Uwe Proske

• PhD, DSc
• Professor Emeritus at Monash University (Australia)

Position sense is arguably more important than any of the other proprioceptive senses, because it provides us with information about the position of our body and limbs in relationship to one another and to our surroundings; it has been considered to contribute to our self‐awareness. There is currently no consensus over the best method of measuring...

Muscle spindles are stretch‐sensitive mechanoreceptors found in the skeletal muscles of most four‐limbed vertebrates. They are unique amongst sensory receptors in the ability to regulate their sensitivity by contraction of the intrafusal muscle fibres on which the sensory endings lie. Muscle spindles have revealed a remarkable diversity of function...

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, thixo...

The senses of limb position and movement become degraded in low gravity. One explanation is a gravity-dependent loss of fusimotor activity. In low gravity, position and movement sense accuracy can be recovered if elastic bands are stretched across the joint. Recent studies using instrumented joysticks have confirmed that aiming and tracking accurac...

In the past, the peripheral sense organs responsible for generating human position sense were thought to be the slowly adapting receptors in joints. More recently, our views have changed and the principal position sensor is now believed to be the muscle spindle. Joint receptors have been relegated to the lesser role of acting as limit detectors whe...

The sense of position of the body and its limbs is a proprioceptive sense. Proprioceptors are concerned with monitoring the body’s own actions. Position sense is important because it is believed to contribute to our self-awareness. This review discusses recent developments in the debate about the sources of peripheral afferent signals contributing...

This is a review of the current state of knowledge of the effects of weightlessness on human proprioception. Two aspects have been highlighted: the sense of limb position and performance in sensorimotor tasks. For the sense of position, an important consideration is that there probably exists more than one sense: one measured in a blindfolded, two-...

New findings: What is the topic of this review? We describe the structure and function of secondary sensory endings of muscle spindles, their reflex action and role in motor control and proprioception. What advances does it highlight? In most mammalian skeletal muscles, secondary endings of spindles are more or much more numerous than primary endi...

This is a review of the current state of knowledge of the effects of weightlessness on human proprioception. Two aspects have been highlighted: the sense of limb position and performance in sensorimotor tasks. For the sense of position, an important consideration is that there probably exists more than one sense: one measured in a blindfolded, two-...

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 f...

This is an account of experiments carried out in my laboratory over more than 20 years, exploring the influence of exercise on human limb position sense. It is known that after intense exercise we are clumsy in the execution of skilled movements. The first question we posed concerned eccentric exercise, where the contracting muscle is forcibly leng...

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...

The kinesthetic senses are the senses of position and movement of the body, senses we are aware of only on introspection. A method used to study kinesthesia is muscle vibration, which engages afferents of muscle spindles to trigger illusions of movement and changed position. When vibrating elbow flexors, it generates sensations of forearm extension...

The kinesthetic senses are the senses of position and movement of the body, senses we are aware of only on introspection. A method used to study kinesthesia is muscle vibration, which engages afferents of muscle spindles to trigger illusions of movement and changed position. When vibrating elbow flexors, it generates sensations of forearm extension...

Isometric exercise is often prescribed during rehabilitation from injury in order 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 contracti...

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 m...

Key points: Position sense at the human forearm can be measured in blindfolded subjects by matching positions of the arms or by a subject pointing to the perceived position of an unseen arm. Effects on position sense tested were: elbow muscle conditioning with a voluntary contraction, muscle vibration, loading the arm and elbow skin stretch. Condi...

These experiments were designed to test the idea that, in a forearm position-matching task, it is the difference in afferent signals coming from the antagonist muscles of the forearm that determines the perceived position of the arm. In one experiment, flexor and then extensor muscles of the reference arm were conditioned by isometric voluntary con...

In this mini-review I have proposed that there are two kinds of position sense, one a sense of the position of one part of the body relative to another, the other a sense of the location in space of our body and its limbs. A common method used to measure position sense is to ask subjects to match with one arm the position adopted by the other. Here...

The traditional view of the neural basis for the sense of muscle force is that it is generated at least in part within the brain. Recently it has been proposed that force sensations do not arise entirely centrally and that there is a contribution from peripheral receptors within the contracting muscle. Evidence comes from experiments on thumb flexo...

This is a brief review of some recent experiments exploring aspects of human limb position sense. It is known that muscle vibration at 80 Hz stimulates muscle spindles to generate a sense of muscle lengthening, representing arm extension for elbow flexors. For the forearm, it was shown in a two-arm position matching task that if the subject was bli...

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 respons...

Most studies of exercise-induced muscle injury have focussed on eccentric contractions, which are known to produce damage and delayed soreness. It is less well known that isometric contractions also produce damage and soreness. This is a study of the damaging effects of isometric exercise. To investigate the contributions of factors such as muscle...

