What if muscle spindles were also involved in the sense of effort?

Effort perception is widely acknowledged to originate from central processes within the brain, mediated by the integration of an efference copy of motor commands in sensory areas. However, in this topical review, we aim to challenge this perspective by presenting evidence from neural mechanisms and empirical studies that suggest that reafferent signals from muscle spindles also play a significant role in effort perception. It is now imperative for future research (a) to investigate the precise mechanisms underlying the interactions between the efference copy and reafferent spindle signals in the generation of effort perception, and (b) to explore the potential for altering spindle sensitivity to affect perceived effort during ecological physical exercise and, subsequently, influence physical activity behaviours.

The contribution of muscle spindles to position sense measured with three different methods.

The sense of limb position is important, because it is believed to contribute to our sense of self-awareness. Muscle spindles, including both primary and secondary endings of spindles, are thought to be the principal position sensors. Passive spindles possess a property called thixotropy which allows their sensitivity to be manipulated. Here, thixotropic patterns of position errors have been studied with three commonly used methods of measurement of position sense. The patterns of errors have been used as indicators of the influence exerted by muscle spindles on a measured value of position sense. In two-arm matching, the blindfolded participant indicates the location of one arm by placement of the other. In one-arm pointing, the participant points to the perceived position of their other, hidden arm. In repositioning, one of the blindfolded participant's arms is placed at a chosen angle and they are asked to remember its position and then, after a delay, reproduce the position. The three methods were studied over the full range of elbow angles between 5° (elbow extension) and 125° (elbow flexion). Different outcomes were achieved with each method; in two-arm matching, position errors were symmetrical about zero and thixotropic influences were large, while in one-arm pointing, errors were biased towards extension. In repositioning, thixotropic effects were small. We conclude that each of the methods of measuring position sense comprises different mixes of peripheral and central influences. This will have to be taken into consideration by the clinician diagnosing disturbances in position sense.

Position sense at the human forearm over a range of elbow angles.

Ten adult participants carried out two experiments on position sense at the forearm, one a two-arm matching task, the other a one-arm pointing task. For matching, both forearms were strapped to paddles which moved in the vertical plane between 0° and 90°. At the start of each trial, the arms were conditioned with a contraction sequence to control for the thixotropic property of muscle and muscle spindles. In the matching task, the blindfolded participant moved their indicator arm from 45° into flexion or extension to match the position of the reference arm placed at one of five test angles, between 5° and 85°. In the pointing task, only the reference arm was strapped to a paddle and conditioned. Participants indicated the position of the arm, hidden by a screen, by moving a pointer paddle or choosing one of a series of trajectory lines drawn on the screen. In matching, where test angles were in the direction of flexion of 45°, errors were small; in the direction of extension larger errors were made, up to 8° into flexion. In pointing trials, except at the most extended position, all errors lay in the direction of extension. It is argued that position sense by matching is concerned with the relative positions of the body and its parts, position sense by pointing gives information about position of the body and limbs in external space.

The neural basis of the senses of effort, force and heaviness.

Effort, force and heaviness are related terms, having in common that they are all sensations associated with the generation of voluntary muscle contractions. Traditionally they have been thought to originate in the brain, as a result of copies of motor commands relayed to sensory areas. A stumbling block for the central hypothesis has been the lack of proportionality between the fall in muscle force from fatigue or paralysis and the increase in sensation generated while trying to achieve the required force. In recent times growing evidence has accumulated supporting a role for peripheral sensory receptors, in particular the muscle spindles, as contributing to these sensations. The review discusses the evidence for participation of sensory receptors and what this means for proprioception. In particular, it is not straightforward to envisage how muscle spindles might provide a reliable force signal in a contracting muscle, with or without support from the fusimotor system. An important additional consideration is the method of measurement. It has emerged that there is evidence of a task-dependency in the composition of the afferent signals contributing to the sense of force. The evidence suggests that the signal used in a two-arm force matching task is not the same as in a one-arm task. It will be important, in the future, to try and obtain more direct evidence about the afferent origins of the senses of effort, force and heaviness, how they might change from one task to another and what implications this has for motor control.

The effects of fast bowling fatigue and adhesive taping on shoulder joint position sense in amateur cricket players in Victoria, Australia.

