Morphology and mechanics of myosepta in a swimming salamander (Siren lacertina)

In contrast to the complex, three-dimensional shape of myomeres in teleost fishes, the lateral hypaxial muscles of salamanders are nearly planar and their myosepta run in a roughly straight line from mid-lateral to mid-ventral. We used this relatively simple system as the basis for a mathematical model of segmented musculature. Model results highlight the importance of the mechanics of myosepta in determining the shortening characteristics of a muscle segment. We used sonomicrometry to measure the longitudinal deformation of myomeres and the dorsoventral deformation of myosepta in a swimming salamander (Siren lacertina). Sonomicrometry results show that the myosepta allow some dorsoventral lengthening, indicating an amplification of myomere shortening that is greater than that produced by muscle fiber angle alone (10% muscle fiber shortening produces 28.7% myomere shortening). Polarized light and DIC microscopy of isolated hypaxial myosepta revealed that the collagen fiber orientation in hypaxial myomeres is primarily mediolateral. The mediolateral collagen fiber orientation, combined with our finding that the hypaxial myosepta lengthen dorsoventrally during swimming, suggests that one possible function of hypaxial myosepta in S. lacertina is to increase the strain amplification of the muscle fibers by reducing the mediolateral bulging of the myomeres and redirecting the bulging toward the dorsoventral direction.     

Patterns of hind limb motor output during walking in the salamander Dicamptodon tenebrosus,with comparisons to other tetrapods

Based on similarity of motor patterns of lizards, crocodiles, birds and mammals, various authors have concluded that a number of homologous muscles across these taxa demonstrate neuromuscular conservatism. This hypothesis remains untested for more basal taxa. Therefore, a quantitative electromyographic study of the hind limb during treadmill walking (mean speed of 0.75 SVL/s) in the salamander Dicamptodon tenebrosus was undertaken. Muscles located ventrally on the hind limb become active just before foot placement on the substrate, and maintain activity through the first half of the stance phase. Dorsally located muscles begin activity at or just before the start of the swing phase, and fire through the first half of swing. Several muscles showed a secondary EMG burst during the stride. The second burst in most ventral muscles occurred in late stance. In all dorsal muscles with double bursts, the second burst occurred in the middle of stance. Comparison of electromyographic onset and offset values for Dicamptodon to those for presumed homologues in other tetrapods reveals similarity in activity patterns for all ventral and two dorsal muscles despite anatomical rearrangements, supporting the hypothesis of neuromuscular conservatism for some muscles but not others.Abbreviations BF biceps femoris muscle - CDF caudofemoralis muscle - CPIT caudalipuboischiotibialis muscle - Dist distal - EDC extensor digitorum communis muscle - EMG electromyogram - EXF extensor cruris et tarsi fibularis muscle - EXT extensor cruris tibialis muscle - FMFB femorofibularis muscle - FPC flexor primordialis communis muscle - Gastroc gastrocnemius muscle - ILFB iliofibularis muscle - ILFM iliofemoralis muscle - ILTA extensor iliotibialis pars anterior muscle - ILTP extensor iliotibialis pars posterior muscle - ISC ischiocaudalis muscle - ISF ischioflexorius muscle - ISFM ischiofemoralis muscle - ITCR iliotrochantericus cranialis muscle - ITM iliotrochantericus medius muscle - MG medial gastrocnemius muscle - PFM pubifemoralis muscle - PIFE puboischiofemoralis externus muscle - PIFI puboischiofemoralis internus muscle - PIT puboischiotibialis muscle - Prox proximal - PTB pubotibialis muscle - Sol soleus muscle - ST semitendinosus muscle - SVL snout-vent length     

