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Biomedical subjects

J E Gregory

Publications and source records attributed to J E Gregory.

At least 37 records · Page 2Linked to original sources

An investigation of the Jendrassik manoeuvre.

Several mechanisms were investigated by which the Jendrassik manoeuvre might potentiate tendon jerk and H-reflexes, recorded here as surface EMG from the triceps surae muscle group. If fusimotor activation is involved, then, after muscle conditioning designed to leave spindles in a slack state, interposing a Jendrassik manoeuvre before reflex testing should have increased reflex amplitude by restoring spindle sensitivity; but this was not the case. The Jendrassik manoeuvre failed to increase facilitation of the soleus H-reflex by a quadriceps volley, contrary to expectations if it operates by presynaptic disinhibition. There was no increase in the level of ongoing EMG during a Jendrassik manoeuvre, indicating that it does not operate by direct facilitation of motoneurones. The Jendrassik manoeuvre produced less reflex potentiation when spindles had a high rate of resting discharge. A remaining, untested mechanism is the modulation of oligosynaptic pathways that may contribute to the largely monosynaptic reflex response.

Electromyography↗

Tension changes in the cat soleus muscle following slow stretch or shortening of the contracting muscle.

1. The permanent extra tension after a stretch and the deficit of tension after a shortening in the soleus muscle of the anaesthetised cat were measured using distributed nerve stimulation across five channels. At low rates of stimulation the optimum length for a contraction was several millimetres longer than that when higher rates of stimulation were used, so that movements applied over the same length range could be on the descending limb of the full activation curve but on the ascending limb of the submaximal activation curve. 2. The extra tension after stretch and the depression after shortening were present only near the peak and on the descending limb of the length-tension curve. Effects on final tension of changing the speed and amplitude of stretches or shortenings were found to be small. 3. Statistical analysis showed that variations in the tension excess or deficit due to changing stimulus rate could be entirely attributed to the effect of stimulus rate on the length-tension relation, as when length was expressed relative to optimum for each rate, stimulus rate was no longer a significant determinant of the tension excess or deficit. 4. The extra tension after stretch and the depression after shortening disappeared if stimulation was interrupted and tension briefly fell to zero. 5. These effects were explained in terms of a non-uniform distribution of sarcomere length changes at long muscle lengths. During stretch some sarcomeres are stretched to beyond overlap while others lengthen hardly at all. During shortening some sarcomeres shorten much further than others. 6. These mechanisms have important implications for exercise physiology and sports medicine.

Analysis of Variance↗

The role of muscle receptors in the detection of movements.

This review discusses the role of muscle receptors, in particular, that of muscle spindles, in the detection of movements, both passive and active. Emphasis is placed on the importance of conditioning the muscles acting at a joint before making measurements of thresholds to passive movements, to take into account muscle's thixotropic property. The detection threshold:movement velocity relation is discussed and described for a number of different joints. Implications for muscle spindles are considered from the generalisation that, when expressed in terms of proportion of muscle fascicle length change, detection thresholds are about the same at different joints. It is concluded that the available data supports the view that muscle spindles lie in parallel with only a portion of a muscle fascicle and not the whole fascicle. At the elbow joint, where it has been tested, movement detection threshold is lower during passive movements than during contraction of elbow muscles. Both peripheral mechanisms and mechanisms operating within the central nervous system may be responsible for the rise in threshold. The signalling of movements by spindles during a contraction raises the question of how the central nervous system is able to extract the length signal under such circumstances, given that there is likely to be co-activation of alpha and gamma motoneurones. The evidence for a central subtraction of fusimotor-evoked impulses and some recent experiments relevant to this idea are described. In conclusion, a number of points of uncertainly have been revealed in this area and these should be the subject of future experiments.

Animals↗

The responses of muscle spindles to small, slow movements in passive muscle and during fusimotor activity.

