Muscular torque generation during imposed joint rotation: torque-angle relationships when subjects' only goal is to make a constant effort

It is a reasonable expectation that voluntarily activated spinal motoneurons will be further excited by increases in spindle afferent activity produced by muscle stretch. Human motor behavior attributed to tonic stretch reflexes and to reflexes recruited by relatively slow joint rotation has been reported from several laboratories. We reinvestigated this issue by rotating the elbow joint over the central portion of its range while subjects focused on keeping their elbow flexion effort constant at one of three different levels and made no attempt to control the position, speed or direction of movement of their forearm. There is evidence that subjects' voluntary motor status is constant under these conditions so that any change in torque would be of involuntary origin. On average, torques rose somewhat and then fell as the elbow was flexed through a range of 80 degrees at 10, 20 and 60 degrees/s and a similar pattern occurred during elbow extension; i.e., both concentric and eccentric torque-angle profiles had roughly similar shapes and neither produced consistent stabilizing cross-range stiffness. The negative stiffness (rising torque) during the early part of a concentric movement and the negative stiffness (falling torque) during the later part of an eccentric movement would not have occurred if a stabilizing stretch reflex had been present. Positive stiffness rarely gave rise to torque changes greater than 20% in either individual or cross-subject averaged data. When angular regions of negative stiffness are combined with regions of low positive stiffness (torque change 10% or less), much of the range of motion was not well stabilized, especially during eccentric movements. The sum of the EMGs from biceps brachii, brachioradialis and brachialis showed a pattern opposite to that expected for a stretch reflex; there was an upward trend in the EMG as the elbow was flexed and a downward trend as the elbow was extended. There was little change in the shape of this EMG-angle relationship with either direction or velocity. The individual EMG-angle relationships were distinctive for each of these three elbow flexor muscles in four of the six subjects; in the remaining two, biceps was distinctive, but brachioradialis and brachialis appeared to be coupled. Although the EMGs of individual muscles were modulated over the angular range, no consistent stretch reflexes could be seen in the individual records. Thus, we could find no clear evidence for stretch reflex stabilization of human subjects maintaining a constant effort. Rather, muscle torque appears to be reflexly modulated across a much used portion of the elbow's angular range so that any appreciable stabilizing stiffness that is sustained for more than fractions of a second is associated with a change in effort.     

Muscular torque generation during imposed joint rotation: torque-angle relationships when subjects' only goal is to make a constant effort

It is a reasonable expectation that voluntarily activated spinal motoneurons will be further excited by increases in spindle afferent activity produced by muscle stretch. Human motor behavior attributed to tonic stretch reflexes and to reflexes recruited by relatively slow joint rotation has been reported from several laboratories. We reinvestigated this issue by rotating the elbow joint over the central portion of its range while subjects focused on keeping their elbow flexion effort constant at one of three different levels and made no attempt to control the position, speed or direction of movement of their forearm. There is evidence that subjects' voluntary motor status is constant under these conditions so that any change in torque would be of involuntary origin. On average, torques rose somewhat and then fell as the elbow was flexed through a range of 80° at 10, 20 and 60°/s and a similar pattern occurred during elbow extension; i.e., both concentric and eccentric torque-angle profiles had roughly similar shapes and neither produced consistent stabilizing cross-range stiffness. The negative stiffness (rising torque) during the early part of a concentric movement and the negative stiffness (falling torque) during the later part of an eccentric movement would not have occurred if a stabilizing stretch reflex had been present. Positive stiffness rarely gave rise to torque changes greater than 20% in either individual or cross-subject averaged data. When angular regions of negative stiffness are combined with regions of low positive stiffness (torque change 10% or less), much of the range of motion was not well stabilized, especially during eccentric movements. The sum of the EMGs from biceps brachii, brachioradialis and brachialis showed a pattern opposite to that expected for a stretch reflex; there was an upward trend in the EMG as the elbow was flexed and a downward trend as the elbow was extended. There was little change in the shape of this EMG-angle relationship with either direction or velocity. The individual EMG-angle relationships were distinctive for each of these three elbow flexor muscles in four of the six subjects; in the remaining two, biceps was distinctive, but brachioradialis and brachialis appeared to be coupled. Although the EMGs of individual muscles were modulated over the angular range, no consistent stretch reflexes could be seen in the individual records. Thus, we could find no clear evidence for stretch reflex stabilization of human subjects maintaining a constant effort. Rather, muscle torque appears to be reflexly modulated across a much used portion of the elbow's angular range so that any appreciable stabilizing stiffness that is sustained for more than fractions of a second is associated with a change in effort.     