Pendular activity of human upper limbs during slow and normal walking

When walking at normal and fast speeds, humans swing their upper limbs in alternation, each upper limb swinging in phase with the contralateral lower limb. However, at slow and very slow speeds, the upper limbs swing forward and back in unison, at twice the stride frequency of the lower limbs. The change from “single swinging” (in alternation) to “double swinging” (in unison) occurs consistently at a certain stride frequency for agiven individual, though different individuals may change at different stride frequencies. To explain this change in the way we use our upper limbs and individual variations in the occurrence of the change, the upper limb is modelled as a compound pendulum. Based on the kinematic properties of pendulums, we hypothesize that the stride frequency at which the change from “single swinging” to “double swinging” occurs will be at or slightly below the natural pendular frequency (NPF) of the upper limbs. Twenty-seven subjects were measured and then filmed while walking at various speeds. The mathematically derived NPF of each subject's upper limbs was compared to the stride frequency at which the subject changed from “single swinging” to “double swinging.” The results of the study conform very closely to the hypothesis, even when the NPF is artificially altered by adding weights to the subjects' hands. These results indicate that the pendulum model of the upper limb will be useful in further investigations of the function of the upper limbs in human walking. © 1994 Wiley-Liss, Inc.     

Variability of neural activation during walking in humans: short heels and big calves

People come in different shapes and sizes. In particular, calf muscle size in humans varies considerably. One possible cause for the different shapes of calf muscles is the inherent difference in neural signals sent to these muscles during walking. In sedentary adults, the variability in neural control of the calf muscles was examined with muscle size, walking kinematics and limb morphometrics. Half the subjects walked while activating their medial gastrocnemius (MG) muscles more strongly than their lateral gastrocnemius (LG) muscles during most walking speeds ('MG-biased'). The other subjects walked while activating their MG and LG muscles nearly equally ('unbiased'). Those who walked with an MG-biased recruitment pattern also had thicker MG muscles and shorter heel lengths, or MG muscle moment arms, than unbiased walkers, but were similar in height, weight, lower limb length, foot length, and exhibited similar walking kinematics. The relatively less plastic skeletal system may drive calf muscle size and motor recruitment patterns of walking in humans.     

Humans exploit the biomechanics of bipedal gait during visually guided walking over complex terrain

How do humans achieve such remarkable energetic efficiency when walking over complex terrain such as a rocky trail? Recent research in biomechanics suggests that the efficiency of human walking over flat, obstacle-free terrain derives from the ability to exploit the physical dynamics of our bodies. In this study, we investigated whether this principle also applies to visually guided walking over complex terrain. We found that when humans can see the immediate foreground as little as two step lengths ahead, they are able to choose footholds that allow them to exploit their biomechanical structure as efficiently as they can with unlimited visual information. We conclude that when humans walk over complex terrain, they use visual information from two step lengths ahead to choose footholds that allow them to approximate the energetic efficiency of walking in flat, obstacle-free environments.     

Elasticity and movements of the cockroach tarsus in walking

Anatomical, kinematic and ablation studies were performed to evaluate the contribution of elasticity in use of the cockroach tarsus (foot) in walking. The distal tarsus (claws and arolium) engages the substrate during the stance phase of walking by the action of a single muscle, the retractor unguis. Kinematic and ablation studies demonstrated that tarsal disengagement occurs at the end of stance, in part via the action of elastic elements at the penultimate tarsal joint. In isolated legs, this joint exhibits very rapid (less than 20 ms duration) recoil to extension when released from the engaged position, and recoil is even more rapid (less than 10 ms) after removal of the retractor tendon (apodeme). The joint also possesses an enlarged cuticular condyle which is the attachment for ligaments and articular membranes, some of which fulfill morphological criteria consistent with the presence of the elastic protein resilin. Measurements of restoring forces generated by joint displacement indicate that they are graded but could readily lift the mass of the distal tarsus. This biomechanical design can facilitate efficient use of the tarsus in walking while under active control by only a single muscle and may also be highly advantageous when cockroaches very rapidly traverse irregular terrain. Accepted: 16 September 1998     