We have previously shown that movement detection thresholds at the human elbow joint were less than a degree of joint rotation in the passive limb but were higher if they were measured while subjects co-contracted elbow muscles [A.K. Wise, J.E. Gregory, U. Proske, J. Physiol., 508 (1998) 325-330]. Here we report observations on the responses of muscle spindles of the soleus muscle of the anaesthetised cat to determine their ability to signal small length changes in the passive muscle and during a contraction, under conditions resembling those of the human experiments. After appropriate conditioning of the muscle to control for history effects, primary endings of muscle spindles showed thresholds to ramp stretch at 20 micrometers s-1 of between less than 5 micrometers and 15 micrometers, which translates to 0.05 degrees -0.15 degrees of human elbow joint rotation. Thresholds were much higher following conditioning to introduce slack in the muscle. Since during a voluntary contraction there is likely to be alpha:gamma co-activation, responses of spindles were also recorded during slow stretches (100 micrometers at 20 micrometers s-1) during static fusimotor stimulation, dynamic fusimotor stimulation, combined fusimotor stimulation and fusimotor plus skeletomotor stimulation. Invariably, responses to passive stretch were larger than during motor stimulation. It is concluded that spindles are sensitive enough to signal fractions of a degree of elbow joint rotation and that the rise in threshold observed during a voluntary contraction may be accounted for by the actions of fusimotor and skeletomotor axons on spindle stretch responses.

Animals↗

Vibration sensitivity of cat muscle spindles at short muscle lengths.

Experiments are described in which the vibration sensitivity was tested for primary and secondary endings of soleus muscle spindles of the anaesthetised cat. The vibratory stimulus was applied longitudinally to the muscle tendon and, after locating the site of the spindle in the muscle, applied transversely to muscle fibres directly overlying the spindle. All measurements were made with the muscle slack, at 20 mm shorter than its maximum physiological length (Lm-20 mm). Spindles were separated into two groups: spontaneous spindles, which maintained background activity at this length, and silent spindles, which did not. Two forms of muscle conditioning were used: hold-long, in which the muscle was stretched by 5 mm, contracted and then returned to the test length, and hold-test, in which a conditioning contraction was given at the test length. After hold-test, most spindles responded to longitudinal vibration; after hold-long, most did not. This difference in responsiveness was attributed to the presence or absence of slack in intrafusal and extrafusal fibres, due to the muscle's thixotropic property. When the vibration was applied transversely, at a site directly overlying the spindle, responses of silent spindles continued to show thixotropic behaviour, whereas responses of spontaneous spindles were almost independent of the form of muscle conditioning. It is proposed that the ability of spontaneous spindles to maintain background activity at short muscle lengths is due to connective tissue or elastic fibre links between the sensory ending and the spindle capsule. The vibratory stimulus, applied transversely, reaches the sensory ending via these strands which, as non-muscle tissue, do not alter their mechanical state as a result of muscle conditioning.

Animals↗

Muscle history, fusimotor activity and the human stretch reflex.

1. The previous history of contraction and length changes of a muscle influences the size of the stretch reflex and H reflex. Here we ask, is this dependence due to changes in mechanical properties of extrafusal fibres, intrafusal fibres of spindles, or both? 2. The soleus muscle of human subjects was conditioned using either a voluntary contraction or a contraction evoked by low-strength electrical stimulation, in the range 0-25 % of maximum. Following conditioning, reflexes were increased by more than twofold above the no-contraction value by a voluntary contraction of 5 % of maximum, or more, but not by electrical stimulation which presumably did not contract the intrafusal fibres of spindles. 3. When the muscle was conditioned with a contraction at a length shorter than the test length, rather than at the test length, a depressing effect on reflexes was attributed to both the burst of impulses generated in spindles when the muscle was stretched back to the test length and to a reduced stretch sensitivity of muscle spindles. 4. The experiments demonstrate the importance of keeping the muscle and its spindles in a defined mechanical state when measuring reflexes. They also point to the powerful facilitating influences of conditioning muscle contractions provided they recruit the intrafusal fibres of spindles.