Long-lasting inhibition of the soleus muscle H reflex related to realization of voluntary arm movements in humans

In healthy humans, we recorded the H reflex induced by transcutaneous stimulation of the tibial nerve (recording from the soleus muscle). In subjects in the lying position, we studied changes in the H reflex values after preceding voluntary arm movements realized with a maximum velocity after presentation of an acoustic signal. On the 200th to 300th msec after forearm flexion, long-lasting inhibition of the H reflex developed following a period of initial facilitation and reached the maximum, on average, 700 msec from the moment of the movement. Flexion of the contralateral upper limb in the elbow joint induced deeper inhibition than analogous movement of the ipsilateral arm. Long-lasting clear inhibition of the H reflex developed after arm flexion in the elbow joint but was slightly expressed after finger clenching. After inhibition reached the maximum, its time course was satisfactorily approximated by a logarithmic function of the time interval between the beginning of the conditioning voluntary movement and presentation of the test stimulus. Durations of inhibition calculated using a regression equation were equal to 6.6 sec and 8.5 sec after ipsilateral and contralateral elbow-joint flexions, respectively. Inhibition was not eliminated under conditions of tonic excitation of motoneurons of the tested muscle upon voluntary foot flexion. Long-lasting inhibition of the H reflex was also observed after electrical stimulation-induced flexions of the upper limb. The obtained data indicate that movements of the upper limb cause reflex long-lasting presynaptic inhibition of the soleus-muscle H reflex that can play a noticeable role in redistribution of the muscle tone during motor activity. Neirofiziologiya/Neurophysiology, Vol. 40, No. 3, pp. 221–227, May–June, 2008.     

Time-course analysis of stretch reflexes in hemiparetic subjects using an on-line spasticity measurement system

Spasticity after a stroke is usually assessed in a score form by subjectively determining the resistance of a joint to an externally imposed passive movement. This work presents a spasticity measurement system for on-line quantifying the stretch reflex of paretic limbs. Four different constant stretch velocities in a ramp-and-hold mode are used to elicit the stretch reflex of the elbow joint in spastic subjects. The subjects are tested at supine position with the upper limb stretched towards the ground, in contrast with the horizontally stretched movement used in other studies. By subtracting the baseline torque, reflex torque measured at a selected low stretch velocity of 5 deg/sec, the influence of gravity torque and inertial in vertical stretching mode can be minimized. The averaged speed-dependent reflex torque (ASRT), defined as the measured torque deviated from the baseline torque, is used for quantifying the spastic hypertonia. Four subjects having incurred cerebrovascular accident (CVA) are recruited for time-course study in which the measurements are taken at 72 hours, one week, one month, three months, and six months after onset of stroke. During the development of spasticity, the changes of ASRT and velocity sensitivity of ASRT of the involved and the intact elbow joints are discussed.     