The impact that muscle fatigue and taping have on proprioception in an applied sporting context remains unclear. The purpose of this study was to investigate disturbances in position sense at the shoulder joint, and asses the effectiveness of adhesive tape in preventing injury and improving performance, after a bout of cricket fast bowling. Among amateur cricket players (N = 14), shoulder position sense, maximum voluntary contraction (MVC) force and bowling accuracy was assessed before and immediately after a fatiguing exercise bout of fast bowling. Participants were tested with the shoulder taped and untapped. Shoulder extension MVC force dropped immediately and 30 min after the exercise (P < 0.05 and P < 0.05, respectively). Position sense errors increased immediately after exercise (P < 0.05), shifting in the direction of shoulder extension for all measurements. Taping had no effect on position errors before exercise, but did significantly reduce position errors after exercise at mid-range shoulder flexion angles (45° and 60°; P < 0.05 and P < 0.05, respectively). Taping had no significant effect on bowling accuracy. These findings may be explained by a body map shift towards a gravity neutral position. Added cutaneous input from the tape is proposed to contribute more to shoulder position sense when muscles are fatigued.

Position sense at the human elbow joint measured by arm matching or pointing.

Position sense at the human elbow joint has traditionally been measured in blindfolded subjects using a forearm matching task. Here we compare position errors in a matching task with errors generated when the subject uses a pointer to indicate the position of a hidden arm. Evidence from muscle vibration during forearm matching supports a role for muscle spindles in position sense. We have recently shown using vibration, as well as muscle conditioning, which takes advantage of muscle's thixotropic property, that position errors generated in a forearm pointing task were not consistent with a role by muscle spindles. In the present study we have used a form of muscle conditioning, where elbow muscles are co-contracted at the test angle, to further explore differences in position sense measured by matching and pointing. For fourteen subjects, in a matching task where the reference arm had elbow flexor and extensor muscles contracted at the test angle and the indicator arm had its flexors conditioned at 90°, matching errors lay in the direction of flexion by 6.2°. After the same conditioning of the reference arm and extension conditioning of the indicator at 0°, matching errors lay in the direction of extension (5.7°). These errors were consistent with predictions based on a role by muscle spindles in determining forearm matching outcomes. In the pointing task subjects moved a pointer to align it with the perceived position of the hidden arm. After conditioning of the reference arm as before, pointing errors all lay in a more extended direction than the actual position of the arm by 2.9°-7.3°, a distribution not consistent with a role by muscle spindles. We propose that in pointing muscle spindles do not play the major role in signalling limb position that they do in matching, but that other sources of sensory input should be given consideration, including afferents from skin and joint.

Muscle thixotropy as a tool in the study of proprioception.

When a muscle relaxes after a contraction, cross-bridges between actin and myosin in sarcomeres detach, but about 1% spontaneously form new, non-force-generating attachments. These bridges give muscle its thixotropic property. They remain in place for long periods if the muscle is left undisturbed and give the muscle a passive stiffness in response to a stretch. They are detached by stretch, but reform at the new length. If the muscle is then shortened, the presence of these bridges prevents muscle fibres from shortening and they fall slack. So, resting muscle can be in one of two states, where it presents in response to a stretch with a high stiffness, if no slack is present, or with a compliant response in the presence of slack. Intrafusal fibres of muscle spindles show thixotropic behaviour. For spindles, after a conditioning contraction, they are left stretch sensitive, with a high level of background discharge. Alternatively, if after the contraction the muscle is shortened, intrafusal fibres fall slack, leaving spindles with a low level of background activity and insensitivity to stretch. Muscle spindles are receptors involved in the senses of human limb position and movement. The technique of muscle conditioning can be used to help understand the contribution of muscle spindles to these senses and how the brain interprets signals arising in spindles. When, in a two-arm position-matching task, elbow muscles of the two arms are deliberately conditioned in opposite ways, the blindfolded subject makes large position errors of which they are unaware. The evidence suggests that the brain is concerned with the difference signal coming from the antagonists acting at the elbow and with the overall difference in signal from the two arms. Another way of measuring position sense is to use a single arm and indicate its perceived position with a pointer. Here, there is no access to a signal from the other limb, and position sense relies on referral to a central map of the body, the postural schema.