Key points When a blindfolded subject holds his or her arm at a particular angle, its reported position shifts over time; this is known as proprioceptive drift. Here, we show that in relation to position sense at the elbow, the direction of perceived shifts is consistent with adaptation in discharge levels of sensory receptors in elbow muscles. Rai...

Recent studies suggested that centrally-generated motor commands contribute to perception of position and movement at the wrist, but not 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 wri...

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 muscl...

This is a review of the proprioceptive senses generated as a result of our own actions. They include the senses of position and movement of our limbs and trunk, the sense of effort, the sense of force, and the sense of heaviness. Receptors involved in proprioception are located in skin, muscles, and joints. Information about limb position and movem...

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,...

Experiments were carried out on blindfolded human subjects to study the contribution of proprioceptive inputs from both arms in a forearm position matching task. Blindfolded matching accuracy was compared with accuracy when the subject could see their indicator (matching) arm, when they used a dummy arm for matching, and when they looked at a mirro...

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 sub...

This review of kinaesthesia, the senses of limb position and limb movement, has been prompted by recent new observations on the role of motor commands in position sense. They make it necessary to reassess the present-day views of the underlying neural mechanisms. Peripheral receptors which contribute to kinaesthesia are muscle spindles and skin str...

to the editor: Perception of effort during dynamic whole body exercise is a subjective interpretation of an integrative signal, which probably resides from peripheral and central mechanisms. Marcora ([1][1]) clearly provides arguments that the perception of effort during exercise might be

Joint position sense is believed to be mediated by muscle afferent signals. Because a "phantom" hand produced by a sensory and motor nerve block appears to move in the direction of voluntary effort, signals of "motor command" or "effort" can influence perceived joint position. To determine whether this occurs when sensory signals are available, thr...

DefinitionChange in sound energy into some other form, usually heat, in passing through a medium or striking a surface.Acoustics

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 b...

Position matching ability at the forearm in young adults was measured after arm muscles had been placed in a defined mechanical state, called conditioning. With flexion conditioning, elbow flexors were contracted isometrically with the arm held flexed; with extension conditioning, extensors were contracted with the arm held extended. When both arms...

This commentary suggests that the distribution and abundance of muscle spindles in different muscles is related to their role as signallers of muscle fascicle length. Large muscles comprising many fascicles will therefore have more spindles than smaller muscles.

In experiments on position sense at the elbow joint in the horizontal plane, blindfolded subjects were required to match the position of one forearm (reference) by placement of their other arm (indicator). Position errors were measured after conditioning elbow muscles of the reference arm with an isometric contraction while the arm was held either...

This is a report of the effects of exercise on position matching at the knee. Young adult subjects were required to step down a set of stairs (792 steps), representing eccentric-biased exercise of the quadriceps muscle, or step up them, concentric-biased exercise. Immediately after eccentric exercise subjects showed a mean force drop of 28% (+/- 6%...

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...

In a forearm position-matching task in the horizontal plane, when one (reference) arm is conditioned by contraction and length changes, subjects make systematic errors in the placement of their other, indicator arm. Here we describe experiments that demonstrate the importance not just of conditioning the reference arm, but of the indicator arm as w...

Eccentric exercise can produce damage to muscle fibres. Here damage indicators are measured in the medial gastrocnemius muscle of the anaesthetised cat after eccentric contractions on the descending limb of the muscle's length-tension relation, compared with eccentric contractions on the ascending limb and concentric contractions on the descending...

The kinesthetic sense, the sense of position and movement of our limbs, has been the subject of speculation for more than 400 years. The present-day view is that it is signaled principally by muscle spindles, with a subsidiary role played by skin and joint receptors. The problem with muscle spindles as position sensors is that they are able to gene...

When blindfolded subjects match the position of their forearms in the vertical plane they rely on signals coming from the periphery as well as from the central motor command. The command signal provides a positional cue from the accompanying effort sensation required to hold the arm against gravity. Here we have asked, does a centrally generated ef...

This is a study of the ability of blindfolded human subjects to match the position of their forearms before and after eccentric exercise. The hypothesis tested was that the sense of effort contributed to forearm position sense. The fall in force after the exercise was predicted to alter the relationship between effort and force and thereby induce p...

The role of afferent inflow and efferent outflow (or command) signals in judgements of limb position has been debated for over a century. One way to assess this is to check for changes during complete paralysis, with the current view being that perceived movements or position changes do not usually accompany attempts to contract paralysed muscles....

We have recently shown that in an unsupported forearm-matching task blindfolded human subjects are able to achieve an accuracy of 2-3 degrees . If one arm was exercised to produce significant fatigue and the matching task was repeated, it led subjects to make position-matching errors. Here that result is confirmed using fatigue from a simple weight...