Using the Threat Probability Task to Assess Anxiety and Fear During Uncertain and Certain Threat

Fear of certain threat and anxiety about uncertain threat are distinct emotions with unique behavioral, cognitive-attentional, and neuroanatomical components. Both anxiety and fear can be studied in the laboratory by measuring the potentiation of the startle reflex. The startle reflex is a defensive reflex that is potentiated when an organism is threatened and the need for defense is high. The startle reflex is assessed via electromyography (EMG) in the orbicularis oculi muscle elicited by brief, intense, bursts of acoustic white noise (i.e., “startle probes”). Startle potentiation is calculated as the increase in startle response magnitude during presentation of sets of visual threat cues that signal delivery of mild electric shock relative to sets of matched cues that signal the absence of shock (no-threat cues). In the Threat Probability Task, fear is measured via startle potentiation to high probability (100% cue-contingent shock; certain) threat cues whereas anxiety is measured via startle potentiation to low probability (20% cue-contingent shock; uncertain) threat cues. Measurement of startle potentiation during the Threat Probability Task provides an objective and easily implemented alternative to assessment of negative affect via self-report or other methods (e.g., neuroimaging) that may be inappropriate or impractical for some researchers. Startle potentiation has been studied rigorously in both animals (e.g., rodents, non-human primates) and humans which facilitates animal-to-human translational research. Startle potentiation during certain and uncertain threat provides an objective measure of negative affective and distinct emotional states (fear, anxiety) to use in research on psychopathology, substance use/abuse and broadly in affective science. As such, it has been used extensively by clinical scientists interested in psychopathology etiology and by affective scientists interested in individual differences in emotion.     

Muscle activity in rapid multi-degree-of-freedom elbow movements: solutions from a musculoskeletal model

The activity of certain muscles that cross the elbow joint complex (EJC) are affected by forearm position and forearm movement during elbow flexion/extension. To investigate whether these changes are based on the musculoskeletal geometry of the joint, a three-dimensional musculotendinoskeletal computer model of the EJC was used to estimate individual muscle activity in multi-degree-of-freedom (df) rapid (ballistic) elbow movements. It is hypothesized that this model could reproduce the major features of elbow muscle activity during multi-df elbow movements using dynamic optimal control theory, given a minimum-time performance criterion. Results from the model are presented and verified with experimental kinematic and electromyographic data from movements that involved both one-df elbow flexion/extension and two-df flexion/extension with forearm pronation/supination. The model demonstrated how the activity of particular muscles is affected by both forearm position and movement, as measured in these experiments and as previously reported by others. These changes were most evident in the flexor muscles and least evident in the extensor muscles. The model also indicated that, for specific one- and two-df movements, activating a muscle that is antagonistic or noncontributory to the movement could reduce the movement time. The major features of muscle activity in multi-df elbow movements appear to be highly dependent on the joint's musculoskeletal geometry and are not strictly based on neural influences or neuroanatomical substrates. Received: 9 May 1997 / Accepted in revised form: 8 December 1998     

Upper limb H reflexes and somatosensory evoked potentials modulated by movement.

In the human lower limb, the magnitudes of both Hoffmann (H) reflexes and primary somatosensory evoked potentials (SEPs) from scalp electrodes, are reduced by active and/or passive movement. We surmised that similar effects occur for the upper limb and specifically hypothesised that amplitudes of median nerve induced flexor carpii radialis H reflexes and cortical SEPs are reduced with passive movement about the wrist or elbow. The results showed (P<0. 05) that either movement significantly attenuated mean magnitudes of SEPs elicited from stimulation at elbow or wrist and that reflex magnitudes attenuated with wrist movement. Thus, the upper limb shows similar movement-induced modulation to the lower limb. These attenuations of fast conducting sensory paths consequent to movement per se, may be a basic level of motor control, initiated from muscle mechanoreceptor discharge. Upon this basic level, more complex modulations then may be laid as appropriate for the particular characteristics of active motor tasks.     

