones were calculated. They lay within the range of stresses calculated for strenuous activities of other vertebrates. The ostrich makes substantial savings of energy in running, by elastic storage in stretched tendons. Pachyornis was a flightless bird, much heavier than ostriches and with massively thick leg bones. These bones are shorter than predicted for its estimated body mass, by extrapolation from allometric equations for flying birds. An attempt is made to calculate the stresses that acted in the leg bones in running, for all possible patterns of leg movement. The stresses were probably rather low, unless Pachyornis was capable of running fast. It is argued that the optimum factor of safety for moo leg bones may have been exceptionally high, as a consequence of the absence of predators.     

The role of tendon elasticity in the locomotion of the camel (Camelus dromedarius)

Several ofthe distal leg muscles ofcamels have very short or even rudimentary muscle fibres. This makes it possible to calculate the elastic extensions of tendons that occur in running, from the leg positions observed in films. A series of experiments have been performed for this purpose on the dissected legs of a camel. The initial conclusions derived from them are modified in the light of estimates of the forces that act on the tendons, and of measurements of the elastic properties of one tendon. Evidence for movement at the intertarsal and tarso-metatarsal joints. and the corresponding joints of the fore leg, is examined. The importance of the various tendons as elastic energy stores in running is assessed.     

Remodelling of Neuromuscular Systems During Insect Metamorphosis

During metamorphosis in the hawkmoth, Manduca sexta, the larvalthoracic legs are replaced by a new set of adult legs that includenew sensory neurons and muscles, and participate in new patternsof locomotor activity. Larval leg motoneurons persist to innervatethe new adult leg muscles, but undergo striking changes in dendriticmorphology that are regulated by the insect steroid, 20-hydroxyecdysone.In the periphery, the motor terminals regress as larval musclesdegenerate, and expand as new adult muscles form from myoblasts.Evidence obtained both in vivo and in vitro suggests that theproliferation of myoblasts during metamorphosis is dependentupon innervation.     

Influence of Achilles tendon vibration on the vertical posture during standing with asymmetrical leg loading

The shift of the common center of pressure (CCP) and the center of pressure (CP) of one leg was studied during the Achilles tendon vibration of one or both legs while the subject was standing with symmetrical load on the legs or with the load transferred to one leg. The CP shift of the standing subject during unilateral Achilles tendon vibration depended on both the side of application of vibration and on the distribution of the leg load. During standing with a asymmetrical load on the legs, the shift of the CCP was larger than when the vibration was applied to the loaded leg. The CP shift of one leg was greater if both vibration and the load were applied to it. Vibration of the unloaded leg caused a CP shift in the loaded contralateral leg. In this case, vibration of the left unloaded leg did not cause any noticeable CP shift of the left leg, while vibration of the unloaded right leg caused a CP shift of the right leg. Under the similar conditions of loading and vibration, the displacement of the CP of the right leg was larger than the displacement of the CP of the left leg. It may be suggested that postural asymmetry and unilateral vibration of the leg muscles change the internal representation of the position of the body axis in relation to the vertical, which affects the displacement of the CP of one leg in response to afferent stimulation of the leg muscles.     

Cupiennius salei: biomechanical properties of the tibia–metatarsus joint and its flexing muscles

Hunting spiders are well adapted to fast locomotion. Space saving hydraulic leg extension enables leg segments, which consist almost soley of flexor muscles. As a result, the muscle cross sectional area is high despite slender legs. Considering these morphological features in context with the spider’s segmented C-shaped legs, these specifics might influence the spider’s muscle properties. Moreover, these properties have to be known for modeling of spider locomotion. Cupiennius salei (n = 5) were fixed in a metal frame allowing exclusive flexion of the tibia–metatarsus joint of the second leg (counted from anterior). Its flexing muscles were stimulated supramaximally using needle electrodes. Accounting for the joint geometry, the force–length and the force–velocity relationships were determined. The spider muscles produce 0.07 N cm maximum isometric moment (corresponding to 25 N/cm² maximum stress) at 160° tibia–metatarsus joint angle. When overextended to the dorsal limit at approximately 200°, the maximum isometric moments decrease to 72%, and, when flexed to the ventral hinge stop at 85°, they drop to 11%. The force–velocity relation shows the typical hyperbolic shape. The mean maximum shortening velocity is 5.7 optimum muscle lengths per second and the mean curvature (a/F _iso) of the Hill-function is 0.34. The spider muscle’s properties which were determined are similar to those of other species acting as motors during locomotion (working range, curvature of Hill hyperbola, peak power at the preferred speeds), but they are relatively slow. In conjunction with the low mechanical advantage (muscle lever/load arm), the arrangement of three considerably actuated joints in series may nonetheless enable high locomotion velocities.     

