Maturation of the flight motor pattern without movement inManduca sexta

The development of the flight motor pattern was studied by recording acutely with fine wire electrodes inserted in the thoracic muscles of pharate moths of known age and by recording chronically for up to 8 days with implanted electrodes. Externally visible morphological characteristics by which the age of a pharateManduca sexta can be established were identified (Table 1). Bouts of activity lasting approximately 30 min to 2 h and alternating with inactive periods of similar duration were recorded as early as the ninth day after pupation and on all successive days until early on the day of eclosion, typically 19 days after pupation (Figs. 1,5). During the 3 days preceding the day of eclosion a rhythmic flight motor pattern was produced (Fig. 2). The rhythmic activity ceased 5^1/2–10^1/2 h before eclosion and only an occasional, large potential change was recorded from the thoracic muscles during this time (Fig. 3). During the 3 days of rhythmic activity the percent-age of time that the animal was active did not change (Fig. 4). The flight motor pattern matured, in that the cycle-time decreased and became less variable (Fig. 6). The approximate flight phase relationship between an elevator muscle and the dorsal longitudinal depressor muscle did not become less variable as the cycle-time improved. The flight motor pattern produced by pharate moths caused neither movement of the scutum nor an increase in thoracic temperature in marked contrast to the consequences of adult motor activity (Fig. 7). Intracellular recording from the dorsal longitudinal muscle of pharate moths 20–30 h before eclosion showed that, after repeated stimulation of the motor nerve at 2/s, only small junctional potentials were elicited (Fig. 8). A burst of 6 stimuli at 50/s elicited 2–5 active membrane responses and a contraction. These observations explain the absence of thoracic movement in immature animals producing the flight motor pattern and the presence of movement in immature animals stimulated to eclose. They also show that the neuromuscular junction matures rapidly during the day before eclosion.     

Direction of movement is encoded in the human primary motor cortex

The present study investigated how direction of hand movement, which is a well-described parameter in cerebral organization of motor control, is incorporated in the somatotopic representation of the manual effector system in the human primary motor cortex (M1). Using functional magnetic resonance imaging (fMRI) and a manual step-tracking task we found that activation patterns related to movement in different directions were spatially disjoint within the representation area of the hand on M1. Foci of activation related to specific movement directions were segregated within the M1 hand area; activation related to direction 0° (right) was located most laterally/superficially, whereas directions 180° (left) and 270° (down) elicited activation more medially within the hand area. Activation related to direction 90° was located between the other directions. Moreover, by investigating differences between activations related to movement along the horizontal (0°+180°) and vertical (90°+270°) axis, we found that activation related to the horizontal axis was located more anterolaterally/dorsally in M1 than for the vertical axis, supporting that activations related to individual movement directions are direction- and not muscle related. Our results of spatially segregated direction-related activations in M1 are in accordance with findings of recent fMRI studies on neural encoding of direction in human M1. Our results thus provide further evidence for a direct link between direction as an organizational principle in sensorimotor transformation and movement execution coded by effector representations in M1.     

The problem of estimating wind drift in migrating birds

Whether migrating birds compensate for wind drift or not is a fundamental question in bird migration research. The procedures to demonstrate and quantitatively estimate wind drift or compensation are fraught with difficulties and pitfalls. In this paper, we evaluate four methods that have been used in several studies over the past decades. We evaluate the methods by analysing a model migratory movement with a realistic scatter in flight directions, for the ideal cases of full drift and complete compensation. Results obtained with the different methods are then compared with the "true behaviour" of the model movement, illustrating that spurious patterns of drift and compensation arise in some cases. We also illustrate and evaluate the different methods of estimating drift for a real case, based on tracking radar measurements of bird migration in relation to winds. Calculating the linear regression of mean geographic track (resulting flight direction) and heading directions (directions of the birds' body axis) of a migratory movement under different wind conditions in relation to the angle alpha (the angle between mean track and heading) always provides robust and reliable results. Comparing mean flight directions between occasions with winds from the left and right of the mean flight direction of the whole migratory movement also always provides expected and correct measures of drift. In contrast, regressions of individual flight directions in relation to alpha (the angle between track and heading for the specific individuals or flocks) are liable to produce biased and spurious results, overestimating compensation/overcompensation if following winds dominate in the analysis and overestimating drift/overdrift if opposed winds are dominating. Comparing mean directions for cases with winds from the left and right in relation to individual flight directions also gives biased and spurious results unless there is full variation in wind directions or an equal distribution of crosswinds from left and right. The results of the methodological evaluation and the analysis of the real case indicate that some earlier analyses of wind drift may have to be re-evaluated.     