A comparison of the effects of agonist and antagonist muscle fatigue on performance of rapid movements

The aim of this study was to investigate the effects of agonist and antagonist muscle fatigue on the performance of rapid, self-terminating movements. Six subjects performed rapid, consecutive elbow flexion and extension movements between two targets prior to and after fatiguing either the elbow flexor or elbow extensor muscles. The experiments demonstrated consistent results. Agonist muscle fatigue was associated with a decrease in peak velocity and peak deceleration, while a decrease in peak acceleration was particularly prominent. Antagonist muscle fatigue, however, was associated with a decrease in peak deceleration, while a decrease in both the peak velocity and peak acceleration was modest and, in some tests, non-significant. The relative acceleration time (i.e. acceleration time as a proportion of the total movement time) increased when agonists were fatigued, but decreased when antagonists were fatigued. Taken together, these results emphasize the mechanical roles of the agonist and antagonist muscles; namely, the fatigue of each muscle group particularly affected the movement phase in which that group accelerated a limb, while changes of the movement kinematics pattern provided more time for action of the fatigued muscles. In addition, the results presented suggest that agonist muscle fatigue affects movement velocity more than antagonist muscle fatigue, even in movements that demonstrate prominently both mechanical and myoelectric activity of the antagonist muscles, such as rapid, self-terminating movements. Accepted: 11 February 1997     

Induction of Enhanced Acoustic Startle Response by Noise Exposure: Dependence on Exposure Conditions and Testing Parameters and Possible Relevance to Hyperacusis

There has been a recent surge of interest in the development of animal models of hyperacusis, a condition in which tolerance to sounds of moderate and high intensities is diminished. The reasons for this decreased tolerance are likely multifactorial, but some major factors that contribute to hyperacusis are increased loudness perception and heightened sensitivity and/or responsiveness to sound. Increased sound sensitivity is a symptom that sometimes develops in human subjects after acoustic insult and has recently been demonstrated in animals as evidenced by enhancement of the acoustic startle reflex following acoustic over-exposure. However, different laboratories have obtained conflicting results in this regard, with some studies reporting enhanced startle, others reporting weakened startle, and still others reporting little, if any, change in the amplitude of the acoustic startle reflex following noise exposure. In an effort to gain insight into these discrepancies, we conducted measures of acoustic startle responses (ASR) in animals exposed to different levels of sound, and repeated such measures on consecutive days using a range of different startle stimuli. Since many studies combine measures of acoustic startle with measures of gap detection, we also tested ASR in two different acoustic contexts, one in which the startle amplitudes were tested in isolation, the other in which startle amplitudes were measured in the context of the gap detection test. The results reveal that the emergence of chronic hyperacusis-like enhancements of startle following noise exposure is highly reproducible but is dependent on the post-exposure thresholds, the time when the measures are performed and the context in which the ASR measures are obtained. These findings could explain many of the discrepancies that exist across studies and suggest guidelines for inducing in animals enhancements of the startle reflex that may be related to hyperacusis.     

Influence of long-latency reflex modulation on the performance of quick adjustment movements

The effect of long-latency reflex modulation on the performance of a quick adjustment movement following a muscle stretch was studied in 26 healthy male subjects. When the subjects felt a sudden angle displacement in the direction of a wrist extension they were required to make an adjustment movement by moving a handlebar, held in the hand, to align with a target position as quickly and as accurately as possible. The index of performance (adjustment time) was the time taken to move the handle to the target position from stretch onset. A DC torque motor was used to evoke electromyographic (EMG) reflex responses on a wrist flexor. Averaging of the rectified EMG, recorded from surface electrodes placed over the flexor, showed short- and long-latency reflexes (M1 and M2 components). For all subjects, the amplitudes of the reflex components decreased during the adjustment movement because the target position for this study was fixed to the extension side of the wrist joint. The decrease in the M2 component, which is considered to be a transcortical reflex, was significantly larger than the decrease in the M1 component, which is spinal reflex. The main finding was of a positive correlation between the length of adjustment time and the degree of reduction of M1 and M2 with the adjustment movement (r = 0.602 for M1, P < 0.01; r = 0.850 for M2, P < 0.001). Moreover, there were correlations between the consistency of the voluntary response onset and the degree of M2 decrease (r = 0.577, P < 0.01), and between the consistency of the voluntary response onset and the length of the adjustment time (r = 0.603, P < 0.01). Therefore, we have concluded that the subjects who were able to perform adjustment movements within a short time could modulate the long-latency reflex of the muscle involved in such movements in order to make the function of their voluntary muscle activity more effective, and thus were able to respond appropriately. Accepted: 19 February 1997     