Mechanics of running of the ostrich (Struthio camelus)

Ostriches have been filmed running fast in their natural habitat. A female ostrich has been dissected and the principal bones, muscles and tendons in a leg have been measured. It is calculated that stresses up to 240 kN m⁻² and 40 MN m⁻², respectively, act in the digital flexor muscles and their tendons during running. Tensile and compressive stresses up to about 70MNm⁻² and 110 MNm⁻² act in the tibiotarsus. A large proportion of the energy which would otherwise be required for running is probably saved by elastic storage in tendons. Comparisons are made with the legs of flying birds and of antelopes.     

Dynamic simulation of insect walking

Insect walking relies on a complex interaction between the environment, body segments, muscles and the nervous system. For the stick insect in particular, previous investigations have highlighted the role of specific sensory signals in the timing of activity of central neural networks driving the individual leg joints. The objective of the current study was to relate specific sensory and neuronal mechanisms, known from experiments on reduced preparations, to the generation of the natural sequence of events forming the step cycle in a single leg. We have done this by simulating a dynamic 3D-biomechanical model of the stick insect coupled to a reduced model of the neural control system, incorporating only the mechanisms under study. The neural system sends muscle activation levels to the biomechanical system, which in turn provides correctly timed propriosensory signals back to the neural model. The first simulations were designed to test if the currently known mechanisms would be sufficient to explain the coordinated activation of the different leg muscles in the middle leg. Two experimental situations were mimicked: restricted stepping where only the coxa-trochanteral joint and the femur-tibia joint were free to move, and the unrestricted single leg movements on a friction-free surface. The first of these experimental situations is in fact similar to the preparation used in gathering much of the detailed knowledge on sensory and neuronal mechanisms. The simulations show that the mechanisms included can indeed account for the entire step cycle in both situations. The second aim was to test to what extent the same sensory and neuronal mechanisms would be adequate also for controlling the front and hind legs, despite the large differences in both leg morphology and kinematic patterns. The simulations show that front leg stepping can be generated by basically the same mechanisms while the hind leg control requires some reorganization. The simulations suggest that the influence from the femoral chordotonal organs on the network controlling levation-depression may have a reversed effect in the hind legs as compared to the middle and front legs. This, and other predictions from the model will have to be confirmed by additional experiments.     

Locomotion and bone strength of the white rhinoceros, Ceratotherium simum

Measurements have been made, of lengths and of geometric properties of cross-sections, of the long bones of the legs of a young white rhinoceros of about 750 kg body mass. These are considered in conjunction with data from film of white rhinoceros trotting and galloping. The stresses developed in the bones in running are rather low, in comparison with other large mammals, suggesting that rhinoceros skeletons may be built to unusually high factors of safety. The long, relatively straight legs of elephants (whose bones experience higher stresses) are contrasted with the shorter, less straight legs of the other graviportal mammals.     

BigDog-inspired studies in the locomotion of goats and dogs

Collision-based expenditure of mechanical energy and the compliance and geometry of the leg are fundamental, interrelated considerations in the mechanical design of legged runners. This article provides a basic context and rationale for experiments designed to inform each of these key areas in Boston Dynamic's BigDog robot. Although these principles have been investigated throughout the past few decades within different academic disciplines, BigDog required that they be considered together and in concert with an impressive set of control algorithms that are not discussed here. Although collision reduction is an important strategy for reducing mechanical cost of transport in the slowest and fastest quadrupedal gaits, walking and galloping, BigDog employed an intermediate-speed trotting gait without collision reduction. Trotting, instead, uses a spring-loaded inverted pendulum mechanism with potential for storage and return of elastic strain energy in appropriately compliant structures. Rather than tuning BigDog's built-in leg springs according to a spring-mass model-based virtual leg-spring constant , a much stiffer distal leg spring together with actuation of the adjacent joint provided good trotting dynamics and avoided functional limitations that might have been imposed by too much compliance in real-world terrain. Adjusting the directional compliance of the legs by adopting a knee-forward, elbow-back geometry led to more robust trotting dynamics by reducing perturbations about the pitch axis of the robot's center of mass (CoM). BigDog is the most successful large-scale, all-terrain trotting machine built to date and it continues to stimulate our understanding of legged locomotion in comparative biomechanics as well as in robotics.     