Coordination of flipflopping neural signals and head turning during pheromone-mediated walking in a male silkworm moth Bombyx mori

Male silkworm moths, Bombyx mori, move their heads side-to-side during zigzag walking toward a source of sex pheromone. High-speed video analysis revealed that changes in walking direction were synchronized with this head turning. Thus the direction of the walking is indicated by the direction of the head turning. Head turning was regulated by neck motor neurons which innervate the cervical ventral muscles and the ventral muscles through the second cervical nerve. To determine the role of the `flipflop' state transition in spike activity carried by descending interneurons from the brain to the thoracic ganglion, we recorded pheromonal responses simultaneously from flipflop descending interneurons and a single cervical ventral 1 neck motor neuron. The activity of the cervical ventral 1 neck motor neuron was synchronized to that of the flipflop descending interneurons. The cervical ventral 1 neck motor neuron was morphologically identified using confocal imaging. Our results demonstrate that the flipflop signals play an important role in instructing turning signals during the pheromone-mediated behavior in a male B. mori. Accepted: 11 June 1998     

An effective balancing response to lateral perturbations at pelvis level during slow walking requires control in all three planes of motion

In this study we investigated balancing responses to lateral perturbations during slow walking (0.85 m/s). A group of seven healthy individuals walked on an instrumented treadmill while being perturbed at the level of waist at left heel strike in outward and inward lateral directions. Centre of mass (COM) and centre of pressure (COP), rotation of pelvis around vertical axis, step lengths, step widths and step times were assessed. The results have shown that beside control of COP in lateral direction, facilitated by adequate step widths, control of COP in sagittal direction, slowing down movement of COM was present after commencement of lateral perturbations. Sagittal component of COM was significantly retarded as compared to unperturbed walking for both inward (4.32 ± 1.29 cm) and outward (9.75 ± 2.17 cm) perturbations. This was necessary since after an inward perturbation first step length (0.29 ± 0.04 m compared to 0.52 ± 0.02 m in unperturbed walking) and step time (0.45 ± 0.05 s compared to 0.61 ± 0.04 s in unperturbed walking) were shortened while after an outward perturbation first two step lengths (0.36 ± 0.05 m and 0.32 ± 0.11 m compared to 0.52 ± 0.03 m in unperturbed walking) were shortened that needed to be accommodated by the described modulation of COP in sagittal plane. In addition pronounced pelvis rotation assisted in bringing swing leg to new location. The results of this study show that counteracting lateral perturbations at slow walking requires adequate response in all three planes of motion.     

Energy transformation during erect and 'bent-hip,bent-knee' walking by humans with implications for the evolution of bipedalism

We have previously reported that predictive dynamic modeling suggests that the 'bent-hip, bent-knee' gait, which some attribute to Australopithecus afarensis AL-288-1, would have been much more expensive in mechanical terms for this hominid than an upright gait. Normal walking by modern adult humans owes much of its efficiency to conservation of energy by transformation between its potential and kinetic states. These findings suggest the question if, and to what extent, energy transformation exists in 'bent-hip, bent-knee' gait.This study calculates energy transformation in humans walking upright, at three different speeds, and walking 'bent-hip, bent-knee'. Kinematic data were gathered from video sequences and kinetic (ground reaction force) data from synchronous forceplate measurement. Applying Newtonian mechanics to our experimental data, the fluctuations of kinetic and potential energy in the body centre of mass were obtained and the effects of energy transformation evaluated and compared. In erect walking the fluctuations of two forms of energy are indeed largely out-of-phase, so that energy transformation occurs and total energy is conserved. In 'bent-hip, bent-knee' walking, however, the fluctuations of the kinetic and potential energy are much more in-phase, so that energy transformation occurs to a much lesser extent. Among all modes of walking the highest energy recovery is obtained in subjectively 'comfortable' walking, the next highest in subjectively 'fast' or 'slow' walking, and the least lowest in 'bent-hip, bent-knee' walking. The results imply that if 'bent-hip, bent-knee' gait was indeed habitually practiced by early bipedal hominids, a very substantial (and in our view as yet unidentified) selective advantage would have had to accrue, to offset the selective disadvantages of 'bent-hip, bent-knee' gait in terms of energy transformation.     