The effects of neonatal saline injections on behaviours in DBA/2 male mice

DBA/2 male mice were exposed to the injections of the saline (0.01 ml/g i.p.) on 1-th, 3-th, 5-th, 7-th, 9-th days after birth. Intact males were used as a control group. Adult saline-treated males displayed the increased number of crossed squares, entries in the centre and time spent in the centre during the open "field" test in comparison with intact animals. The time spent in the light compartment of the light-dark box was decreased in saline treated mice compared with intact animals. During the test of acoustic startle response the magnitude of startle reflex and prepulse inhibition didn't change the startle reflex. Saline administration in males did not affect corticosterone basal level. Sexual motivation was revealed to decrease in saline treated males. These data suggest that neonatal administration of saline induced a stable behavioral syndrome in adult DBA/2 male mice: hyperactivity, a decrease of open space fear and simultaneously an increase of some indices of anxiety.     

The effect of neonatal administration of monosodium glutamate on behavior and blood corticosterone level

DBA/2 male mice were treated with monosodium glutamate (MSG) in a dose of 4 mg/g on 1, 3, 5, 7, 9 days after birth. Saline treated and intact males were used as control groups. MSG treated males displayed decreased number of crossed squares, rearings, entries in the centre and time in the centre of open field in comparison with saline-treated but not intact animals. Time in the light compartment of the light-dark box was increased in MSG-treated mice versus both saline treated and intact animals. MSG administration reduced acoustic startle response but did not affect the magnitude of prepulse inhibition of the startle reflex. Sexual motivation in male mice was reduced by MSG, the same trend was observed after saline treatment. MSG administration increased corticosterone basal level 4-fold while saline treatment did not affect it. These data suggest that neonatal administration of MSG decreases locomotion, exploratory activity, anxiety in male mice, while corticosterone level is increased. Saline treatment increases these parameters (except sexual motivation), and this augmentation is not connected to changes in corticosterone basal level.     

Control of the Elbow Extensor Muscles in Slow Targeted Extensions of the Arm in Humans

Relations between the kinematic parameters of slow (non-ballistic) targeted extension movements in the elbow joint of humans and characteristics of the movement-related EMG activity in the two heads of the m. triceps brachii were analyzed. Test movements were performed under conditions of application of non-inertional external loadings directed toward flexion. It was shown that the movement-related EMG activity of the elbow extensors, similarly to what was observed in the flexors at flexion movements with the same parameters, demonstrates a complex structure and includes dynamic and stationary phases. In the former phase, in turn, initial and main components can be differentiated. The rising edge and decay of the main component of the dynamic extensor EMG phase could be approximated by exponential functions; this component was never split into a few subcomponents. Dependences between the amplitudes of m. triceps brachii EMG phases and the amplitude of the movement (or external loading) were, as a rule, nonlinear but monotonic. An increase in the test movement velocity led to an increase in the rate of rise of the rising edge of the dynamic EMG phase, while an increment in the amplitude was less significant. Under the used test conditions, the activity of the elbow extensors was usually accompanied by some coactivation of the antagonists (m. biceps brachii). It is concluded that motor commands coming to the elbow extensors at performance of the extension test movements differ from motor commands to the flexors at analogous flexion test movements by a simpler structure and more tonic pattern. Biomechanical specificities of fixation of the mentioned muscle groups to the arm bones (stability of the moment for application of the extensor force under conditions of changing the joint angle vs variable moment of the flexor force) are considered one of the main reasons for such specificity of the patterns of the extensor and flexor motor commands.     

