Testing steering functions of the frontal zone in the locomotion of amoeba proteus

Light and chemical stimuli were asymmetrically applied to the advancing front of amoeba without affecting any body region behind the frontal zone. Stimulation limited to the front alone was sufficient to control the frontal expansion and, as a further consequence, the locomotion and shape of the whole cell. Contracting factors applied locally to the front inhibited it, whereas the relaxing agents activated its expansion.     

Rat muscle blood flows during high-speed locomotion

We previously studied blood flow distribution within and among rat muscles as a function of speed from walking (15 m/min) through galloping (75 m/min) on a motor-driven treadmill. The results showed that muscle blood flows continued to increase as a function of speed through 75 m/min. The purpose of the present study was to have rats run up to maximal treadmill speeds to determine if blood flows in the muscles reach a plateau as a function of running speed over the animals' normal range of locomotory speeds. Muscle blood flows were measured with radiolabeled microspheres at 1 min of running at 75, 90, and 105 m/min in male Sprague-Dawley rats. The data indicate that even at these relatively high treadmill speeds there was still no clear evidence of a plateau in blood flow in most of the hindlimb muscles. Flows in most muscles continued to increase as a function of speed. These observed patterns of blood flow vs. running speed may have resulted from the rigorous selection of rats that were capable of performing the high-intensity exercise and thus only be representative of a highly specific population of animals. On the other hand, the data could be interpreted to indicate that the cardiovascular potential during exercise is considerably higher in laboratory rats than has normally been assumed and that inadequate blood flow delivery to the muscles does not serve as a major limitation to their locomotory performance.     

Fundamental patterns of bilateral muscle activity in human locomotion

Human gait is characterized by smooth, regular and repeating movements but the control system is complex: there are many more actuators (i.e. muscles) than degrees of freedom in the system. Statistical pattern-recognition techniques have been applied to examine muscle activity signals, but these have all concentrated exclusively on unilateral gait. We report here the application of factor analysis to the electromyographic patterns of 16 muscles (eight bilateral pairs) in ten normal subjects. Consistent with our prior work, we have established two factors, named loading response and propulsion, which correspond with important phases in the gait cycle. In addition, we have also discovered a third factor, which we have named the coordinating factor, that maintains the phase shift between the left and right sides. These findings suggest that the central nervous system solves the problem of high dimensionality by generating a few fundamental signals which control the major muscle groups in both legs.     

Myosin dilated cardiomyopathy mutation S532P disrupts actomyosin interactions,leading to altered muscle kinetics,reduced locomotion,and cardiac dilation in Drosophila

Dilated cardiomyopathy (DCM), a life-threatening disease characterized by pathological heart enlargement, can be caused by myosin mutations that reduce contractile function. To better define the mechanistic basis of this disease, we employed the powerful genetic and integrative approaches available in Drosophila melanogaster. To this end, we generated and analyzed the first fly model of human myosin–induced DCM. The model reproduces the S532P human β-cardiac myosin heavy chain DCM mutation, which is located within an actin-binding region of the motor domain. In concordance with the mutation’s location at the actomyosin interface, steady-state ATPase and muscle mechanics experiments revealed that the S532P mutation reduces the rates of actin-dependent ATPase activity and actin binding and increases the rate of actin detachment. The depressed function of this myosin form reduces the number of cross-bridges during active wing beating, the power output of indirect flight muscles, and flight ability. Further, S532P mutant hearts exhibit cardiac dilation that is mutant gene dose–dependent. Our study shows that Drosophila can faithfully model various aspects of human DCM phenotypes and suggests that impaired actomyosin interactions in S532P myosin induce contractile deficits that trigger the disease.     

Role of speed vs. grade in relation to muscle pump function at locomotion onset.

