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Torque-speed relationship of the flagellar rotary motor of Escherichia coli.

The output of a rotary motor is characterized by its torque and speed. We measured the torque-speed relationship of the flagellar rotary motor of Escherichia coli by a new method. Small latex spheres were attached to flagellar stubs on cells fixed to the surface of a glass slide. The angular speeds of the spheres were monitored in a weak optical trap by back-focal-plane interferometry in solutions containing different concentrations of the viscous agent Ficoll. Plots of relative torque (viscosity x speed) versus speed were obtained over a wide dynamic range (up to speeds of approximately 300 Hz) at three different temperatures, 22.7, 17.7, and 15.8 degrees C. Results obtained earlier by electrorotation (, Biophys. J. 65:2201-2216) were confirmed. The motor operates in two dynamic regimes. At 23 degrees C, the torque is approximately constant up to a knee speed of nearly 200 Hz, and then it falls rapidly with speed to a zero-torque speed of approximately 350 Hz. In the low-speed regime, torque is insensitive to changes in temperature. In the high-speed regime, it decreases markedly at lower temperature. These results are consistent with models in which torque is generated by a powerstroke mechanism (, Biophys. J. 76:580-587).

Biomechanical Phenomena↗

Study of the torque of the bacterial flagellar motor using a rotating electric field.

Bacterial flagella are driven by a rotary motor that is energized by an electrochemical ion gradient across the cell membrane. In this study the torque generated by the flagellar motor was measured in tethered cells of a smooth-swimming Escherichia coli strain by using rotating electric fields to determine the relationship between the torque and speed over a wide range. By measuring the electric current applied to the sample cell and combining the data obtained at different viscosities, the torque of the flagellar motor was estimated up to 55 Hz, and also at negative rotation rates. By this method we have found that the torque of the flagellar motor linearly decreases with rotation rate from negative through positive rate of rotation. In addition, the dependence of torque upon temperature was also investigated. We showed that torque at the high speeds encountered in swimming cells had a much steeper dependence on temperature that at the low speeds encountered in tethered cells. From these results, the activation energy of the proton transfer reaction in the torque-generating unit was calculated to be about 7.0 x 10(-20) J.

Biomechanical Phenomena↗

Isometric muscle length-tension curves do not predict angle-torque curves of human wrist in continuous active movements.

In this study we tested the hypothesis that during steady contractions of human wrist extensors or flexors, the torque-angle relationship during movements imposed about the wrist is predicted by the classical isometric muscle length-tension curve, with ascending, descending and ascending limbs. Angle-torque relationships were measured during steady muscle activation (10% of maximal voluntary contraction: MVC), elicited either by electrical stimulation or voluntary regulation of the electromyogram (EMG). Flexion-extension movements of constant speed (+/-10 degrees /s) were imposed on the subjects' hands with a servo actuator, either through the full physiological range of motion +/-50 degrees, or through +/-10 degrees. During extensor contractions, angle-torque curves in +/-50 degrees movements had ascending, descending and ascending limbs, as in isometric contractions. However, in +/-10 degrees movements, torque always increased with increasing muscle length and decreased with decreasing length, even over angles corresponding to the descending limb of isometric curves. For flexor activation, angle-torque curves had similar properties, though descending limbs were less obvious or absent. During imposed movements, hysteresis was observed in the angle-torque curves. This was attributed to non-linearities of the active muscles. Hysteresis reached a maximum at intermediate wrist angles and declined at maximal muscle length, contradicting the recent hypothesis that sarcomere non-uniformity is responsible for the hysteresis. We conclude that the classical isometric length-tension curve, with its prominent descending limb, does not predict angle-torque curves of human wrist muscles in continuous movements. A more appropriate model is one in which stiffness about the wrist is always positive and hysteresis is a significant factor.

Adult↗

Ankle and hip postural strategies defined by joint torques.