Experiments were carried out to test the hypothesis that, in the absence of vision, position sense at the human forearm is generated by the combined input from muscle spindles in elbow flexor muscles and signals of central origin giving rise to a sense of effort. In a forearm position-matching task, to remove a possible contribution from the sense...

A commonly used method for warm-up before exercise is to stretch muscles. How this benefits performance remains uncertain. After a period of eccentric exercise, there is muscle damage accompanied by an increase in passive tension, perceived as a sensation of increased stiffness in the exercised muscles. We have tested the idea that warm-up stretche...

The role of muscle afferents is discussed in terms of their contribution to kinesthesia, the senses of position and movement of the limbs. It is argued that muscle spindles are not well suited as position sensors, on several grounds. Yet we know from muscle vibration experiments that they do contribute to kinesthesia. A number of recent experiments...

Evidence is provided for a mechanical event as the first step in the process leading to muscle damage after a series of eccentric contractions. Aspects discussed include the decline in active tension, increase in passive tension, shift in length-tension relation, soreness, swelling, and disturbed proprioception.

Eccentric exercise is unique in that it can lead to muscle damage and soreness. Concentric exercise is not accompanied by evidence of damage. There are reports in the literature that muscle fatigue is a factor determining the amount of damage from eccentric exercise. Our theory for the damage process predicts that susceptibility for damage is indep...

Nine participants performed two bouts of a step exercise, during which the quadriceps muscle of one leg acted eccentrically. Before and after the exercise, isokinetic torque was measured over a range of knee angles to determine the optimum angle for torque. Immediately after the first bout of exercise, the quadriceps showed a significant (P < 0.05)...

During eccentric exercise contracting muscles are forcibly lengthened, to act as a brake to control motion of the body. A consequence of eccentric exercise is damage to muscle fibres. It has been reported that following the damage there is disturbance to proprioception, in particular, the senses of force and limb position. Force sense was tested in...

Subjecting a muscle to a series of eccentric contractions in which the contracting muscle is lengthened results in a number of changes in its mechanical properties. These include a fall in isometric tension that is particularly pronounced during low-frequency stimulation, a phenomenon known as low-frequency depression (LFD). Reports of LFD have not...

1. Exercise that involves stretching a muscle while active cause microscopic areas of damage, delayed onset muscle soreness and adaptation to withstand subsequent similar exercise. 2. Longer muscle lengths are associated with greater damage and recent animal experiments show that it is the length relative to optimum that determines the damage. 3. I...

1. One common soft-tissue injury in sports involving sprinting and kicking a ball is the hamstring strain. Strain injuries often occur while the contracting muscle is lengthened, an eccentric contraction. We have proposed that the microscopic damage to muscle fibres that routinely occurs after a period of unaccustomed eccentric exercise can lead to...

To investigate the effects of eccentric exercise on the signalling properties of muscle spindles, experiments were done using the medial gastrocnemius muscle of cats anaesthetised with 40 mg/kg sodium pentobarbitone, i.p. Responses were recorded from single afferent nerve fibres in filaments of dorsal root during slow stretch of the passive muscle...

After a period of eccentric exercise of elbow flexor muscles of one arm in young, adult human subjects, muscles became fatigued and damaged. Damage indicators were a fall in force, change in resting elbow angle and delayed onset of soreness. After the exercise, subjects were asked to match the forearm angle of one arm, whose position was set by the...

Eccentric exercise, where the contracting muscle is lengthened, produces microscopic damage in muscle fibers, and sensations of stiffness and soreness, the next day. These normally resolve within a week. A more major sports injury is the muscle strain. Because strain injuries are known to occur during eccentric contractions, it is hypothesized that...

This is a review of the structure and innervation of the mechanosensory organ, the push-rod, in skin of the platypus bill and echidna snout. Four receptor types can. be identified in association with push rods in platypus and echidna: (i) central vesicle chain receptors, (ii) peripheral vesicle chain receptors, (iii) Merkel endings and (iv) pacinif...

In the platypus, electroreceptors are located in rostro-caudal rows in skin of the bill, while mechanoreceptors are uniformly distributed across the bill. The electrosensory area of the cerebral cortex is contained within the tactile somatosensory area, and some cortical cells receive input from both electroreceptors and mechanoreceptors, suggestin...

In human subjects the triceps surae of one leg was exercised eccentrically by asking subjects to walk backwards on an inclined treadmill. Before the exercise controlled local pressure, applied to the muscle with an electromagnet, produced mild soreness, which was reduced when the pressure was combined with vibration. When delayed-onset muscle soren...

This is a report of experiments carried out on the medial gastrocnemius muscle of the anesthetized cat, investigating the effects of eccentric contractions carried out at different muscle lengths on the passive and active length-tension relationships. In one series of experiments, the motor supply to the muscle was divided into three approximately...