We sought to clarify the roles of contraction frequency (speed) and contraction force (grade) in the rise in muscle blood flow at the onset of locomotion. Shoemaker et al. (Can J Physiol Pharmacol 76: 418-427, 1998) explored this relationship in human handgrip exercise and found that the time course of the rise in muscle vascular conductance was similar when a light weight was lifted in a fast cadence and a heavy weight was lifted in a slow cadence (total work constant). This indicates that muscle pumping (contraction frequency) was of limited importance in governing the time course. Rather, vasodilator substances released in proportion to the total work performed appeared to determine the pattern and extent of the rise in conductance. We hypothesized that conductance would rise faster during locomotion at a high speed (frequency) and low grade (force) than at a low speed and high grade, despite similar total increases in conductance, owing to more effective muscle pumping at faster contraction rates. Seven male rats performed nine 1-min bouts of treadmill locomotion across a combination of three speeds (5, 10, and 20 m/min) and three grades (-10, 0, and +15 degrees ) in random order. Locomotion at 10 m/min and 0 degrees grade and 20 m/min and -10 degrees grade led to an equal rise in terminal aortic vascular conductance. However, the equal rise was achieved more quickly at the higher running speed, suggestive of more effective muscle pumping. Across the nine combinations of exercise, speed began to exert a statistically significant influence on conductance by the 3rd s of locomotion. Grade did not begin to exert an influence until the 12th s of locomotion (similar to the delays reported for arteriolar dilation to muscle contraction). Additional experiments in dogs provided similar results. Thus the muscle pump appears to initiate the increase in blood flow in proportion to contraction frequency at locomotion onset.     

Functional diversification within and between muscle synergists during locomotion

Locomotion arises from the complex and coordinated function of limb muscles. Yet muscle function is dynamic over the course of a single stride and between strides for animals moving at different speeds or on variable terrain. While it is clear that motor unit recruitment can vary between and within muscles, we know little about how work is distributed within and between muscles under in vivo conditions. Here we show that the lateral gastrocnemius (LG) of helmeted guinea fowl (Numida meleagris) performs considerably more work than its synergist, the medial gastrocnemius (MG) and that the proximal region of the MG (pMG) performs more work than the distal region (dMG). Positive work done by the LG was approximately twice that of the proximal MG when the birds walked at 0.5 ms -1, and four times when running at 2.0 m s-1. This is probably due to different moments at the knee, as well as differences in motor unit recruitment. The dMG performed less work than the pMG because its apparent dynamic stiffness was greater, and because it exhibited a greater recruitment of slow-twitch fibres. The greater compliance of the pMG leads to increased stretch of its fascicles at the onset of force, further enhancing force production. Our results demonstrate the capacity for functional diversity between and within muscle synergists, which increases with changes in gait and speed.     

Influence of locomotion speed on biomechanical subtask and muscle synergy

This paper investigates the relationship of biomechanical subtasks, and muscle synergies with various locomotion speeds. Ground reaction force (GRF) of eight healthy subjects is measured synchronously by force plates of treadmill at five different speeds ranging from 0.5 m/s to 1.5 m/s. Four basic biomechanical subtasks, body support, propulsion, swing, and heel strike preparation, are identified according to GRF. Meanwhile, electromyography (EMG) data, used to extract muscle synergies, are collected from lower limb muscles. EMG signals are segmented periodically based on GRF with the heel strike as the split points. Variability accounted for (VAF) is applied to determine the number of muscle synergies. We find that four muscle synergies can be extracted in all five situations by non-negative matrix factorization (NMF). Furthermore, the four muscle synergies and biomechanical subtasks keep invariant as the walking speed changes.     

Effects of surface grade on proximal hindlimb muscle strain and activation during rat locomotion.

Sonomicrometry and electromyography were used to determine how surface grade influences strain and activation patterns in the biceps femoris and vastus lateralis of the rat. Muscle activity is generally present during much of stance and is most intense on an incline, intermediate on the level, and lowest on a decline, where the biceps remains inactive except at high speeds. Biceps fascicles shorten during stance, with strains ranging from 0.07-0.30 depending on individual, gait, and grade. Shortening strains vary significantly among grades (P = 0.05) and average 0.21, 0.16, and 0.14 for incline, level, and decline walking, respectively; similar trends are present during trotting and galloping. Vastus fascicles are stretched while active over the first half of stance on all grades, and then typically shorten over the second half of stance. Late-stance shortening is highest during galloping, averaging 0.14, 0.10, and 0.02 in the leading limb on incline, level, and decline surfaces, respectively. Our results suggest that modulation of strain and activation in these proximal limb muscles is important for accommodating different surface grades.     

Artificial neural network model for the generation of muscle activation patterns for human locomotion.

