The integrated function of muscles and tendons during locomotion

The mechanical roles of tendon and muscle contractile elements during locomotion are often considered independently, but functionally they are tightly integrated. Tendons can enhance muscle performance for a wide range of locomotor activities because muscle-tendon units shorten and lengthen at velocities that would be mechanically unfavorable for muscle fibers functioning alone. During activities that require little net mechanical power output, such as steady-speed running, tendons reduce muscular work by storing and recovering cyclic changes in the mechanical energy of the body. Tendon stretch and recoil not only reduces muscular work, but also allows muscle fibers to operate nearly isometrically, where, due to the force-velocity relation, skeletal muscle fibers develop high forces. Elastic energy storage and recovery in tendons may also provide a key mechanism to enable individual muscles to alter their mechanical function, from isometric force-producers during steady speed running to actively shortening power-producers during high-power activities like acceleration or uphill running. Evidence from studies of muscle contraction and limb dynamics in turkeys suggests that during running accelerations work is transferred directly from muscle to tendon as tendon stretch early in the step is powered by muscle shortening. The energy stored in the tendon is later released to help power the increase in energy of the body. These tendon length changes redistribute muscle power, enabling contractile elements to shorten at relatively constant velocities and power outputs, independent of the pattern of flexion/extension at a joint. Tendon elastic energy storage and recovery extends the functional range of muscles by uncoupling the pattern of muscle fiber shortening from the pattern of movement of the body.     

Vestibular responses of neurons of the descending tracts during locomotion

Responses of vestibulo-, reticulo-, and rubro-spinal neurons to tilting decerebrate cats in the frontal plane were investigated. Tilting was carried out both at rest and during induced locomotion (walking and running on a treadmill). During locomotion the vestibular responses were greatly reduced or they disappeared completely.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 4, No. 3, pp. 311–316, May–June, 1972.     

Experimental antimony toxicity on lower motor neurons and muscles of mice

The occurrence of muscular weakness and wasting in patients with schistosomal infections treated with tartar emetic prompted this investigation of the role of antimony (potassium tartrate) on the lower motor neurons and muscle of mice (lumbar cord, sciatic nerve, myoneural junction, and gastrocnemius). Motor weakness could be demonstrated in the animals and could be correlated with the dosage by testing muscle endurance in an activity cage. The pathological changes in the anterior horn cells are vacuolar degeneration, chromatolysis, satellitosis, and swelling of the nuclei. The edematous sciatic nerves have hydropic degeneration with vacuolation, and fragmentation of axons. The myoneural junctions show swollen ramifications of the axis cylinder and an increase in the number of end-plate nuclei. Some muscle fibers (gastrocnemius) have hyaline and granular alterations with vesicular nuclei, and other atrophic fibers with pyknotic nuclei, but the main changes are in the anterior horn cells, sciatic nerves, and myoneural junction.     

EMG of scapulohumeral muscles in the chimpanzee during reaching and "arboreal" locomotion

Current views on the function of the deltoid and rotator cuff muscles emphasize their roles in arm-raising as participants in a scapulohumeral force "couple." The acceptance of such a mechanism is based primarily on a 1944 EMG study of human shoulder muscle action. More recently, it has been suggested that shoulder joint stabilization constitutes a second and equally important function of the cuff musculature, especially in nonhuman primates which habitually use their forelimbs in overhead postural and locomotor activities. Few comparative data exist, however, on the actual recruitment patterns of these muscles in different species. In order to assess the general applicability of a scapulohumeral force couple model, and the functional significance of the differential development of the scapulohumeral musculature among primate species, we have undertaken a detailed study of shoulder muscle activity patterns in nonhuman primates employing telemetered electromyography, which permits examination of unfettered natural behaviors and locomotion. The results of our research on the chimpanzee, Pan troglodytes, on voluntary reaching and two forms of "arboreal" locomotion reveal four ways in which previous perceptions of the function of the scapulohumeral muscles must be revised: 1) the posterior deltoid is completely different in function from the middle and anterior regions of this muscle; 2) the integrity of the glenohumeral joint during suspensory postures is not maintained solely by osseoligamentous structures; 3) the function of teres minor is entirely different from that of the other rotator cuff muscles and is more similar to the posterior deltoid and teres major; and 4) each remaining member of the rotator cuff plays a distinct, and often unique, role during natural behaviors. These results clearly refute the view that the muscles of the rotator cuff act as a single functional unit in any way, and an alternative to the force couple model is proposed.     

Topography and distribution of motor neurons supplying uropod muscles of a crayfish

• 1.1. The pathway and distribution of motor neurons in the uropod muscles of the crayfish, Procambarus clarkii, was investigated electrophysiologically and histologically.
• 2.2. There were three crossing points of motor neurons between the peripheral motor bundle originating from the second and third roots of the sixth abdominal ganglion.
• 3.3. It seems that there are no anatomical and functional regularity in the innervation pattern of the uropod muscles.