Single administration of a psychedelic [(R)-DOI] influences coping strategies to an escapable social stress.

Psychedelic compounds have potentially rapid, long-lasting anxiolytic, antidepressive and anti-inflammatory effects. We investigated whether the psychedelic compound (R)-2,5-dimethoxy-4-iodoamphetamine [(R)-DOI], a selective 5-HT2A receptor partial agonist, decreases stress-related behavior in male mice exposed to repeated social aggression. Additionally, we explored the likelihood that these behavioral changes are related to anti-inflammatory properties of [(R)-DOI]. Animals were subjected to the Stress Alternatives Model (SAM), an escapable social stress paradigm in which animals develop reactive coping strategies - remaining in the SAM arena (Stay) with a social aggressor, or dynamically initiated stress coping strategies that involve utilizing the escape holes (Escape) to avoid aggression. Mice expressing these behavioral phenotypes display behaviors like those in other social aggression models that separate animals into stress-vulnerable (as for Stay) or stress-resilient (as for Escape) groups, which have been shown to have distinct inflammatory responses to social stress. These results show that Stay animals have heightened cytokine gene expression, and both Stay and Escape mice exhibit plasma and neural concentrations of the inflammatory cytokine tumor necrosis factor-α (TNFα) compared to unstressed control mice. Additionally, these results suggest that a single administration of (R)-DOI to Stay animals in low doses, can increase stress coping strategies such as increasing attention to the escape route, promoting escape behavior, and reducing freezing during socially aggressive interaction in the SAM. Lower single doses of (R)-DOI, in addition to shifting behavior to suggest anxiolytic effects, also concomitantly reduce plasma and limbic brain levels of the inflammatory cytokine TNFα.

The contribution of motor commands to position sense differs between elbow and wrist.

Recent studies have suggested that centrally generated motor commands contribute to the perception of position and movement at the wrist, but not at the elbow. Because the wrist and elbow experiments used different methods, this study was designed to resolve the discrepancy. Two methods were used to test both the elbow and wrist (20 subjects each). For the wrist, subjects sat with their right arm strapped to a device that restricted movement to the wrist. Before each test, voluntary contraction of wrist flexor or extensor muscles controlled for muscle spindle thixotropy. After relaxation, the wrist was moved to a test angle. Position was indicated either with a pointer, or by matching with the contralateral wrist, under two conditions: when the reference wrist was relaxed or when its muscles were contracted isometrically (30% maximum). The elbow experiment used the same design to measure position sense in the passive elbow and with elbow muscles contracting (30% maximum). At the wrist when using a pointer, muscle contraction altered significantly the perceived wrist angle in the direction of contraction by 7 deg [3 deg, 12 deg] (mean [95% confidence interval]) with a flexor contraction and 8 deg [4 deg, 12 deg] with an extensor contraction. Similarly, in the wrist matching task, there was a change of 13 deg [9 deg, 16 deg] with a flexor contraction and 4 deg [1 deg, 8 deg] with an extensor contraction. In contrast, contraction of elbow flexors or extensors did not alter significantly the perceived position of the elbow, compared with rest. The contribution of central commands to position sense differs between the elbow and the wrist.

The senses of force and heaviness at the human elbow joint.