Coding proprioceptive information to control movement to a target: Simulation with a simple neural network

When the stick insect walks, the middle and rear legs step to positions immediately behind the tarsus of the adjacent rostral leg. Previous reports have described this movement to a target as a relationship between the tarsus positions of the two legs in a Cartesian coordinate system. However, leg proprioceptors measure the position of the target leg in terms of joint angles and leg muscles bring the tarsus of the moving leg to the proper end-point by establishing appropriate angles at the joints. Representation of this task in Cartesian coordinates requires non-linear coordinate transformations; realizing such a transformation in the nervous system appears to require many neurons. The present simulation using the back-propagation algorithm shows that a simple network of only nine units — 3 sensory input units, 3 motor output units, and 3 hidden units — suffices. The simulation also shows that an analytic coordinate transformation can be replaced by a direct association of joint configurations in the moving leg with those in the target leg.     

Bed rest attenuates sympathetic and pressor responses to isometric exercise in antigravity leg muscles in humans

Although spaceflight and bed rest are known to cause muscular atrophy in the antigravity muscles of the legs, the changes in sympathetic and cardiovascular responses to exercises using the atrophied muscles remain unknown. We hypothesized that bed rest would augment sympathetic responses to isometric exercise using antigravity leg muscles in humans. Ten healthy male volunteers were subjected to 14-day 6 degrees head-down bed rest. Before and after bed rest, they performed isometric exercises using leg (plantar flexion) and forearm (handgrip) muscles, followed by 2-min postexercise muscle ischemia (PEMI) that continues to stimulate the muscle metaboreflex. These exercises were sustained to fatigue. We measured muscle sympathetic nerve activity (MSNA) in the contralateral resting leg by microneurography. In both pre- and post-bed-rest exercise tests, exercise intensities were set at 30 and 70% of the maximum voluntary force measured before bed rest. Bed rest attenuated the increase in MSNA in response to fatiguing plantar flexion by approximately 70% at both exercise intensities (both P < 0.05 vs. before bed rest) and reduced the maximal voluntary force of plantar flexion by 15%. In contrast, bed rest did not alter the increase in MSNA response to fatiguing handgrip and had no effects on the maximal voluntary force of handgrip. Although PEMI sustained MSNA activation before bed rest in all trials, bed rest entirely eliminated the PEMI-induced increase in MSNA in leg exercises but partially attenuated it in forearm exercises. These results do not support our hypothesis but indicate that bed rest causes a reduction in isometric exercise-induced sympathetic activation in (probably atrophied) antigravity leg muscles.     

The tracheae of water mites

The water mites of standing waters have evolved a novel respiratory system consisting of numerous independent tracheae of tracheolar dimensions. Each trachea has a portion of its length lying directly under the cuticle and one or both ends of the trachea turn into the body to supply some organ. There is no fusion of tracheae to form trunks. Areas of dense tracheation dorsal to the legs supply the leg muscles, and sometimes there is a distinct area of the venter that supplies the muscles of the mouthparts.     

Leg kinematics and muscle activity during treadmill running in the cockroach, Blaberus discoidalis : I. Slow running

We have combined high-speed video motion analysis of leg movements with electromyogram (EMG) recordings from leg muscles in cockroaches running on a treadmill. The mesothoracic (T2) and metathoracic (T3) legs have different kinematics. While in each leg the coxa-femur (CF) joint moves in unison with the femur-tibia (FT) joint, the relative joint excursions differ between T2 and T3 legs. In T3 legs, the two joints move through approximately the same excursion. In T2 legs, the FT joint moves through a narrower range of angles than the CF joint. In spite of these differences in motion, no differences between the T2 and T3 legs were seen in timing or qualitative patterns of depressor coxa and extensor tibia activity. The average firing frequencies of slow depressor coxa (Ds) and slow extensor tibia (SETi) motor neurons are directly proportional to the average angular velocity of their joints during stance. The average Ds and SETi firing frequency appears to be modulated on a cycle-by-cycle basis to control running speed and orientation. In contrast, while the frequency variations within Ds and SETi bursts were consistent across cycles, the variations within each burst did not parallel variations in the velocity of the relevant joints. Accepted: 24 May 1997