A method for estimating torque-vector directions of shoulder muscles using surface EMGs

In this study, a new method is proposed to estimate the torque-vector directions of each shoulder muscle. The method is based on a multiple regression model that reconstructs shoulder torque, which is calculated from the hand force and posture, from the surface EMG of many muscles recorded simultaneously. The torque-vector directions of eleven shoulder muscles of four subjects were obtained at up to 30 different arm postures with this method. The mean confidence interval ( p< 0.05) of the estimated torque-vector direction of each subject was 7.7-10.6 degrees. The correlation coefficient between the measured shoulder torque and reconstructed shoulder torque was between 0.76-0.84. The results for majority of the muscles were in accordance with previous studies, and reasonable from the viewpoint of anatomy. The torque-vector directions of a muscle, which are estimated with this method, have more of a functional meaning than a pure anatomical or mechanical one. These indicate the direction of the shoulder torque accompanying the muscle activation for a normal shoulder action that involves the cooperative contraction of many muscles.     

A method for estimating torque-vector directions of shoulder muscles using surface EMGs

In this study, a new method is proposed to estimate the torque-vector directions of each shoulder muscle. The method is based on a multiple regression model that reconstructs shoulder torque, which is calculated from the hand force and posture, from the surface EMG of many muscles recorded simultaneously. The torque-vector directions of eleven shoulder muscles of four subjects were obtained at up to 30 different arm postures with this method. The mean confidence interval (p < 0.05) of the estimated torque-vector direction of each subject was 7.7-10.6 degrees. The correlation coefficient between the measured shoulder torque and reconstructed shoulder torque was between 0.76-0.84. The results for majority of the muscles were in accordance with previous studies, and reasonable from the viewpoint of anatomy. The torque-vector directions of a muscle, which are estimated with this method, have more of a functional meaning than a pure anatomical or mechanical one. These indicate the direction of the shoulder torque accompanying the muscle activation for a normal shoulder action that involves the cooperative contraction of many muscles.     

Heterogeneity of muscle activation in relation to force direction: A multi-channel surface electromyography study on the triceps surae muscle

Several skeletal muscles can be divided into sub-modules, called neuromuscular compartments (NMCs), which are thought to be controlled independently and to have distinct biomechanical functions. We looked for distinct muscle activation patterns in the triceps surae muscle (TS) using surface electromyography (EMG) during voluntary contraction. Nine subjects performed isometric and isotonic plantar flexions combined with forces along pre-defined directions. Besides the forces under the ball of the foot, multi-channel surface EMG was measured with electrodes homogeneously distributed over the entire TS. Using principal component analysis, common (global) components were omitted from the EMG signals, thereby estimating muscle activity sufficiently accurate to track fine fluctuations of force during an isotonic contraction (r = 0.80 ± 0.09). A subsequent cluster analysis showed a topographical organization of co-activated parts of the muscle that was different between subjects. Low and negative correlations between the EMG activity within clusters were found, indicating a substantial heterogeneity of TS activation. The correlations between cluster time series and forces at the foot in specific directions differed substantially between clusters, showing that the differentially activated parts of the TS had specific biomechanical functions.     