The present-day view of the neural basis for the senses of muscle force and heaviness is that they are generated centrally, within the brain, from copies of motor commands. A corollary of the motor discharge generates a sense of effort which underlies these sensations. In recent experiments on force and heaviness sensations using thumb flexor muscles, a rather different explanation has been invoked: Subjects were proposed to rely predominantly on inputs of a peripheral origin, in particular, the signals of muscle spindles. The present experiments have been carried out at the elbow joint to determine whether these new ideas apply more widely. The effects of fatigue of elbow flexor muscles have been studied in force and heaviness matching tasks using three exercise regimes, a sustained maximum voluntary contraction (MVC), a maintained contraction of 35 % MVC, and a maintained contraction of 35 % MVC combined with muscle vibration at 80 Hz. In force-matching experiments, subjects were required to contract both arms and while the reference arm generated the target force under visual control, it was matched by the indicator arm without visual feedback. During the 100 % MVC exercise, force in the exercising reference arm fell rapidly to almost a half of its original value over 90 s while force in the indicator did not fall, leading to a significant overestimation of the reference force. During the 35 % MVC exercise, subjects also overestimated the reference force and this persisted at 5 and 10 min after the exercise. When 35 % MVC was combined with vibration, the amount by which the indicator arm overestimated the reference force was significantly reduced. In heaviness matching experiments, subjects could move their arms through a small range. The reference arm was loaded with a weight, and weights were added or removed from the indicator until heaviness felt the same in the two arms. There was a small, but significant fall in the matching weight used after 100 % MVC exercise, that is, the weight held by the fatigued arm felt lighter. The 35 % exercise did not alter heaviness sensation while 35 % MVC exercise with vibration led to a significant reduction in perceived heaviness. To conclude, while the results of these experiments on elbow flexors are not as clear cut as for thumb flexors, the central effort hypothesis falls short, in a number of respects in explaining the data which are able to be interpreted in terms of a peripheral afferent contribution to the senses of force and heaviness.

Population based case-control study of serious non-fatal motorcycle crashes.

BACKGROUND: Motorcycle sales, registration and use are increasing in many countries. The epidemiological literature on risk factors for motorcycle injury is becoming outdated, due to changes in rider demography, licensing regulations, traffic mix and density, road environments, and motorcycle designs and technologies. Further, the potential contribution of road infrastructure and travel speed has not yet been examined. METHODS/DESIGN: A population based case-control study together with a nested case-crossover study is planned. Cases will be motorcycle riders who are injured but not killed in a motorcycle crash on a public road within 150 km radius of Melbourne, Australia, and admitted to one of the study hospitals. Controls will be motorcycle riders who ride through the crash site on the same type of day (weekday or weekend) within an hour of the crash time. Data on rider, bike, and trip characteristics will be collected from the participants by questionnaire. Data on crash site characteristics will be collected in a structured site inspection, and travel speed for the cases will be estimated from these data. Travel speed for the controls will be measured prior to recruitment with a radar traffic detection device as they ride through the crash site. Control sites for the case-crossover study will be selected 1 km upstream from the crash site and matched on either intersection status or road curvature (either straight or cornered). If the initial site selected does not match the case site on these characteristics, then the closest matching site on the case route will be selected. Conditional multivariate logistic regression models will be used to compare risk between the matched case and control riders and to examine associations between road infrastructure and road environment characteristics and crash occurrence. Interactions between type of site and speed will be tested to determine if site type is an effect modifier of the relationship between speed and crash risk. The relationship between rider factors and travel speed generally will be assessed by multivariate regression methods. DISCUSSION: In the context of the changing motorcycling environment, this study will provide evidence on contemporary risk factors for serious non-fatal motorcycle crashes.

An evaluation of measurement uncertainty of trajectory angles using a 3D laser scanner.

Measuring trajectory angles of bullet defects at a crime scene is an important part of crime scene investigation as these angles can be used for shooting incident reconstructions. The Washington State Patrol Crime Scene Response Team (WSP CSRT) had a desire to report and use measured trajectory angles. To report quantitative measurements, a statement of uncertainty for the measurement must also be reported due to an accreditation requirement through the ANSI National Accreditation Board (ANAB), through which the WSP CSRT is accredited. This evaluation examined the measurement uncertainty of the 3D laser scanning method that the WSP CSRT utilizes for measuring the vertical and azimuth angles from trajectory rods fitted to bullet defects. Three studies were performed which examined the variation of vertical and azimuth angle measurements from trajectory rods on flat surfaces, the variation of azimuth angle measurements from a trajectory rod on a curved surface, and a traceability study with known vertical and azimuth angle measurements. Seven common substrates were selected as target materials and were shot with 9 mm Luger and 0.45 Auto caliber bullets. A vehicle with one bullet defect was utilized for the curved surface study. The WSP's current fleet of Trimble X7 3D laser scanners, the WSP Criminal Investigation Division (CID) Detectives who operate these scanners, and the Forensic Scientists responsible for trajectory rod placement and angle measurements were utilized. An overall measurement uncertainty of +/- 2.6 degrees at an approximate 95% confidence interval was determined for all trajectory angles measured from trajectory rods.