Previous studies have identified two discrete strategies for the control of posture in the sagittal plane based on EMG activations, body kinematics, and ground reaction forces. The ankle strategy was characterized by body sway resembling a single-segment-inverted pendulum and was elicited on flat support surfaces. In contrast, the hip strategy was characterized by body sway resembling a double-segment inverted pendulum divided at the hip and was elicited on short or compliant support surfaces. However, biomechanical optimization models have suggested that hip strategy should be observed in response to fast translations on a flat surface also, provided the feet are constrained to remain in contact with the floor and the knee is constrained to remain straight. The purpose of this study was to examine the experimental evidence for hip strategy in postural responses to backward translations of a flat support surface and to determine whether analyses of joint torques would provide evidence for two separate postural strategies. Normal subjects standing on a flat support surface were translated backward with a range of velocities from fast (55 cm/s) to slow (5 cm/s). EMG activations and joint kinematics showed pattern changes consistent with previous experimental descriptions of mixed hip and ankle strategy with increasing platform velocity. Joint torque analyses revealed the addition of a hip flexor torque to the ankle plantarflexor torque during fast translations. This finding indicates the addition of hip strategy to ankle strategy to produce a continuum of postural responses. Hip torque without accompanying ankle torque (pure hip strategy) was not observed. Although postural control strategies have previously been defined by how the body moves, we conclude that joint torques, which indicate how body movements are produced, are useful in defining postural control strategies. These results also illustrate how the biomechanics of the body can transform discrete control patterns into a continuum of postural corrections.

Adult↗

Torque pull-out load characteristics of skull traction calipers of 'ice tong' type (with particular reference to the cone type of caliper). Technical note.

The relationship between the applied torque and the load required to pull off the calipers was established on cadavers. Torques of 0.7, 0.8 and 0.9 Newton-metres were applied to a specially adapted skull traction caliper whilst attached to the skull in the normal position when used clinically, and loads were applied in steps of 2 lb up to a maximum of 60 lb. It was found that at a torque of 0.7 Newton-metres, the caliper became detached at the maximum load, but still held during traction at torques above this value. Taking into account the fall in load with time that may occur clinically and the limiting torque for pull-out with a traction load of 60 lb, it is suggested for complete clinical safety that a torque of 0.9 Newton-metres should be applied. An open-ended torque release spanner was set for 0.9 Newton-metres for clinical use such that the compressive load is sufficient to prevent pull-out even in the unlikely event of 60 lb being applied for up to 21 days.

Humans↗

The effect of dynamic constant external resistance training on the isokinetic torque-velocity curve.

The purpose of this investigation was to examine the effects of dynamic constant external resistance (DCER) training on isokinetic peak torque (PT) and constant joint angle (CJA) torque at velocities from zero to 5.03 rad.s-1. Twelve male volunteers (mean age +/- SD = 21.67 +/- 2.61 years) were trained using DCER exercises (leg extension and squat) for eight weeks. The subjects were tested pre-training and post-training for isokinetic leg extension PT as well as CJA torque (0.26 rad below the horizontal plane) at multiple velocities (zero, 0.84, 1.68, 2.51, 3.35, 4.19 and 5.03 rad.s-1) using a Cybex II isokinetic dynamometer. Film analyses indicated that the mean training velocities for leg extension and squat were 1.50 +/- 0.20 rad.s-1 and 1.41 +/- 0.26 rad.s-1, respectively. Univariate two-way repeated measures (within subjects design) ANOVAs [time (pre-training versus posttraining) x velocity] were used to analyze the changes in PT and CJA torque across the training period. There was no interaction but a significant (p < 0.05) main effect for time for the CJA torque measures. The increases in CJA torque ranged from 6.3 to 10.7%. For PT, there was a significant (p < 0.05) time x velocity interaction. Post-hoc comparisons showed that PT increased significantly at all velocities (p < 0.0083) as a result of the training (8.1 to 10.9%). The results of this investigation indicated that DCER training significantly increased isokinetic PT and CJA torque values at velocities up to 3.53 rad.s-1 above the training velocity.

Adult↗

Effect of 12 and 20 weeks of resistance training on lumbar extension torque production.

This study compared the effect of varied training frequencies on the development of isometric lumbar extension torque (strength) over 12- and 20-week training periods. Fifty-six subjects were randomly assigned to training once every other week (training group 1, n = 10), once per week (training group 2, n = 12), twice per week (training group 3, n = 12), or three times per week (training group 4, n = 7) or to a nontraining control group (n = 15). Training consisted of one set of 8 to 12 variable-resistance lumbar extensions to volitional muscular fatigue. Prior to and following 12 and 20 weeks of training, subjects were given a test that evaluated isolated isometric lumbar extension torque in a seated position at seven positions (angles) through a 72-degree range of motion. The control group showed no change in isometric torque. All training groups showed significant increases in lumbar extension torque at 12 and 20 weeks of training, whereas no significant differences were found among the groups with respect to the magnitude of torque gained. Pooled training showed a significant time x angle interaction at 12 weeks and a continuing trend at 20 weeks, indicating that the shape of the isometric torque-angle curve changed as a result of training. This effect was due to greater increases in isometric torque at the fully extended position than at the fully flexed position at 12 weeks (92% versus 16%, respectively) and at 20 weeks (123% versus 17%, respectively).(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Correlations between screw hole preparation, torque of insertion, and pullout strength for spinal screws.