弥猴单侧软腭肌肉对针式电极刺激的反应

目的建立针电极口内刺激猴软腭肌肉诱发腭咽闭合运动的模式,取得软腭肌肉运动的有效刺激数值,为软腭肌肉功能重建奠定基础。方法通过解剖成年猕猴软腭的五组肌肉,确定其体表位置;利用实验动物用腭部肌肉电极定位刺激器及针式电极对软腭肌肉进行有效刺激;结合鼻咽纤维镜、头颅侧位X片及软腭造影技术观察、记录肌肉收缩及腭咽闭合动作。结果在猕猴口内定位目标肌肉进行针电极刺激可诱发肌肉收缩。刺激电压为3 V、刺激频率为20 Hz时均能诱发单侧软腭肌肉的有效收缩;单侧腭帆提肌在刺激电压为5 V、20 Hz时可发生腭咽闭合动作。咽腭肌、舌腭肌在刺激电压5 V、刺激频率100 Hz时发生软腭下降动作。腭帆张肌仅发生收缩,而未发生腭咽闭合。应用鼻咽纤维镜和X线成像技术配合能记录腭咽闭合动作。结论弥猴可作为研究软腭肌肉运动模式的实验动物。应用电极刺激软腭肌肉,可初步建立腭咽闭合的动作模式。     

Turning manoeuvres in free-flying locusts: two-channel radio-telemetric transmission of muscle activity

A device has been constructed allowing the simultaneous transmission of two separate electrical signals in unrestrained small animals. We employed this device to investigate the motor output in free-flying locusts. The activation pattern of several combinations of different muscles was recorded, including bilateral symmetric muscles and pairs of antagonists. Particular attention was paid to the recruitment of a specific set of flight muscles in both winged segments during rolling manoeuvres. The relationship of the muscle activation with wing movement was analysed in combination with a high-speed video-monitoring. The muscles are activated in advance of the relevant stroke directions, in opposition to previous studies of tethered flying locusts. During turning manoeuvres a statistically significant difference in timing of the bilateral symmetric muscles is not apparent; this contrasts with the distinct difference revealed for the bilateral wing movement. It is discussed that rolling might rely on the fine tuned interaction of several major flight muscles or on the precise activation of a specific wing hinge muscle. Correspondence with investigations of bird flight is discussed.     

A nerve clamp electrode design for indirect stimulation of skeletal muscle

A nerve clamp electrode was developed to indirectly stimulate skeletal muscle innervated by α motor neurons as an alternative to conventional electrodes. The stimulating electrode device consists of a spring coil-activated nerve clamp mounted inside a 1-mL syringe barrel. Supramaximal pulses were generated by a Grass stimulator and delivered to the nerve segment via the nerve clamp electrode. The salient feature of the electrode is its ability to produce muscle contractions indirectly through stimulation of the attached nerve. Indirect muscle stimulation is critical for studying the paralytic actions of presynaptic-acting toxins such as botulinum neurotoxins (BoNT), a potent inhibitor of acetylcholine (ACh) release from α motor neurons. This device enables stimulation of muscle contraction indirectly as opposed to contraction from direct muscle stimulation. The electrode is able to stimulate indirect muscle contraction when tested on ex vivo preparations from rodent phrenic nerve-hemidiaphragm muscle in similar fashion to conventional electrodes. In addition, the electrode stimulated external intercostal nerve-muscle preparations. This was confirmed after applying BoNT serotype A, a potent inhibitor of ACh release, to induce muscle paralysis. Alternative methods, including suction and bipolar loop electrodes, were unsuccessful in stimulating indirect muscle contraction. Therefore, this novel electrode is useful for physiological assessment of nerve agents and presynaptic actions of toxins that cause muscle paralysis. This electrode is useful for stimulating nerve-muscle preparations for which the length of nerve is a concern.     