The fall in force after exercise disturbs position sense at the human forearm.

We reported previously that concentric or eccentric exercise can lead to errors in human limb position sense. Our data led us to conclude that the errors, post-exercise, were not due to an altered responsiveness of the proprioceptive afferents, and we proposed that they resulted from central changes in the processing of the afferent input. However, it remained uncertain what was responsible for triggering those changes, the volume of afferent traffic during the exercise or the developing fatigue. The afferent traffic hypothesis was tested by subjects carrying out a series of 250 lightly loaded concentric contractions of elbow flexors that produced little fatigue (6 %). This did not lead to significant position errors. In a second experiment, a series of fatiguing isometric contractions, which kept movements of the muscle to a minimum, led to a 24 % fall in force and significant position errors (3°, direction of extension). In the third experiment, at 24 h after eccentric exercise, when the short-term effects of fatigue and accumulated metabolites were gone, but force was still 28 % below control values, this was accompanied by significant position errors in the direction of extension, 3.2° in the relaxed arm and 3.3° in the self-supported arm. It is concluded that it is the fall in force accompanying exercise which is responsible for disturbing limb position sense. It is suggested that the exercise effects are generated in the brain, perhaps as a result of an alteration of the body map, triggered by the fall in force.

Motorcycle curve assist: A novel approach based on active speed control for crash injury reduction.

OBJECTIVE: Safely negotiating curves with a powered-two-wheeler (PTW) requires a high level of skill, and a significant proportion of PTW crashes have a curve involvement. This study aimed to estimate the applicability, potential benefits and feasibility of novel Motorcycle Curve Assist (MCA). The system is designed to operate an emergency control of the speed of a motorcycle approaching a bend at an inappropriate speed. METHODS: First, the MCA system intervention was defined. Second, the applicability of the system and an estimate of its potential benefits was performed based on a PTW crash database. Motorcyclists' injury risk estimates, MCA working parameters and timing of intervention were employed to estimate the potential injury reduction of applicable crash types. Third, a field test campaign involving 29 common riders as participants was conducted to investigate the real-world applicability and acceptability among end-users of the system deployment in one relevant riding condition adopting a range of parameters of intervention. RESULTS: In the crash database, 23% of cases had curve involvement and after detailed analysis, 14% resulted to be suitable for MCA (60% of cases with curve involvement). The potential relative injury risk reduction considering only the benefits due to crash speed reduction ranged from 3-9% for MAIS2+ to 9-27% for MAIS3+ injuries. Field tests were performed in corners approached at an average speed of 28.7 km/h and an average lean angle of 20°. The system provided a mean deceleration of 0.33 g reached with a fade-in jerk of 1.73 g/s, for an average total duration of 0.59 s. For the field test component, participants reported good controllability of the system, with no incipient loss of control recorded nor reported by participants. CONCLUSIONS: The proposed approach for MCA implementation showed considerable potential benefits in terms of injury reduction. The intervention with the defined working parameters was considered feasible by a sample of end-users. When integrated with an intervention logic capable of predicting emergency situations while approaching curves, MCA will be a technology capable of assisting PTW riders in conditions where other available active safety systems do not.

Improving injury surveillance data quality: a study based on hospitals contributing to the Victorian Emergency Minimum Dataset.

OBJECTIVE: In this paper, we describe the design and baseline data of a study aimed at improving injury surveillance data quality of hospitals contributing to the Victorian Emergency Minimum Dataset (VEMD). METHODS: The sequential study phases include a baseline analysis of data quality, direct engagement and communication with each of the emergency department (ED) hospital sites, collection of survey and interview data and ongoing monitoring. RESULTS: In 2019/20, there were 371,683 injury-related ED presentations recorded in the VEMD. Percentage unspecified, the indicator of (poor) data quality, was lowest for 'body region' (2.7%) and 'injury type' (7.4%), and highest for 'activity when injured' (29.4%). In the latter, contributing hospitals ranged from 3.0-99.9% unspecified. The 'description of event' variable had a mean word count of 10; 16/38 hospitals had a narrative word count of <5. CONCLUSIONS: Baseline hospital injury surveillance data vary vastly in data quality, leaving much room for improvement and justifying intervention as described. IMPLICATIONS FOR PUBLIC HEALTH: Hospital engagement and feedback described in this study is expected to have a marked effect on data quality from 2021 onwards. This will ensure that Victorian injury surveillance data can fulfil their purpose to accurately inform injury prevention policy and practice.