1. Experiments were performed to test the ability of human subjects to match forces in their elbow flexor muscles following eccentric exercise of one arm and, in a second series, after biceps brachii of one arm had been made sore by injection of hypertonic saline. 2. In the force-matching task, the elbow flexors of one arm, the reference arm, gener...

Eccentric contractions, where the active muscle is stretched, can lead to muscle damage. One of the signs of damage is a rise in the whole-muscle passive tension. Here we have asked, how many eccentric contractions are necessary to produce a measurable rise in passive tension and can this be detected by the muscle's tension sensors, the tendon orga...

Human subjects generated a specified level of isometric torque with elbow flexor muscles of one arm, the reference arm, under visual feedback. They were then asked to generate what they perceived to be the same level, with the other arm, the indicator, but with no visual feedback. A number of torque levels, between 2% and 30% of maximum were used i...

These experiments are concerned with the ability of human subjects to match isometric torque in their elbow flexor muscles when biceps of one arm is made sore. Pain was induced by injection of hypertonic saline. Subjects were asked to generate a level of torque, 30% of maximum, with one arm, the reference arm. To achieve the required torque, subjec...

To test the possible role of ATP in transducing or modulating touch sensation, an isolated skin-nerve preparation from the toad, Bufo marinus, perfused on the inner side, was used to examine the effects of ATP on slowly adapting (SA) and rapidly adapting (RA) mechanoreceptors, identified by ramp and hold indentation of the skin. ATP (1-50 mM) did n...

Slow-twitch motor units in the medial gastrocnemius muscle of the anesthetized cat were found to have an average optimum length for active tension that was 0.8 +/- 0.5 (SE) mm longer than the whole muscle optimum. For fast-twitch units (time to peak < 50 ms), the average optimum was 1.3 +/- 0.3 mm shorter than the whole muscle optimum. After the mu...

The length-tension curve of muscle is one of the important descriptors of mechanical performance, and also a direct reflection of the underlying structure, particularly the number of sarcomeres connected in series in muscle fibres. This number is one of the most plastic properties of muscle, changing within days after changes in activity patterns....

Some important issues for muscle receptors remain unresolved. For muscle spindles it is uncertain how responses to combined static and dynamic fusimotor stimulation may summate. Such summation may occur during certain phases of locomotion. Two mechanisms considered here include electrotonic spread of generator current between sources of impulse act...

To investigate the possibility of a peripheral contribution to the perturbations of force sensation reported to occur after eccentric exercise, responses to passive and active tension were recorded from Golgi tendon organs in the medial gastrocnemius muscle of the anaesthetised cat, before and after a series of eccentric contractions. After the ecc...

In eccentric exercise the contracting muscle is forcibly lengthened; in concentric exercise it shortens. While concentric contractions initiate movements, eccentric contractions slow or stop them. A unique feature of eccentric exercise is that untrained subjects become stiff and sore the day afterwards because of damage to muscle fibres. This revie...

This collection of contributions on the subject of the neural mechanisms of sensorimotor control resulted from a conference held in Cairns, Australia, September 3-6, 2001. While the three of us were attending the International Union of Physiological Sciences (IUPS) Congress in St Petersburg, Russia, in 1997, we discussed the implications of the nex...

1. This is a report on the history dependence of the passive mechanical properties of the medial gastrocnemius muscle of the anaesthetised cat. 2. The muscle was conditioned with an isometric contraction at the test length, or at 3 mm longer than the test length and then returned to the test length, where the level of resting tension was measured,...

Muscles subjected to eccentric exercise, in which the contracting muscle is forcibly lengthened, become sore the next day (delayed onset muscle soreness). In subjects who had their triceps surae of 1 leg exercised eccentrically by walking backwards on an inclined moving treadmill, mapping the muscle 48 hours later with a calibrated probe showed sen...

Tendon jerk and H-reflexes are both potentiated by the Jendrassik manoeuvre, but the mechanism of potentiation remains uncertain. We investigated several possibilities in human subjects. Evidence for fusimotor activation during the Jendrassik manoeuvre was sought by recording the tendon jerk reflex as surface EMG in triceps surae after the muscles...

This is a report of experiments on ankle extensor muscles of human subjects and a parallel series on the medial gastrocnemius of the anaesthetised cat, investigating the origin of the rise in passive tension after a period of eccentric exercise. Subjects exercised their triceps surae of one leg eccentrically by walking backwards on an inclined, for...

Tendon jerk and H-reflexes are both potentiated by the Jendrassik manoeuvre, but the mechanism of potentiation remains uncertain. We investigated several possibilities in human subjects. Evidence for fusimotor activation during the Jendrassik manoeuvre was sought by recording the tendon jerk reflex as surface EMG in triceps surae after the muscles...