Skilled locomotor behaviour requires information from various levels within the central nervous system (CNS). Mathematical models have permitted researchers to simulate various mechanisms in order to understand the organization of the locomotor control system. While it is difficult to adequately characterize the numerous inputs to the locomotor control system, an alternative strategy may be to use a kinematic movement plan to represent the complex inputs to the locomotor control system based on the possibility that the CNS may plan movements at a kinematic level. We propose the use of artificial neural network (ANN) models to represent the transformation of a kinematic plan into the necessary motor patterns. Essentially, kinematic representation of the actual limb movement was used as the input to an ANN model which generated the EMG activity of 8 muscles of the lower limb and trunk. Data from a wide variety of gait conditions was necessary to develop a robust model that could accommodate various environmental conditions encountered during everyday activity. A total of 120 walking strides representing normal walking and ten conditions where the normal gait was modified in terms of cadence, stride length, stance width or required foot clearance. The final network was assessed on its ability to predict the EMG activity on individual walking trials as well as its ability to represent the general activation pattern of a particular gait condition. The predicted EMG patterns closely matched those recorded experimentally, exhibiting the appropriate magnitude and temporal phasing required for each modification. Only 2 of the 96 muscle/gait conditions had RMS errors above 0.10, only 5 muscle/gait conditions exhibited correlations below 0.80 (most were above 0.90) and only 25 muscle/gait conditions deviated outside the normal range of muscle activity for more than 25% of the gait cycle. These results indicate the ability of single network ANNs to represent the transformation between a kinematic movement plan and the necessary muscle activations for normal steady state locomotion but they were also able to generate muscle activation patterns for conditions requiring changes in walking speed, foot placement and foot clearance. The abilities of this type of network have implications towards both the fundamental understanding of the control of locomotion and practical realizations of artificial control systems for use in rehabilitation medicine.     

Neck muscle fatigue and spatial orientation during stepping in place in humans.

Neck proprioceptive input, as elicited by muscle vibration, can produce destabilizing effects on stance and locomotion. Neck muscle fatigue produces destabilizing effects on stance, too. Our aim was to assess whether neck muscle fatigue can also perturb the orientation in space during a walking task. Direction and amplitude of the path covered during stepping in place were measured in 10 blindfolded subjects, who performed five 30-s stepping trials before and after a 5-min period of isometric dorsal neck muscle contraction against a load. Neck muscle electromyogram amplitude and median frequency during the head extensor effort were used to compute a fatigue index. Head and body kinematics were recorded by an optoelectronic system, and stepping cadence was measured by sensorized insoles. Before the contraction period, subjects normally stepped on the spot or drifted forward. After contraction, some subjects reproduced the same behavior, whereas others reduced their forward progression or even stepped backward. The former subjects showed minimal signs of fatigue and the latter ones marked signs of fatigue, as quantified by the dorsal neck electromyogram index. Head position and cadence were unaffected in either group of subjects. We argue that the abnormal fatigue-induced afferent input originating in the receptors transducing the neck muscle metabolic state can modulate the egocentric spatial reference frame. Notably, the effects of neck muscle fatigue on orientation are opposite to those produced by neck proprioception. The neck represents a complex source of inputs capable of modifying our orientation in space during a locomotor task.     

Rat hindlimb muscle blood flow during level and downhill locomotion

Duringeccentrically biased exercise (e.g., downhill locomotion), whole bodyoxygen consumption and blood lactate concentrations are lower thanduring level locomotion. These general systemic measurements indicatethat muscle metabolism is lower during downhill exercise. This studywas designed to test the hypothesis that hindlimb muscle blood flow iscorrespondingly lower during downhill vs. level exercise. Muscle bloodflow (determined by using radioactive microspheres) was measured inrats after 15 min of treadmill exercise at 15 m/min on the level (L,0°) or downhill (D, 17°). Blood flow to ankle extensormuscles was either lower (e.g., white gastrocnemius muscle: D, 9 ± 2; L, 15 ± 1 ml · min¹ · 100 g¹) or not different(e.g., soleus muscle: D, 250 ± 35; L, 230 ± 21 ml · min¹ · 100 g¹) in downhill vs. levelexercise. In contrast, blood flow to ankle flexor muscles was higher(e.g., extensor digitorum longus muscle: D, 53 ± 5; L, 31 ± 6 ml · min¹ · 100 g¹) during downhill vs.level exercise. When individual extensor and flexor muscle flows weresummed, total flow to the leg was lower during downhill exercise (D,3.24 ± 0.08; L, 3.47 ± 0.05 ml/min). These data indicate thatmuscle blood flow and metabolism are lower during eccentrically biasedexercise but are not uniformly reduced in all active muscles; i.e.,flows are equivalent in several ankle extensor muscles and higher inankle flexor muscles.     