     

Control of ground reaction forces by hindlimb muscles during cat locomotion

It has been proposed that biarticular muscles are primarily responsible for the control of the direction of external forces, as their activation is closely related and highly sensitive to the direction of external forces. This functional role for biarticular muscles has been supported qualitatively by experimental evidence, but has never been tested quantitatively for lack of a mathematical/mechanical formulation of this theory and the difficulty of measuring individual muscle forces during voluntary movements. The purposes of this study were: (1) to define rules for muscular coordination based on the control of external forces; (2) to develop a model of the cat hindlimb that allows for the calculation of the magnitude and direction of the ground reaction forces (GRFs) produced by individual hindlimb muscles; and (3) to test if the coordination of mono- and biarticular cat hindlimb muscles is related to the control of the resultant GRF. We measured the GRF, hindlimb kinematics, selected muscle forces and activations during cat locomotion. Then, the measured muscle forces were used as input to the hindlimb model to compute the muscle-induced GRF. We assume that if activation (and possibly force) increased as the muscle-induced component of GRF approximated the resultant GRF, then that muscle was used by the central nervous system (CNS) to help control the direction of the external GRF. During cat walking, medial gastrocnemius (MG) and plantaris (PL) forces increased with increasing proximity to the GRF, while soleus (SOL) forces and vastus lateralis (VL) activations did not. SOL and VL activation were most strongly related to the vertical and parallel (braking/accelerating) component of the GRF, respectively. We concluded from these results that MG and PL are primarily responsible for the control of the direction of the GRF, while SOL primarily functions as an anti-gravity muscle, and VL as an acceleration/deceleration muscle.     

Classification of lumbosacral neurons by their discharge pattern during evoked locomotion

Unit activity was recorded in the lumbosacral division of the spinal cord during evoked locomotion in mesencephalic cats with the afferent fibers from their hind limbs intact or divided. If the afferent fibers were intact, all neurons recorded showed modulation of activity during locomotion in the rhythm of stepping movements. In experiments on cats with afferent fibers from the hind limbs divided modulation was absent in 30% of neurons, while in the modulated neurons, the frequencies in the excitation phase were approximately the same as when the limb innervation was intact. Modulation of activity in some neurons occurred in response to stimulation of the locomotor region even before stepping movements began. The tuning of the spinal generator of stepping movements is discussed.M. V. Lomonosov Moscow State University. Institute of Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 4, pp. 410–417, July–August, 1972.     

Differential sympathetic outflow and vasoconstriction responses at kidney and skeletal muscles during fictive locomotion

We compared sympathetic and circulatory responses between kidney and skeletal muscles during fictive locomotion evoked by electrical stimulation of the mesencephalic locomotor region (MLR) in decerebrate and paralyzed rats (n = 8). Stimulation of the MLR for 30 s at 40-microA current intensity significantly increased arterial pressure (+38 +/- 6 mmHg), triceps surae muscle blood flow (+17 +/- 3%), and both renal and lumbar sympathetic nerve activities (RSNA +113 +/- 16%, LSNA +31 +/- 7%). The stimulation also significantly decreased renal cortical blood flow (-18 +/- 6%) and both renal cortical and triceps surae muscle vascular conductances (RCVC -38 +/- 5%, TSMVC -17 +/- 3%). The sympathetic and vascular conductance changes were significantly dependent on current intensity for stimulation at 20, 30, and 40 microA. The changes in LSNA and TSMVC were significantly less than those in RSNA and RCVC, respectively, at all current intensities. At the early stage of stimulation (0-10 s), decreases in RCVC and TSMVC were significantly correlated with increases in RSNA and LSNA, respectively. These data demonstrate that fictive locomotion induces less vasoconstriction in skeletal muscles than in kidney because of less sympathetic activation. This suggests that a neural mechanism mediated by central command contributes to blood flow distribution by evoking differential sympathetic outflow during exercise.     

Context-dependent changes in motor control and kinematics during locomotion: modulation and decoupling

Successful locomotion through complex, heterogeneous environments requires the muscles that power locomotion to function effectively under a wide variety of conditions. Although considerable data exist on how animals modulate both kinematics and motor pattern when confronted with orientation (i.e. incline) demands, little is known about the modulation of muscle function in response to changes in structural demands like substrate diameter, compliance and texture. Here, we used high-speed videography and electromyography to examine how substrate incline and perch diameter affected the kinematics and muscle function of both the forelimb and hindlimb in the green anole (Anolis carolinensis). Surprisingly, we found a decoupling of the modulation of kinematics and motor activity, with kinematics being more affected by perch diameter than by incline, and muscle function being more affected by incline than by perch diameter. Also, muscle activity was most stereotyped on the broad, vertical condition, suggesting that, despite being classified as a trunk-crown ecomorph, this species may prefer trunks. These data emphasize the complex interactions between the processes that underlie animal movement and the importance of examining muscle function when considering both the evolution of locomotion and the impacts of ecology on function.