Motorcycle Autonomous Emergency Braking (MAEB) employed as enhanced braking: Estimating the potential for injury reduction using real-world crash modeling.

OBJECTIVE: Recent field-tests on Motorcycle Autonomous Emergency Braking system (MAEB) showed that higher levels of deceleration to improve its effectiveness were feasible. However, the potential of MAEB in mitigating rider injuries is not well understood, particularly in scenarios where the efficacy of standard MAEB is limited because the rider is manually braking. The purpose of this study was first, to assess the injury mitigation potential of MAEB and second, to test MAEB as an enhanced braking system applied in circumstances where the rider is braking before a crash. METHODS: Data from previously investigated motorcycle injury crashes that occurred on public roads in Victoria, Australia were reconstructed using a 2D model. The intervention of MAEB was applied in the simulations to test both MAEB standard and MAEB working as enhanced braking system. The effects of MAEB in mitigating crashes were separated by crash configuration and evaluated based on the modeled reductions in impact speed and injury risk, employing injury risk functions available in the literature. RESULTS: After modeling was applied, MAEB was found to be applicable in 30 cases (91% of those in which was estimated as "possibly applicable"). The modeled Impact Speed Reduction (ISR) among the 30 cases averaged 5.0 km/h. In the cases without manual braking, the mean ISR due to standard MAEB was 7.1 km/h, whereas the relative injury risk reduction ranged from 10% for MAIS2+ to 22% for fatal injuries. In the 14 cases with manual braking, the modeled application of MAEB as enhanced braking led to an average ISR ranging from 5.3 km/h to 7.3 km/h. This resulted in an injury risk reduction ranging from 9% to 12% for MAIS2+ and from 16% to 21% for fatal injuries, depending on the different modes of MAEB. CONCLUSIONS: This study modeled the potential benefits of the highest levels of intervention for MAEB field-tested to date. The findings estimate the degree to which MAEB could mitigate motorcycle crashes and reduce injury risks for motorcyclists. New strategies for MAEB intervention as enhanced braking were modeled through crash simulations, and suggest improvements in the benefits of MAEB when riders are braking before the crash. This highlighted the requirement to perform new field-based tests to assess the feasibility of MAEB deployed as enhanced braking system.

The effect of fatigue from exercise on human limb position sense.

We have previously shown, in a two-limb position-matching task in human subjects, that exercise of elbow flexors of one arm led the forearm to be perceived as more extended, while exercise of knee extensors of one leg led the lower leg to be perceived as more flexed. These findings led us to propose that exercise disturbs position sense because subjects perceive their exercised muscles as longer than they actually are. In order to obtain further support for this hypothesis, in the first experiment reported here, elbow extensors were exercised, with the prediction that the exercised arm would be perceived as more flexed after exercise. The experiment was carried out under three load conditions, with the exercised arm resting on a support, with it supporting its own weight and with it supporting a load of 10% of its voluntary contraction strength. For each condition, the forearm was perceived as more extended, not more flexed, after exercise. This result was confirmed in a second experiment on elbow flexors. Again, under all three conditions the exercised arm was perceived as more extended. To explore the distribution of the phenomenon, in a third experiment finger flexor muscles were exercised. This had no significant effect on position sense at the elbow. In a fourth experiment, position sense at the knee was measured after knee flexors of one leg were exercised and, as for knee extensors, it led subjects to perceive their exercised leg to be more flexed at the knee than it actually was. Putting all the observations together, it is concluded that while the influences responsible for the effects of exercise may have a peripheral origin, their effect on position sense occurs centrally, perhaps at the level of the sensorimotor cortex.

Changes in passive tension after stretch of unexercised and eccentrically exercised human plantarflexor muscles.