Efferent activity in the phrenic nerve during startle reflex in chloralose anesthetized cats

Efferent activity was investigated in the phrenic nerve during startle reflex manifesting as somatic nerve discharges (lower intercostal nerves and the nerve endings) in chloralose anesthetized cats. Inhibition (usually of short duration, lasting 23–36 msec) of inspiration activity was found to be the main component of response in the phrenic nerve in the shaping of "low threshold" startle reflex produced by acoustic and tactile stimuli and stimulation of low threshold peripheral afferents. Reflex discharge prevailed amongst the response patterns produced in the phrenic nerve by stimulating high threshold afferents, i.e., early (propriospinal) and late (suprasegmental, arising from stimulating intercostal nerve) or late only (when stimulating the hindlimb nerves). Two patterns of late response could be distinguished, one on inspiration (found in roughly 3 out of 4 experiments) and other on exhalation — the respiratory homologs of somatic startle reflex. Response pattern is described throughout the respiratory cycle. Structure and respiratory modulation of reflex responses produced in the phrenic nerve by stimulating bulbar respiratory structure are also examined. Possible neurophysiological mechanisms underlying phrenic response during the shaping of startle reflex are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 4, pp. 473–482, July–August, 1987.     

A forward genetic screen identifies Dolk as a regulator of startle magnitude through the potassium channel subunit Kv1.1

The acoustic startle response is an evolutionarily conserved avoidance behavior. Disruptions in startle behavior, particularly startle magnitude, are a hallmark of several human neurological disorders. While the neural circuitry underlying startle behavior has been studied extensively, the repertoire of genes and genetic pathways that regulate this locomotor behavior has not been explored using an unbiased genetic approach. To identify such genes, we took advantage of the stereotypic startle behavior in zebrafish larvae and performed a forward genetic screen coupled with whole genome analysis. We uncovered mutations in eight genes critical for startle behavior, including two genes encoding proteins associated with human neurological disorders, Dolichol kinase (Dolk), a broadly expressed regulator of the glycoprotein biosynthesis pathway, and the potassium Shaker-like channel subunit Kv1.1. We demonstrate that Kv1.1 and Dolk play critical roles in the spinal cord to regulate movement magnitude during the startle response and spontaneous swim movements. Moreover, we show that Kv1.1 protein is mislocalized in dolk mutants, suggesting they act in a common genetic pathway. Combined, our results identify a diverse set of eight genes, all associated with human disorders, that regulate zebrafish startle behavior and reveal a previously unappreciated role for Dolk and Kv1.1 in regulating movement magnitude via a common genetic pathway.     

Differences in stretch reflex responses of elbow flexor muscles during shortening, lengthening and isometric contractions

Stretch reflexes were evoked in elbow flexor muscles undergoing three different muscle contractions, i.e. isotonic shortening (SHO) and lengthening (LEN), and isometric (ISO) contractions. The intermuscle relationships for the magnitude of the stretch reflex component in the eletromyographic (EMG) activities of two main elbow flexor muscles, i.e. the biceps brachii (BB) and the brachioradialis (BRD), were compared among the three types of contractions. The subjects were requested to move their forearms sinusoidally (0.1 Hz) against a constant pre-load between elbow joint angles of 10° (0° = full extension) and 80° during SHO and LEN, and to keep an angle of 45° during the ISO. The perturbations were applied at the elbow angle of 45° in pseudo-random order. The EMG signals were rectified and averaged over a period of 100 ms before and 400 ms after the onset of the perturbation 40–50 times. From the ensemble averaged EMG waveform, the background activity (BGA), short (20–50 ms) and long latency (M2, 50–80, M3, 80–100 ms) reflex and voluntary activity (100–150 ms) components were measured. The results showed that both BGA and reflex EMG activity of the two elbow flexor muscles were markedly decreased during the lengthening contraction compared to the SHO and ISO contractions. Furthermore, the changes of reflex EMG components in the BRD muscle were more pronounced than those in the BB muscle, i.e. the ratios of M2 and M3 magnitudes between BRD and BB (BRD:BB) were significantly reduced during the LEN contractions. These results would suggest that the gain of long latency stretch reflex EMG activities in synergistic muscles might be modulated independently according to the model of muscle contraction. Accepted: 1 September 1997     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