Sprawling locomotion in the lizard Sceloporus clarkii: the effects of speed on gait, hindlimb kinematics, and axial bending during walking

Although the hindlimb is widely considered to provide the propulsive force in lizard locomotion, no study to date has analysed kinematic patterns of hindlimb movements for more than one stride for a single individual and no study has considered limb and axial kinematics together. In this study, kinematic data from several individuals of the Sceloporus clarkii are used to describe the movement patterns of the axial skeleton and hindlimb at different speeds, to analyse how kinematics change with speed, and to compare and contrast these findings with the inferred effects of speed cited in the literature. Angular limb movements and axial bending patterns (standing wave with nodes on the girdles) did not change with speed. Only the relative speed of retracting the femur and flexing the knee during limb retraction changes with speed. Based on these data and similar results from a recent study of salamanders, it appears that, over a range of speeds involving a walking trot, sprawling vertebrates increase speed by simply retracting the femur relatively faster, thus this simple functional adjustment may be a general mechanism to increase speed in tetrapods. The demonstration that femoral retraction alone is the major speed effector in Sceloporus clarkii lends strong functional support to ecomorphological implications of limb length (and especially femur length and caudifemoralis size) in locomotory ecology and performance in phrynosomatid lizards. It also lends support to inferences about the caudifemoralis muscle as a preadaptation to terrestrial locomotion and as a key innovation in the evolution of bipedalism.     

Cineradiographic study of forelimb movements during quadrupedal walking in the brown lemur (Eulemur fulvus, Primates: Lemuridae)

Movements of forelimb joints and segments during walking in the brown lemur (Eulemur fulvus) were analyzed using cineradiography (150 frames/sec). Metric gait parameters, forelimb kinematics, and intralimb coordination are described. Calculation of contribution of segment displacements to stance propulsion shows that scapular retroversion in a fulcrum near the vertebral border causes more than 60% of propulsion. The contribution by the shoulder joint is 30%, elbow joint 5%, and wrist joint 1%. Correlation analysis was applied to reveal the interdependency between metric and kinematic parameters. Only the effective angular movement of the elbow joint during stance is speed-dependent. Movements of all other forelimb joints and segments are independent of speed and influence, mainly, linear gait parameters (stride length, stance length). Perhaps the most important result is the hitherto unknown and unexpected degree of scapular mobility. Scapular movements consist of ante-/retroversion, adduction/abduction, and scapular rotation about the longitudinal axis. Inside rotation of the scapula (60 degrees -70 degrees ), together with flexion in the shoulder joint, mediates abduction of the humerus, which is not achieved in the shoulder joint, and is therefore strikingly different from humeral abduction in man. Movements of the shoulder joint are restricted to flexion and extension. At touch down, the shoulder joint of the brown lemur is more extended compared to that of other small mammals. The relatively long humerus and forearm, characteristic for primates, are thus effectively converted into stride length. Observed asymmetries in metric and kinematic behavior of the left and right forelimb are caused by an unequal lateral bending of the spinal column.     

Assessment of the lower limb soft tissue artefact at marker-cluster level with a high-density marker set during walking

The estimation of joint kinematics from skin markers is hindered by the soft tissue artefact (STA), a well-known phenomenon although not fully characterized. While most assessments of the STA have been performed based on the individual skin markers displacements, recent assessments were based on the marker-cluster geometrical transformations using, e.g., principal component or modal analysis. However, these marker-clusters were generally made of 4–6 markers and the current findings on the STA could have been biased by the limited number of skin makers analysed. The objective of the present study was therefore to confirm them with a high-density marker set, i.e. 40 markers placed on the segments.A larger number of modes than found in the literature was required to describe the STA. Nevertheless, translations and rotations of the marker-cluster remained the main STA modes, archetypally the translation along the proximal-distal and anterior-posterior axes for the shank and the translation along the proximal-distal axis and the rotation about the medial-lateral axis for the thigh. High correlations were also found between the knee flexion angle and the amplitude of these modes for the thigh whereas moderate ones were found for the shank.These findings support the current re-orientation of the STA compensation methods, from bone pose estimators which typically address the non-rigid components of the marker-cluster to kinematic-driven rigid-component STA models.     