Acoustic myography during voluntary isometric contraction reveals non-propagative lateral vibration

When isolated muscle synchronous contraction is evoked during in vitro twitches, mechanical vibrations at the surface of the muscle reflect resonant behavior. In contrast, voluntary contraction corresponds to the asynchronous contraction of recruited motor units, therefore, this kind of excitation could lead to different muscle vibrational behavior. We have studied human biceps brachii muscle during voluntary contraction in 10 healthy subjects. Low and high levels of voluntary contraction were explored with simultaneous recording of surface vibration by two sensors located longitudinally or perpendicular to the muscle's main axis. Cross-correlation and coherence functions were computed. Coherence functions revealed a common vibration frequency band between 17 and 28 Hz. Cross correlation functions revealed in-phase vibration for longitudinal sensors and opposite phase vibration for perpendicular sensors thus confirming a lateral bending movement. This behavior suggests that the acoustic myogram is the response of the muscle as a global resonant structure to the local fluctuations of pressure.     

Optimum timing of muscle activation for simple models of throwing.

In diverse throwing activities, muscles contract in sequence, starting with those furthest from the hand. This paper uses simple mathematical models, each with just two muscles, to investigate the consequences of this sequential contraction. One model was suggested by shot putting, another by underarm throwing and the third by overarm throwing, but all are much simpler than real human movements. In each case there is an optimum delay between activation of the more proximal muscle and of the more distal one, that maximizes the speed at which the missile leaves the hand. If the delay is shorter than optimal, the throw is completed sooner and less time is available for contraction of the proximal muscle: it may shorten faster, exerting less torque, or through less than its full range of movement, and so do less work. If it is longer than optimal, less time is available for contraction of the distal muscle, which therefore does less work. The optimal delay is in some cases longer than would maximize total work because the delay influences the proportion of the work that appears as kinetic energy of the missile.     

Dependence of elbow joint stiffness measurements on speed,angle, and muscle contraction level

Elbow joint stiffness is critical to positioning the hand. Abnormal elbow joint stiffness may affect a person's ability to participate in activities of daily living. In this work, elbow joint stiffness was measured in ten healthy young adults with a device adapted from one previously used to measure stiffness in other joints. Measurements of elbow stiffness involved applying a constant-velocity rotational movement to the elbow and measuring the resultant displacement, torque, and acceleration. Elbow stiffness was then computed using a previously-established model for joint stiffness. Measurements were made at two unique elbow joint angles, two speeds, and two forearm muscle contraction levels. The results indicate that the elbow joint stiffness is significantly affected by both rotational speed and forearm muscle contraction level.     

Muscles within muscles: Coordination of 19 muscle segments within three shoulder muscles during isometric motor tasks.

The aim of the present study was to determine how the intra-muscular segments of three shoulder muscles were coordinated to produce isometric force impulses around the shoulder joint and how muscle segment coordination was influenced by changes in movement direction, mechanical line of action and moment arm (ma). Twenty male subjects (mean age 22 years; range 18-30 years) with no known history of shoulder pathologies, volunteered to participate in this experiment. Utilising an electromyographic technique, the timing and intensity of contraction within 19 muscle segments of three superficial shoulder muscles (Pectoralis Major, Deltoid and Latissimus Dorsi) were studied and compared during the production of rapid (e.g. approximately 400ms time to peak) isometric force impulses in four different movement directions of the shoulder joint (flexion, extension, abduction and adduction). The results of this investigation have suggested that the timing and intensity of each muscle segment's activation was coordinated across muscles and influenced by the muscle segment's moment arm and its mechanical line of action in relation to the intended direction of shoulder movement (e.g. flexion, extension, abduction or adduction). There was also evidence that motor unit task groups were formed for individual motor tasks which comprise motor units from both adjacent and distant muscles. It was also confirmed that for any particular motor task, individual muscle segments can be functionally classified as prime mover, synergist or antagonist - classifications which are flexible from one movement to the next.     