The bone-screw interface is critical in the use of spinal instrumentation. The purpose of these experiments described here was twofold. First, to determine whether a correlation existed between torque generated during screw insertion and the pullout strength. Second, to determine how differing surgical methods of screw hole preparation influenced torque of insertion and screw pullout strength. A series of experiments were carried out in which screws were inserted into synthetic bone (experiment 1) and into calf vertebrae (experiment 2). The method of screw hole preparation (i.e., diameter of entrance hole and pilot hole) was varied while the resulting torque of insertion and the pullout strength of the screw was measured in each case. A torque screwdriver was used to measure the torque of insertion of the screws. Screw pullout strength was measured using a materials testing machine. Two important results emerged from these experiments. First, a higher torque of insertion correlated with a higher screw pullout force. This correlation may be useful intraoperatively in evaluating fixation. Second, torque of insertion and pullout force were more influenced by cortex over-drill diameter than pilot hole diameter. These experiments show the importance of the dorsal cortex in pedicle screw fixation.

Animals↗

Torque behavior in schizophrenics, elderly persons, and other special groups.

The incidence of clockwise drawing (torque) was compared for 90 schizophrenics, 57 institutionalized elderly individuals, 24 general medical patients, 50 adults functioning well in the community, 71 graduate students, and 72 undergraduate college students. In view of Blau's (Blau, T. H. Torque and schizophrenic vulnerability: As the world turns. Am. Psychol., 32: 997-1005, 1977) previous finding that torque behavior in children is substantially associated with subsequent schizophrenic pathology an attempt was made to verify the pathological significance of torque. Results indicated that for all Ss combined 38 per cent produced torque. Only the elderly as compared with undergraduate students displayed a significant difference in incidence of clockwise circling, with the elderly producing more torque. The absence of any significant differences between adult schizophrenics and any of the other comparison groups raises serious doubts about the pathological importance of torque in adults. Moreover, since groups who would be expected to be neurologically more intact performed no different from those who would be expected to be less intact, the neural integrative defect explanation proposed by Blau would seem to be unwarranted.

Adult↗

Trunk extensor EMG-torque relationship.

The integrated surface electromyogram (IEMG) of the lumbar erectores spinae and the torque generated were simultaneously recorded from 27 healthy subjects in the standing posture while they pulled isometrically against resistance provided by a harness around the shoulders. The IEMG-torque ratio (efficiency of electrical activity, or EEA) was used to characterize each subject. Individual recordings showed evidence of nonlinearity of the IEMG-torque relationship in that a statistically better fit to experimental recordings was obtained by using two straight lines with a breakpoint between them. However, with repeated testing, the gradients of these two lines were more variable than the slope of the single straight line fitted to the entire recording. The slope of the best fit line (EEA) was less for recordings made during torque decrease than for increasing-torque recordings. This also showed as a "hysteresis" pattern in the recordings. The coefficient of variability (within subjects) of the EEA was greater in day-to-day testing (24%) than with repeated pulls at the same testing session (14%). This was similar to variability of the maximum generated torque. About 25% of the variability between subjects was found to be due to anthropometric differences. The residual variability of the relationship would limit the accuracy of IEMG as a measure of muscular effort under changing torque conditions. However, the EEA may be useful for characterizing muscle performance, especially when maximum effort cannot be achieved.

Adult↗

Isometric torque about the knee joint generated by microstimulation of the cat L6 spinal cord.

Isometric torque was generated about the knee joint by microstimulation of the cat L6 spinal cord using a single microelectrode. The torque responses varied with microstimulation location. Appreciable extension torque was generated by microstimulation in ventrolateral locations of the L6 spinal cord. Stimulation parameters (intensity, frequency and pulse-width) also influenced the extension torque. Specific stimulation parameters (100 microA intensity, 40 Hz frequency and 0.20 ms pulse-width) appear best suited for mapping the spinal cord based on knee joint torque responses. Low levels of cocontraction of the extensor and flexor could be achieved when extension torque was produced, but also varied with the stimulation locations. There are locations in the L6 ventral horn where microstimulation could evoke sustained extension for at least 4 min with only a slight change in torque. This study suggests the possibility of restoring lower limb function in patients with spinal cord injury above the lumbar level.

Animals↗

The mean frequency of the EMG of the knee extensors is torque dependent both in the unfatigued and the fatigued states.