Delay of muscle vasodilation to changes in work rate (treadmill grade) during locomotion.

Previous studies examining the delay to the onset of vasodilation have primarily focused on the onset of exercise, a setting complicated by the fact that the muscle pump and the vasodilator systems are both activated, making it difficult to attribute changes in blood flow to one or both. The goal here was to determine the delay to the onset of vasodilation after changes in work rate imposed by changes in treadmill grade (work intensity) during locomotion at a steady speed. The rationale was that constant speed would help ensure constant muscle pump activity (contraction frequency) such that vasodilator responses could be examined in isolation. Seven Sprague-Dawley rats underwent three trials each in which treadmill incline was suddenly ( approximately 1 s) elevated from -10 degrees to +10 degrees. The delay to the onset of vasodilation averaged 5.0 +/- 1.8 s, and this delay was not altered by inhibition of nitric oxide synthase. Similar or longer delays were seen during sinusoidal exercise. Thus there is a significant delay before the onset of vasodilation after an increase in work intensity (muscle force) during locomotory exercise at constant speed.     

Visuomotor coordination in locomotion.

This article reviews the recent literature concerning the role of visual information in the control of locomotion with an emphasis on the neurophysiological mechanisms that underlie visually triggered, voluntary, gait modifications. Data are presented to show how these gait modifications may be encoded by the motor cortex, and how they may interact with the basic locomotor rhythm.     

Mechanical efficiency of locomotion in females during different kinds of muscle action

The mechanical efficiencies (ME) of pure positive and pure negative work as well as of stretch-shortening cycle (SSC) exercise were investigated with a special sledge apparatus. The subjects were 20 young females who performed six different types of submaximal exercise: two of pure concentric exercise (positive work), two of pure eccentric exercise (negative work) and two SSC exercises. The work intensities were determined individually, from the recordings of distance obtained during a single maximal concentric exercise. Each exercise involved 60 muscle actions lasting a total of 3 min per testing condition. The MEs of pure positive work with intensities of 30% and 60% maximum (C30 and C60 respectively) were 15.5%, SD 2.6% and 14.3%, SD 1.9%, respectively. In pure negative work, when the dropping heights were 20 cm (E20) and 80 cm (E80), MEs were 28.4%, SD 6.9% and 47.9%, SD 10.1%, respectively. In SSC-exercise, the MEs during the positive phase of the take-off were 31.3%, SD 6.3% (E20/C90) and 35.0%, SD 7.0% (E80/C69). The total MEs in SSC-exercise were 29.1%, SD 4.0% (E20/C90) and 40.1%, SD 5.2% (E80/C60 x 100). In pure negative work, the increased stretching velocity increased the value of ME. In the concentric phase of SSC-exercise, the integrated electromyographic activity (iEMG) of vastus lateralis (VL) and vastus medialis (VM) muscles were lower (P less than 0.05) than in pure concentric work, when the mechanical work was the same (C60 vs E80/C60). During pure eccentric work, iEMGs were lower in comparison to the eccentric phase of SSC-exercise.(ABSTRACT TRUNCATED AT 250 WORDS)     

Trunk muscle electromyography and whole body vibration

By measuring the electromyographic (EMG) activity of the paraspinal muscles, we have estimated the average and peak-to-peak torque imposed on the spine during whole body vibration. Six subjects had surface electrodes placed on their erector spinae muscles at the L3 level. The EMG-torque relationship was estimated by having each subject perform isometric horizontal pulls in an upright seated posture. The subject was then vibrated vertically and sinusoidally in a controlled, flexed, slightly lordotic seated posture, in 1 Hz increments from 3 to 10 Hz at a 0.1 g RMS seat acceleration level. Between vibration readings taken at each frequency, a static reading was also taken with the subject maintaining the same posture. The entire vibration-static 3-10 Hz test was repeated for reliability purposes. Specialized digital signal processing techniques were developed for the EMG signals to enhance the measured cyclic muscle activity and to allow automatic measurement of the time relationship between the mechanical displacement and the estimated torque. We found significantly more average and peak-to-peak estimated torque at almost all frequencies for vibration vs static sitting.