Adult↗

Sensory receptors in monotremes.

This is a summary of the current knowledge of sensory receptors in skin of the bill of the platypus, Ornithorhynchus anatinus, and the snout of the echidna, Tachyglossus aculeatus. Brief mention is also made of the third living member of the monotremes, the long-nosed echidna, Zaglossus bruijnii. The monotremes are the only group of mammals known to have evolved electroreception. The structures in the skin responsible for the electric sense have been identified as sensory mucous glands with an expanded epidermal portion that is innervated by large-diameter nerve fibres. Afferent recordings have shown that in both platypuses and echidnas the receptors excited by cathodal (negative) pulses and inhibited by anodal (positive) pulses. Estimates give a total of 40,000 mucous sensory glands in the upper and lower bill of the platypus, whereas there are only about 100 in the tip of the echidna snout. Recording of electroreceptor-evoked activity from the brain of the platypus have shown that the largest area dedicated to somatosensory input from the bill, S1, shows alternating rows of mechanosensory and bimodal neurons. The bimodal neurons respond to both electrosensory and mechanical inputs. In skin of the platypus bill and echidna snout, apart from the electroreceptors, there are structures called push rods, which consist of a column of compacted cells that is able to move relatively independently of adjacent regions of skin. At the base of the column are Merkel cell complexes, known to be type I slowly adapting mechanoreceptors, and lamellated corpuscles, probably vibration receptors. It has been speculated that the platypus uses its electric sense to detect the electromyographic activity from moving prey in the water and for obstacle avoidance. Mechanoreceptors signal contact with the prey. For the echidna, a role for the electrosensory system has not yet been established during normal foraging behaviour, although it has been shown that it is able to detect the presence of weak electric fields in water. Perhaps the electric sense is used to detect moving prey in moist soil.

Animals↗

Summation of responses of cat muscle spindles to combined static and dynamic fusimotor stimulation.

This is a study of the process of interaction between the responses of muscle spindles to stimulation of two fusimotor fibres. Combined stimulation of a static and a dynamic fusimotor fibre supplying the same muscle spindle in the soleus muscle of the anaesthetised cat gave a response which was larger than from stimulating each fibre separately, but less than their sum. A similar summation process was observed with pairs of static fusimotor fibres. The mean summation coefficient for the responses to stimulation of 14 pairs of static fusimotor fibres was 0.29 (range 0.14-0.52; S.D. 0.09), while for 42 static:dynamic pairs it was 0.30 (range 0.07-0.89; S.D. 0.20). Mechanisms considered for the summation process were probabilistic mixing of impulse traffic from two or more impulse generators within the terminals of the primary ending of the spindle, the spread of generator current from one encoding site to another and mechanical interactions between contracting intrafusal fibres. In an experiment where single static and dynamic fusimotor fibres were stimulated together, and then stimulation of the static fibre stopped, the size of the continuing dynamic response was larger than when the dynamic fibre had been stimulated alone. This finding suggested some kind of mechanical interaction between the contracting intrafusal fibres and implies that static and dynamic fusimotor effects within a spindle cannot be considered to be entirely independent of one another.

Action Potentials↗

Detection of movements of the human forearm during and after co-contractions of muscles acting at the elbow joint.

1. We report here observations on the effects of muscle contraction history on thresholds for the detection of movements at the elbow joint of human subjects. Detection thresholds were measured in the direction of flexion or extension to movements of the relaxed forearm at a speed of 0.2 deg s-1 with the elbow at 90 deg. 2. As reported previously, thresholds for movements in the direction of extension were lower than in the direction of flexion after a conditioning co-contraction of elbow muscles with the arm flexed by 30 deg from its mid-position (hold-short). After a co-contraction with the arm held extended by 30 deg (hold-long), thresholds were lower in the direction of flexion. 3. Here we have made two additional observations. Thresholds for movements of the passive forearm after a co-contraction at the 90 deg test position (hold-test) were low, both in the direction of flexion and extension. Secondly, when thresholds were measured while subjects were carrying out a co-contraction of forearm muscles (15-20 % maximum voluntary contraction), thresholds were much higher. 4. It is concluded that muscle contraction history is an important factor to consider when making measurements of movement thresholds at the relaxed elbow joint. It is speculated that during an active contraction increases in muscle spindle discharges evoked by fusimotor activity lead to the rise in movement detection threshold.