The present study aimed to investigate the shift in the mean frequency (fmean) of the power density spectrum of the electromyogram (EMG) during endurance tests of the knee extensors at three different torque levels (70% MVC, 25% MVC and 10% MVC). Fourteen clinically healthy men performed the three endurance tests until exhaustion. Surface EMG(s) were obtained from the vastus medialis, the vastus lateralis and the rectus femoris muscles and fmean and signal amplitude (RMS) were computed. The subjects rated the perception of fatigue in the knee extensor muscles throughout the three tests using a 10-point graded scale. The endurance time increased with decreasing torque level. A significantly higher perception of fatigue was found at the low torque level (10% MVC) at the end of the endurance time. Individual time series analysis (using linear regression) was performed. It was found that linear models described well the behaviour of fmean throughout the endurance time at the three torque levels. When the endurance times were normalized significant differences in the rate of decrease in fmean were found for vastus lateralis and rectus femoris; the rate decreased with decreasing torque level. For these two muscles significant differences were also found in fmean in the unfatigued state; at 10% MVC significantly lower fmean was found. The present study demonstrated both that the unfatigued fmean and the rate of decrease in fmean (with normalized endurance time) were torque dependent. The latter finding, combined with our result that the subjective fatigue was highest at the lowest torque level, calls into question the use of fmean as a valid indicator of peripheral fatigue.

Adult↗

A measurement of torque values obtained with hand-held drivers in a simulated clinical setting.

PURPOSE: This study measures the amount of torque that can be applied to the heads of implant screws via hand-held drivers and wrenches in a simulated clinical setting. MATERIALS AND METHODS: Sixty second-year dental students applied their maximum controlled torque to the head of a hex instrument attached to a Tonichi torque gauge mounted in a mannequin. The torque values obtained were recorded, and a mean torque value was calculated RESULTS; The mean torque value obtained in this study was 11.55 N.cm. CONCLUSIONS: Small-diameter, 10 N.cm gold screws can be adequately tightened with manufacturer-supplied instrumentation. Larger screws requiring more than 10 N.cm of torque cannot be tightened manually using currently available hand-held screwdrivers and hand wrenches.

Dental Implants↗

Contraction specific changes in passive torque in human skeletal muscle.

The present investigation examined passive torque and electromyographic response in human skeletal muscle during passive static stretch within 60 s after maximal repetitive eccentric and concentric contractions. Passive torque (Nm) offered by the hamstring muscle group was measured during passive knee extension in a modified dynamometer in 10 subjects. The distal thigh was elevated to 0.52 rad from horizontal and the backrest was positioned at 1.57 rad. The lever arm moved the leg passively at 0.09 rad s-1 from a starting position of 1.48 rad below horizontal to the final position where it remained stationary for 90 s. Gross electrical activity of the human hamstring muscle group was recorded simultaneously. The effect of concentric or eccentric contraction was tested on separate days. Two stretch manoeuvres with a 45 min hiatus were administered on a control and experimental side. The experimental side performed 40 maximal effort repetitive concentric or eccentric hamstring muscle contractions at 1.05 rad s-1 prior to the second stretch. Passive torque during the 90 s stretch declined 30-35% on the experimental and control side in all stretches, P < 0.001, without a significant effect of prior contraction mode. Passive peak and final torques were lower on the experimental side, P < 0.01 after concentric contractions. Passive peak and final torques remained unchanged after eccentric contractions on the experimental side. The low level EMG response of the hamstring muscle during the stretch was unchanged after maximal repetitive concentric or eccentric contractions. These data demonstrate acute contraction specific alteration in passive torque in human skeletal muscle, which cannot be accounted for by EMG activity. Furthermore, the lack of difference on the control side implies that one 90 s stretch has no effect on passive torque of the muscle 45 min later.

Adult↗

Torque potentiation and myosin light-chain phosphorylation in human muscle following a fatiguing contraction.

Indices of electrically stimulated and maximal voluntary isometric muscle torgue and the phosphate content of myosin phosphorylatable light chains (P light chains) were studied during recovery following a 60-s maximal voluntary isometric contraction (MVC) in 21 human subjects. Analysis of muscle biopsy samples revealed that immediately after the 60-s MVC there were significant decreases in ATP (-15%) and phosphocreatine (-82%), and lactate concentration increased by 17-fold. All indices of muscle torque production were reduced by the 60-s MVC, but the twitch torque and torque at 10 Hz were relatively less reduced compared with the torque at 20 and 50 Hz or a 1-s MVC. Between 3 and 6 min of recovery, twitch torque and torque at 10 Hz stimulation were significantly potentiated, reaching peak values of 125 and 134%, respectively, compared with rest. Phosphate content of the fast and two slow P light chains was significantly increased over rest levels immediately after and 4 min after the 60-s MVC. These results suggest that myosin P light-chain phosphorylation could provide a mechanism to increase human muscle torque under conditions of submaximal contractile element activation following fatigue.