Coordination of wing motion and walking suggests common control of zigzag motor program in a male silkworm moth

The male silkworm moth, Bombyx mori, exhibits a zigzagging pattern as it walks upwind to pheromones. This species usually does not fly, but obvious wing-beating accompanies the pheromone-mediated walking. Males supported by a `sled', after having their legs removed, also moved upwind in a pheromone plume along zigzagging tracks, indicating that wing-generated thrust and torque result in locomotory paths similar to those observed from walking moths. Using a high-speed video system we investigated the correlation between the wing movements and zigzag walking. The wing ipsilateral to the direction of the turn showed a greater degree of retraction with respect to the contralateral wing. The timing of the wing retraction pattern was synchronized with changes of direction in the walking track. Coordination of wing movements and walking pattern was not dependent on visual feedback or sensory feedback generated from neck movements associated with turning. The results presented here, taken together with our previous studies of descending interneurons suggest that the coordination of wing movements with the walking pattern may result from the activity of a set of identified interneurons descending from the brain to the thoracic ganglia and/or may be coordinated by coupling of oscillating circuits for walking and wing beating. Accepted: 15 May 1997     

Light-evoked walking in crayfish: Behavioral and neuronal responses triggered by the caudal photoreceptor

The caudal photoreceptors (CPRs) of crayfish (Procambarus clarkii) can trigger walking and abdominal movements by their response to light.

Forest succession and prey availability influence the strength and scale of terrestrial‐aquatic linkages in a headwater salamander system

1. Trophic linkages between terrestrial and aquatic ecosystems are common and sensitive to disruption. However, there is little information on what causes variation in the strength and spatial scale of these linkages. 2. In the highly aquatic adults of the headwater salamander Gyrinophilus porphyriticus (family Plethodontidae), use of terrestrial prey decreases along a gradient from early‐ to late‐successional riparian forests. To understand the cause of this relationship, we tested the predictions that (i) terrestrial prey abundance is lower in late‐successional forests, and (ii) G. porphyriticus adults cannot move as far from the stream to forage in late‐successional forests, thus limiting access to terrestrial prey. 3. We established 100‐m long study reaches on six headwater streams in the Hubbard Brook Experimental Forest, New Hampshire. Three reaches were in early‐successional forests and three were in late‐successional forests. We conducted pitfall trapping for invertebrate prey in June and July of 2005, with three traps at 0, 2, 5 and 10 m from the stream at each reach. In June, July and August of 2004 and 2005, nighttime salamander surveys were conducted at each reach along ten, 10‐m long by 2.5‐m wide transects perpendicular to the stream. 4. Abundance of terrestrial prey was consistently lower in late‐successional forests, suggesting that consumption of terrestrial prey by G. porphyriticus is affected by prey abundance. Contrary to our prediction, G. porphyriticus adults moved farther from the stream in late‐successional forests, suggesting that habitat conditions in late‐successional forests do not limit movement away from the stream, and that lower abundances of terrestrial prey in these forests may cause salamanders to move farther from streams. 5. Our results provide novel insight on the extent of terrestrial habitat use by G. porphyriticus. More broadly, these results indicate that major habitat gradients, such as forest succession, can affect the strength and scale of terrestrial‐aquatic linkages. Application of this insight to the design of vegetation buffers along headwater streams would have widespread benefits to freshwater ecosystems.