Adult↗

A comparison of the effects of concentric versus eccentric exercise on force and position sense at the human elbow joint.

It is generally accepted that our sense of limb position and movement is provided, in part, by signals from muscle spindles, while the sense of muscle force derives from signals in tendon organs. Experiments are described here, using human subjects, in which the effects of eccentric and concentric exercise of elbow flexor muscles are compared on the sense of forearm position and the sense of tension in elbow flexors. Subjects were required to compress a preloaded spring with one arm, carrying out a concentric contraction in elbow flexors, then flexors of the other arm released the spring from compression and thereby carried out an eccentric contraction. The force of the spring was adjusted to be 20% maximum voluntary contraction (MVC), and each subject carried out a minimum of 120 contractions. Position sense was measured in blindfolded subjects by placing one forearm at a set angle and asking subjects to match it by positioning the other arm. Over 4 days postexercise, subjects placed the eccentrically exercised arms in a more extended position than the concentrically exercised arm suggesting that they thought the muscle was shorter than it actually was. In a force-matching task, subjects systematically undershot the target 10% MVC with their eccentrically exercised arm. Since it is known that eccentric exercise is associated with damage to muscle fibres, it is postulated that this leads to a disturbance of muscle receptors, the muscle spindles and tendon organs.

Adolescent↗

The influence of muscle spindle discharge on the human H reflex and the monosynaptic reflex in the cat.

1. Experiments were carried out to test the effect of changes in spindle resting discharge on the size of monosynaptic reflexes in the cat and on the H reflex in humans. Resting discharge was altered by contracting the triceps surae muscle at longer (hold-long) or shorter (hold-short) lengths than that at which the reflex was tested. 2. The reflex in the cat was larger after hold-long than after hold-short conditioning, and the difference, after an initial decline, was well maintained. For the human H reflex a similar pattern was observed except that 15 s after muscle conditioning the difference in reflex size had disappeared. 3. Monosynaptic reflex depression immediately after hold-long conditioning, when most of the muscle spindles are silent, was attributed to the high level of spindle discharge during the immediately preceding hold-long period. The time course of this inhibition was too long to be accounted for by presynaptic inhibition. 4. In the cat heteronymous muscle conditioning was used to test whether presynaptic inhibition could be responsible for reflex depression using the synergist muscle pair lateral gastrocnemius-soleus and medial gastrocnemius. Conditioning one of the pair did not affect the reflex in the other, the opposite result to that expected with presynaptic inhibition. A similar experiment in which the triceps H reflex in human subjects was facilitated by a quadriceps volley gave the same result. 5. Thus this study presents evidence that monosynaptic reflexes are depressed by the on-going discharge of muscle spindles in the homonymous muscle, but that this depression does not appear to involve "classical' presynaptic inhibition.

Adult↗

The effects of muscle conditioning on movement detection thresholds at the human forearm.

We have used the muscle history dependence of the sensitivity of muscle spindles to stretch, to provide evidence for their contribution to kinaesthesia, the sense of position and movement. Stretch sensitivity is altered depending on whether or not slack has been introduced in intrafusal fibres [13]. At the human elbow joint detection threshold was measured to passive movements applied at different speeds to the forearm after a conditioning co-contraction of muscles of the upper arm, with the arm held either flexed ('hold short') or extended ('hold long'). Test measurements were made with the elbow joint at 90 degrees. For the three speeds of movement, 2 degrees s-1, 0.2 degree s-1 and 0.02 degree s-1, after 'hold short' conditioning thresholds were lower for movements into extension, after 'hold long' conditioning they were lower for movements into flexion. It is concluded that when muscle conditioning introduces slack in the intrafusal fibres of muscle spindles, this must be taken up by the test movements before they can be detected by the subject. It means that whenever detection thresholds to passive movements are measured at a joint, the contraction history of the muscles acting at that joint must be taken into account.