The study measured the effect of stretch on passive mechanical properties in unexercised and eccentrically exercised plantarflexor muscles, to obtain insight into how stretch might serve athletes as a warm-up strategy. Passive torque, voluntary contraction strength and muscle soreness were measured before and after a large amplitude stretch given before and after a period of eccentric exercise and at 0, 1, 2 and 24 h later. Stretch of the unexercised muscle led to a 20% fall in passive torque which recovered within an hour. About 40% of the fall could be recovered immediately with a voluntary contraction. After eccentric exercise there was a rise in passive torque by 20% at 2 h post-exercise. This rise was postulated to result from an injury contracture in muscle fibres damaged by the exercise. It was accompanied by a fall in maximum voluntary torque and the development of muscle soreness at 24 h. Stretch of the exercised muscle led to a fall in passive torque and rise in pain threshold. It is proposed that in response to a stretch there is a fall in passive tension in the muscle due to stable cross-bridges in sarcomeres which could be recovered with a voluntary contraction and an additional component attributable to the elastic filament, titin. The size of the fall was not significantly different between exercised and unexercised muscle. These observations provide a physiological basis for the effects of passive stretches on skeletal muscle and help to explain why they are used as a popular warm-up strategy.

Muscle damage produced by isometric contractions in human elbow flexors.

Isometric exercise is often prescribed during rehabilitation from injury to maintain muscle condition and prevent disuse atrophy. However, such exercise can lead to muscle soreness and damage. Here we investigate which parameters of isometric contractions are responsible for the damage. Bouts of 30 repetitions of maximum voluntary contractions of elbow flexors in 38 subjects were carried out and peak force, soreness, and tenderness were measured before the exercise, immediately afterwards, at 2 h, and at 24 h postexercise. When one arm was held near the optimum angle for force generation (90°), the force it produced was greater by 28% than by the other arm held at a longer length (155°). However, despite the smaller contraction forces of the muscle held at the longer length, after the exercise it exhibited a greater fall in force that persisted out to 24 h (20% fall) and more delayed soreness than the muscle exercised at 90° (7% fall at 24 h). The result indicates a length dependence of the damage process for isometric contractions at maximum effort. In four additional experiments, evidence was provided that the damage occurred during the plateau of the contraction and not the rising or relaxation phases. The damage had a prompt onset and was cumulative, continuing for the duration of the contraction. We interpret our findings in terms of the nonuniform lengthening of sarcomeres during the plateau of the contractions and conclude that muscle damage from isometric exercise is minimized if carried out at lengths below the optimum, using half-maximum or smaller contractions. NEW & NOTEWORTHY Isometric exercise, where muscle contracts while the limb is held fixed, is often possible for individuals rehabilitating from injury and can help maintain muscle condition. Such exercise has been reported to cause some muscle damage and soreness. We confirm this and show that to minimize damage, exercising muscles should be held at shorter than the optimum length for force and carried out at half-maximum effort or less.

Eccentric exercise increases EMG amplitude and force fluctuations during submaximal contractions of elbow flexor muscles.

The purpose of this study was to determine the effect of eccentric exercise on the ability to exert steady submaximal forces with muscles that cross the elbow joint. Eight subjects performed two tasks requiring isometric contraction of the right elbow flexors: a maximum voluntary contraction (MVC) and a constant-force task at four submaximal target forces (5, 20, 35, 50% MVC) while electromyography (EMG) was recorded from elbow flexor and extensor muscles. These tasks were performed before, after, and 24 h after a period of eccentric (fatigue and muscle damage) or concentric exercise (fatigue only). MVC force declined after eccentric exercise (45% decline) and remained depressed 24 h later (24%), whereas the reduced force after concentric exercise (22%) fully recovered the following day. EMG amplitude during the submaximal contractions increased in all elbow flexor muscles after eccentric exercise, with the greatest change in the biceps brachii at low forces (3-4 times larger at 5 and 20% MVC) and in the brachialis muscle at moderate forces (2 times larger at 35 and 50% MVC). Eccentric exercise resulted in a twofold increase in coactivation of the triceps brachii muscle during all submaximal contractions. Force fluctuations were larger after eccentric exercise, particularly at low forces (3-4 times larger at 5% MVC, 2 times larger at 50% MVC), with a twofold increase in physiological tremor at 8-12 Hz. These data indicate that eccentric exercise results in impaired motor control and altered neural drive to elbow flexor muscles, particularly at low forces, suggesting altered motor unit activation after eccentric exercise.