Characteristics and lateral dominance of hand grip and elbow flexion powers in young male adults

This study aimed to clarify the characteristics and the lateral dominance of hand grip power and elbow flexion power. The subjects were 15 healthy young males (mean age 22.1+/-0.7 yr, mean height 171.3+/-3.4 cm, mean mass 64.5+/-4.1 kg). All subjects were right-handed. Peak power was measured by both hands with 6 different loads of 20%-70% of maximum voluntary contraction. The maximum voluntary contraction of hand grip movement and elbow flexion movement was significantly larger in the dominant hand. Peak power of the dominant hand was larger in all loads in hand grip movement and in loads of 20% and 30% of maximum voluntary contraction in elbow flexion movement. In short, lateral dominance was confirmed. Peak power was significantly larger in hand grip movement than in elbow flexion movement in both hands. Peak velocity decreased with increasing loads in both movements, but peak power increased until about 50% of maximum voluntary contraction and then decreased. The peak power ratio of the dominant hand to the nondominant hand was significantly larger in hand grip movement than in elbow flexion movement in all loads and the peak power ratio in elbow flexion movement was more marked in light loads. In conclusion, both powers showed lateral dominance. Lateral dominance is more marked in hand grip power.     

Optimal design of vertebrate and insect sarcomeres

This paper offers a model for the normalized length-tension relation of a muscle fiber based upon sarcomere design. Comparison with measurements published by Gordon et al. ('66) shows an accurate fit as long as the inhomogeneity of sarcomere length in a single muscle fiber is taken into account. Sequential change of filament length and the length of the cross-bridge-free zone leads the model to suggest that most vertebrate sarcomeres tested match the condition of optimal construction for the output of mechanical energy over a full sarcomere contraction movement. Joint optimization of all three morphometric parameters suggests that a slightly better (0.3%) design is theoretically possible. However, this theoretical sarcomere, optimally designed for the conversion of energy, has a low normalized contraction velocity; it provides a poorer match to the combined functional demands of high energy output and high contraction velocity than the real sarcomeres of vertebrates. The sarcomeres in fish myotomes appear to be built suboptimally for isometric contraction, but built optimally for that shortening velocity generating maximum power. During swimming, these muscles do indeed contract concentrically only. The sarcomeres of insect asynchronous flight muscles contract only slightly. They are not built optimally for maximum output of energy across the full range of contraction encountered in vertebrate sarcomeres, but are built almost optimally for the contraction range that they do in fact employ.     

Fast actions in small animals: springs and click mechanisms

Small animals that jump or perform predatory strikes depend on much higher limb accelerations than larger animals. To overcome the temporal restrictions of muscle contraction, some arthropod muscles slowly load spring-like structures with potential energy. In flight, sound generation, jumping, or predatory strikes arthropods employ different strategies to transform muscular action to the desired movement. Click mechanisms control the frequency of oscillating spring — muscle systems while other accessory structures such as snap mechanism or latches with trigger mucles determined the stability and control the timing of the instantaneous discharge in catapult mechanisms.     

Electromyography of superficial and deep neck muscles during isometric, voluntary, and reflex contractions

Increasingly complex models of the neck neuromusculature need detailed muscle and kinematic data for proper validation. The goal of this study was to measure the electromyographic activity of superficial and deep neck muscles during tasks involving isometric, voluntary, and reflexively evoked contractions of the neck muscles. Three male subjects (28-41 years) had electromyographic (EMG) fine wires inserted into the left sternocleidomastoid, levator scapulae, trapezius, splenius capitis, semispinalis capitis, semispinalis cervicis, and multifidus muscles. Surface electrodes were placed over the left sternohyoid muscle. Subjects then performed: (i) maximal voluntary contractions (MVCs) in the eight directions (45 deg intervals) from the neutral posture; (ii) 50 N isometric contractions with a slow sweep of the force direction through 720 deg; (iii) voluntary oscillatory head movements in flexion and extension; and (iv) initially relaxed reflex muscle activations to a forward acceleration while seated on a sled. Isometric contractions were performed against an overhead load cell and movement dynamics were measured using six-axis accelerometry on the head and torso. In all three subjects, the two anterior neck muscles had similar preferred activation directions and acted synergistically in both dynamic tasks. With the exception of splenius capitis, the posterior and posterolateral neck muscles also showed consistent activation directions and acted synergistically during the voluntary motions, but not during the sled perturbations. These findings suggest that the common numerical-modeling assumption that all anterior muscles act synergistically as flexors is reasonable, but that the related assumption that all posterior muscles act synergistically as extensors is not. Despite the small number of subjects, the data presented here can be used to inform and validate a neck model at three levels of increasing neuromuscular-kinematic complexity: muscles generating forces with no movement, muscles generating forces and causing movement, and muscles generating forces in response to induced movement. These increasingly complex data sets will allow researchers to incrementally tune their neck models' muscle geometry, physiology, and feedforward/feedback neuromechanics.     