Adult↗

Studies of mechanoreceptors in skin of the snout of the echidna Tachyglossus aculeatus.

The echidna Tachyglossus aculeatus, together with the platypus, belongs to the monotremes, a group of mammals with a number of reptilian characteristics. A structure unique to the skin of monotremes is the push rod-a compacted column of epidermal cells that is 20 microns wide and 100 microns long with its tip at the skin surface, and that is able to move relatively independently of adjacent tissue. At the base of each push rod is a cluster of encapsulated nerve endings. Push rods are common in skin of the snout and have been postulated to have a mechanosensory function. Experiments were carried out on four anesthetized echidnas with the aim of determining the function of push rods. Recordings made from the infraorbital nerve, which supplies the skin of the upper jaw, yielded responses from a total of 46 afferents. Two were electroreceptors; the others were mechanoreceptors. Within the group of mechanoreceptors with rapidly adapting responses, three responded to high-frequency vibration and resembled pacinian corpuscles. There were 26 slowly adapting (SA) mechanoreceptors, which, based on the regularity of their discharge, could be divided into two groups: SA I or Merkel type, and SA II or Ruffini type. SA I receptors had very discrete receptive fields with diameters of 100 microns. The receptive fields of two SA I receptors were marked, and after histological processing, one was seen to lie near two push rods. It is concluded that mechanoreceptor responses in the echidna's snout skin resemble those in other mammals in many aspects. We could not unequivocally associate responses to mechanical stimulation with the push rods.

Afferent Pathways↗

The responses of secondary endings of cat soleus muscle spindles to succinyl choline.

This report describes the effects of succinylcholine (SCh) on the secondary endings of cat soleus muscle spindles and attempts to explain them in terms of the action of the drug on intrafusal fibres. All but 2 of 41 secondary endings studied in detail showed a significant response to a single intravenous injection of 200 micrograms kg-1 SCh. This consisted of a rise in the resting rate or development of a resting discharge if the spindle had previously been silent and an increase in the response to stretch. The increases in the responses to stretch were weaker than those observed for primary endings of spindles, but were much larger than those of tendon organs, which showed very little effect with this concentration of drug. The response to SCh showed two features consistent with its action being mediated via an intrafusal muscle fibre contraction rather than a direct depolarising action on the afferent nerve ending. In the presence of SCh, secondary endings were able to maintain a discharge during muscle shortening at rates, on average, more than 5 times greater than under control conditions. Secondly, the increase in spindle discharge produced by SCh showed a length dependence similar to that for fusimotor stimulation. Further support for the action of SCh being principally via an intrafusal fibre contraction was provided by the observation that its effects were abolished by the neuromuscular blocker gallamine triethiodide.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Fusimotor activity and the tendon jerk in the anaesthetised cat.

This is a study of the tendon jerk reflex elicited by a brief stretch applied to the triceps surae muscle group in the chloralose-anaesthetised cat. The size of the recorded reflex depended on stretch parameters (optimum at 300 microns amplitude at a rate of 100 mm/s) and on how the muscle had been conditioned. A reflex elicited after a conditioning contraction at the test length was often twice as large as after a contraction carried out at a length longer than the test length. This difference was attributed to the amount of slack introduced in the intrafusal fibres of muscle spindles by conditioning. The question was posed, did ongoing fusimotor activity exert any influence on the size of the tendon jerk? Depolarization indices (DPI) were calculated from responses of muscle spindles to stretch and correlated with the level of reflex tension. Values of DPI obtained from afferent responses with and without repetitive stimulation of identified fusimotor fibres suggested that with the stretch parameters used here the main influence of fusimotor activity was that it removed any pre-existing slack in muscle spindles and thereby increased reflex tension. In the absence of intrafusal slack, stimulation of static and dynamic fusimotor fibres had little additional influence on the size of the reflex. It is concluded that much of the variability typically seen with tendon jerks is due to muscle history effects. Since in muscles which have not been deliberately conditioned there is commonly some slack present in spindles, activity in fusimotor fibres is likely to reduce slack and therefore increase reflex size.