Multisegmental motor activity in the segmentally restricted gin trap behavior in Manduca sexta pupae

Stimulation of sensory neurons innervating hairs in the gin traps on the abdomen of Manduca sexta pupae evokes a rapid bending of the abdomen that is restricted to one or more of the three articulating posterior segments. However, electrical stimulation of the gin trap sensory nerve in an isolated abdominal nerve cord evokes characteristic motor neuron activity in every abdominal segment. To determine if the segmentally distributed motor activity also occurred in intact animals and how it contributed to the segmentally restricted reflex movement, mechanical stimulation of the sensory hairs in intact animals was used to evoke reflex responses that were recorded as electromyograms synchronized with video recordings of the behavior. Motor activity was monitored during movements to determine if there was activity in many segments when the movement was restricted to one segment. Coordinated muscle activity was evoked throughout the abdomen in response to stimulation of any of the three gin traps, even when movement was restricted to one segment. Differences in the timing of ipsilateral and contralateral motor activity among segments allowed the closing of gin traps to be segmentally restricted. These findings suggest that the neural circuit underlying the gin trap reflex is distributed throughout the abdominal nerve cord. This network generates a complex, yet coordinated, motor pattern with muscular activity in many abdominal segments that produces a localized bending reflex. Accepted: 10 January 1997     

Improving elbow torque output of stroke patients with assistive torque controlled by EMG signals

This paper develops an assistive torque system which uses homogeneic surface electromyogram (EMG) signals to improve the elbow torque capability of stroke patients by applying an external time-varying assistive torque. In determining the magnitude of the torque to apply, the incorporated assistive torque algorithm considers the difference between the weighted biceps and triceps EMG signals such that the applied torque is proportional to the effort supplied voluntarily by the user. The overall stability of the assistive system is enhanced by the incorporation of a nonlinear damping element within the control algorithm which mimics the physiological damping of the elbow joint and the co-contraction between the biceps and triceps. Adaptive filtering of the control signal is employed to achieve a balance between the bandwidth and the system adaptability so as to ensure a smooth assistive torque output. The innovative control algorithm enables the provision of an assistive system whose operation is both natural to use and simple to learn. The effectiveness of the proposed assistive system in assisting elbow movement performance is investigated in a series of tests involving five stroke patients and five able-bodied individuals. The results confirm the ability of the system to assist all of the subjects in performing a number of reaching and tracking tasks with reduced effort and with no sacrifice in elbow movement performance.     

Interaction between acoustic startle and habituated neck postural responses in seated subjects.

Postural and startle responses rapidly habituate with repeated exposures to the same stimulus, and the first exposure to a seated forward acceleration elicits a startle response in the neck muscles. Our goal was to examine how the acoustic startle response is integrated with the habituated neck postural response elicited by forward accelerations of seated subjects. In experiment 1, 14 subjects underwent 11 sequential forward accelerations followed by 5 additional sled accelerations combined with a startling tone (124-dB sound pressure level) initiated 18 ms after sled acceleration onset. During the acceleration-only trials, changes consistent with habituation occurred in the root-mean-square amplitude of the neck muscles and in the peak amplitude of five head and torso kinematic variables. The subsequent addition of the startling tone restored the amplitude of the neck muscles and four of the five kinematic variables but shortened onset of muscle activity by 9-12 ms. These shortened onset times were further explored in experiment 2, wherein 16 subjects underwent 11 acceleration-only trials followed by 15 combined acceleration-tone trials with interstimulus delays of 0, 13, 18, 23, and 28 ms. Onset times shortened further for the 0- and 13-ms delays but did not lengthen for the 23- and 28-ms delays. These temporal and spatial changes in EMG can be explained by a summation of the excitatory drive converging at or before the neck muscle motoneurons. The present observations suggest that habituation to repeated sled accelerations involves extinguishing the startle response and tuning the postural response to the whole body disturbance.