Effects of different movement directions on electromyography recorded from the shoulder muscles while passing the target positions

PurposeWe compared electromyography (EMG) recorded from the shoulder joint muscles in the same position for different movement directions.MethodsFifteen healthy subjects participated. They performed shoulder elevation from 0° to 120°, shoulder depression from 120° to 0°, shoulder horizontal adduction from ?15° to 105°, and shoulder horizontal abduction from 105° to ?15°. The target positions were 90° shoulder elevation in the 0°, 30°, 60°, and 90° planes (0°, 30°, 60°, and 90° positions). EMG signals were recorded from the supraspinatus (SSP) muscle by fine-wire electrodes. EMG signals from the infraspinatus (ISP), anterior deltoid, middle deltoid, and posterior deltoid muscles were recorded using active surface electrodes.ResultsDuring elevation and horizontal abduction, the SSP showed significantly higher activity than that shown during depression and during horizontal adduction in the 0°, 30°, and 60° positions. During elevation, the ISP showed significantly higher activity than during depression and during horizontal adduction in the 90° position. During horizontal abduction, the ISP showed significantly higher activity than during depression in the 90° position.ConclusionsWhen the movement tasks were performed in different movement directions at the same speed, each muscle showed characteristic activity.     

Work organization of a group of human motoneurons during voluntary muscular contraction

Needle electrodes were used to record action potentials of motor units of the rectus femoris muscle during isometric contraction (up to 50% of maximal). Up to 10 motor units working simultaneously could be identified. Under strictly stable conditions of muscular contraction the recruitment order of the motoneurons was constant. The firing rate was inversely proportional as a rule to this recruitment order. As a rule the changes in frequency connected with voluntary contraction of measured strength were in the same direction for different motoneurons. Statistical analysis of the frequency fluctuations observed during contraction of constant strength revealed direct correlation between them. The behavior of the motoneurons as described above is regarded as the result of the diffuse, indeterminate distribution of the synaptic input in the group of motoneurons innervating the muscle studied. It was also shown that even under stable conditions individual motoneurons or groups of them sometimes fired independently. During the performance of different types of movements, the firing rates of the recruited motoneurons varied in different directions and some motoneurons were replaced by others. This shows that when motoneurons function under natural conditions they use not only a common (indeterminate) but also a determinate input.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 5, No. 1, pp. 77–87, January–February, 1973.     

The myosin motor: muscle contraction and in vitro movement

The molecular mechanism of in vitro movement is assumed, by most investigators, to be identical to that of muscle contraction. We discuss this view, which raises various problems. We believe there are mechanisms for muscle contraction (in this case considerable forces are developed, with small displacements) and other mechanisms for in vitro movement (giving large displacements, without necessarily generating substantial forces). Hybrid models may explain muscle contraction. The traditional swinging-crossbridge model may explain in vitro movement. For muscle contraction, movement may result partly from the swinging-crossbridge mechanism and partly from other factors. Comparisons of different fibres at different moments of the Mg-ATPase cycle suggest that both the value of the isometric force in muscle and in vitro and that of the Mg-ATPase activity used in vitro need to be reconsidered. The recently reported dependence of the isometric active tension of smooth skinned fibres on temperature appears to be weaker than predicted by the swinging-crossbridge theory alone. This recent observation is compatible with the existence of other forces (electrostatic repulsions) decreasing with temperature as has been known for some years. From recent experimental data, we think the biochemistry of myosin and actomyosin should be reassessed, to try to find new details of the mechanisms of muscle contraction and in vitro motility.