Adenosine Triphosphate↗

Electrically evoked isokinetic plantar flexor torque in males.

The involuntary angle-specific isokinetic plantar flexor torques of seven male subjects aged 18-21 yr were measured using a Cybex II dynamometer (Lumex) modified by the addition of a strain-gauge load cell to improve the dynamic response of the instrument. Supramaximal electrical stimuli were used to evoke a maximal tetanic response from the triceps surae and ensure constant muscle activation at each angular velocity studied. Angle-specific torques were measured over a range (0.5-5.0 rad/s) of preset velocities, torque decreasing in a nonlinear manner with increasing angular velocity. The torque-velocity data was adequately described by an exponential equation of the form: V = a(e-1/b - e-Po/b) where V = velocity (rad/s), P = torque (N.m), Po = isometric torque (N.m), and a and b are constants. The mean intrasubject coefficient of variation of torque over the range of velocities studies was 7.9 +/- 1.88% (SD).

Adult↗

Response to sudden torques about ankle in man. II. Postmyotatic reactions.

1. Torques were applied to dorsiflex or plantarflex the ankle joint of normal human subjects who were instructed to react to them in various ways. Myoelectric and mechanical responses were recorded and analyzed. 2. Myoelectric responses in triceps surae or tibialis anterior with times of onset between 100 and 200 ms are conditional on the instruction to the subject and not the direction of the applied torque. These responses are termed postmyotatic responses. 3. The latency of the postmyotatic response is the same in both triceps surae and tibialis anterior. It is not dependent on the direction of the applied torque and consequently is not a "stretch reflex," but a triggered reaction. 4. When the subject is sitting relaxed and instructed to resist the applied torque, the magnitude of the postmyotatic response is highly and linearly correlated with the rate of ankle rotation produced by the disturbing torque. 5. Tonic contraction of the muscles prior to the onset of a stretching torque has only a slight effect on the magnitude of the postmyotatic response and usually reduces it. Prior tonic contraction of the antagonist also has a modest effect in reducing the response. 6. Instructing the subject to resist a torque as quickly and strongly as possible increases the magnitude of the postmyotatic response but has little effect on its linear functional relationship to the magnitude of the mechanical stimulus. 7. The postmyotatic response, although voluntary and kinesthetically triggered, is not completely equivalent to a visually triggered voluntary reaction because under experimental situations designed to measure simple reaction times and choice reaction times of the postmyotatic response, no significant difference is found between the two conditions. 8. It is suggested that the peripheral afferent pathways of the stretch reflex play a dual role in determining postmyotatic responses. First, they provide to supraspinal centers one early proprioceptive signal about limb perturbation that can trigger a previously selected mode of response. Second, they modulate that response by convergence, at the segmental level, of peripheral afferent signals with the descending command signals.

Electromyography↗

Inhibition of maximal voluntary isokinetic torque production following stretching is velocity-specific.

Recent research has shown that a regimen of stretching provides an acute inhibition of maximal force production by the stretched muscle group. To further characterize this phenomenon, the effect of an acute stretching regimen on maximal isokinetic knee-extension torque at 5 specific movement velocities (1.05, 1.57, 2.62, 3.67, and 4.71 rad x s(-1)) was examined in 10 men and 5 women (22-28 years). Each person's 5 baseline maximal isokinetic knee-extension torques (dominant leg) were measured on a Cybex NORM dynamometer. Following the baseline torque measurements, the participants stretched the dominant quadriceps for 15 minutes using 1 active and 3 passive stretching exercises. Once the stretching exercises were completed, the maximal torque measurements were repeated. Poststretch maximal torque at 1.05 rad x s(-1) was significantly reduced (p < 0.05) from 218 +/- 47 Nm (mean +/- SD) to 199 +/- 49 Nm (7.2% decrease). At 1.57 rad x s(-1), a similar decrease (p < 0.05) was also seen (204 +/- 48 Nm vs. 195 +/- 47 Nm; 4.5% decrease), but at the other velocities (2.62, 3.67, and 4.71 rad x s(-1)), poststretch maximal torque was unaltered (p > 0.05). It appears, therefore, that the deleterious impact of stretching activities on maximal torque production might be limited to movements performed at relatively slow velocities.

Adult↗