Anesthesia↗

Blockade of intrafusal neuromuscular junctions of cat muscle spindles with gallamine.

This is a report on the resistance to block of the motor terminals on intrafusal fibres of cat soleus muscle spindles using the drug gallamine triethiodide (Flaxedil). To minimize diffusion barriers and to permit accurate measurements of time courses, rather slow rates of gallamine infusion were used (0.15 mg min-1). The main finding made was that after gallamine infusion, when extrafusal tension had dropped to half, all dynamic fusimotor effects and eight of twenty static effects had fallen to 40% or less of their control value. The remaining static effects persisted at 60-80% of their control value. Where fusimotor fibres were stimulated together with one or two skeletomotor fibres, the influence of the skeletomotor axons was significant only after spindle biasing had fallen to low levels. When gallamine infusion was stopped extrafusal tension returned to control levels within 20-75 min, depending on the length of the block, while fusimotor responses did not fully recover within the recording period of up to 150 min. The combination for some fusimotor responses of an early fall and a late recovery when compared with extrafusal tension, suggested a greater sensitivity of these endings to the drug. A comparison of spindle responses to the drug succinyl choline (SCh) and to fusimotor stimulation in the presence of gallamine showed that SCh responses were rapidly reduced by gallamine and had a long recovery time course, as were some fusimotor responses. From this it is argued that fusimotor effects with a high sensitivity to gallamine blockade were associated with nuclear bag fibre contractions and the more resistant effects with nuclear chain fibre contraction. It is generally believed that intrafusal neuromuscular junctions are more resistant to neuromuscular blockers than extrafusal junctions. The present experiments provide evidence to the contrary for some intrafusal junctions. Since muscle relaxants are often used in general anaesthesia it is interesting to speculate about the recovery of function of proprioceptive reflexes and of kinaesthesia during the immediate post-anaesthetic period, in view of the large difference in recovery time for transmission at intrafusal and extrafusal junctions.

Animals↗

Correlating resting discharge with small signal sensitivity and discharge variability in primary endings of cat soleus muscle spindles.

1. In a previous report we proposed that primary endings of cat soleus muscle spindles can be separated into two kinds. One kind, called by us silent endings, at muscle lengths shorter than Lm -10 (maximum body length -10 mm), fell silent after a 5 mm shortening step. Spontaneous endings, on the other hand, were able to resume a resting discharge after a brief pause at all muscle lengths down to Lm -20. This report examines further differences between the two kinds of endings. 2. There were consistent differences in the muscle length dependence of the maintained level of resting discharge of the two kinds of endings, measured after a conditioning contraction or a contraction followed by a shortening step. The resting discharge of spindles with spontaneous endings, after both forms of conditioning increased progressively with length. For silent endings, after a conditioning contraction, resting discharge fell slightly at longer lengths. 3. Discharge variability, measured at a number of muscle lengths, showed a dependence both on mean interimpulse interval and on spindle type, being higher in silent than spontaneous spindles. 4. Small signal sensitivity was measured with the use of 1 Hz sinusoidal stretches applied longitudinally to the tendon. Sine wave amplitude was adjusted to give a 30% depth of modulation of the resting discharge. Spontaneous endings were consistently less sensitive to the stretches than silent endings at all muscle lengths. Average sensitivities, measured over a range of lengths between Lm -4 and Lm -20 mm were 0.30 imp.s-1.microns -1 for spontaneous endings and 0.66 imp.s-1.microns -1 for silent endings.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