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R M Reinking

Publications and source records attributed to R M Reinking.

At least 19 recordsLinked to original sources

Fatigue-related changes in motor unit action potentials of adult cats.

The purpose of this study was to quantify the changes in motor-unit action potentials (MUAP) and force during a standard motor-unit fatigue test. MUAP waveforms were characterized by the measurement of amplitude, duration, area, and shape (as reflected in a coefficient of proportionality). Fatigue-resistant motor units exhibited small, but statistically significant, changes in MUAP amplitude and area during the fatigue test, whereas fatigable motor units displayed variable changes in MUAP amplitude, duration, and area. For all motor-unit types, the coefficient of proportionality did not change, and hence the change in MUAP area was proportional to the combined changes in amplitude and duration. The between- and within-train changes in MUAP were also distinct for the fatigue-resistant and fatigable motor units. Although several mechanisms could be responsible for the changes in the MUAP as the fatigue test proceeded, the dissociation of the time courses for MUAP and force indicated that these MUAP changes were not the principal reason for the decline in force under these conditions.

Action Potentials

The effect of the stimulation pattern on the fatigue of single motor units in adult cats.

1. The main purpose of this study was to examine the effects of two subtly different stimulus patterns on the force developed by fast-twitch, fatiguable motor units in a cat hindlimb muscle during control (pre-fatigue) and fatiguing contractions. 2. The peak force and the force-time integral responses of nineteen high fatigue (FF) and three intermediate fatigue (FI) motor units of the tibialis posterior muscle in five deeply anaesthetized adult cats were measured at selected times during the course of a 360-s fatigue test. 3. The fatigue test involved a pseudo-random alternation of two patterns of stimulation. One pattern (regular) was composed of a train of stimuli with constant interpulse intervals, set at 1.8 x the twitch contraction time of each unit (interval range, 27-51 ms), and delivered for 500 (or 400) ms. For the total (FF + FI) motor-unit sample, the mean (+/- S.D.) stimulation frequency was 26 +/- 4 Hz (range, 19-37 Hz). The other stimulus pattern (optimized) consisted of three initial stimuli with short (10 ms) interpulse intervals, followed by a constant interpulse-interval train that was adjusted (interval range, 29-62 ms; frequency, 23 +/- 5 Hz; frequency range, 16-36 Hz) such that the total train had the same number of pulses, and the same average frequency and duration as the regular train. 4. The stimulus trains were delivered at 1 s-1 for 360 s, using three-train sequences of each pattern, randomly alternating with one another. The response of the third train in each sequence was selected for the force measurements. The force profile obtained from the fatigue test was subsequently decomposed into two profiles: one attributable to regular and one to optimized stimulation. 5. During the initial responses to the fatigue test, the optimized stimulus pattern produced significantly more force than the regular stimulus pattern. For FF units, the mean increase in peak force (141%) was significantly greater than the increase in the force-time integral (59%). 6. All motor units exhibited an initial potentiation of peak force with the regular stimulation pattern, whereas peak force declined monotonically with the optimized pattern. In contrast, the force-time integral potentiated in the first 30 s for both regular and optimized stimulus patterns. 7. Each motor unit maintained an increased force response to optimized stimulation during the fatigue test, with the greatest relative increase occurring about 120 s into the test, well after the potentiation effect had subsided.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Measurement systems calibration: microcomputer implementation.

Measurement systems used in the collection and processing of laboratory data must be calibrated periodically to obtain accurate results. Because calibration factors can change over time or may be reset to optimize measurements for specific tests, care must be taken to assure that calibration factors and data are aligned correctly. Users should be able to process current data or re-process older data using appropriate calibration factors. The alignment of calibration factors and data should occur in a simple, automatic and transparent way. This document describes one approach to calibration procedures and computer programs used to collect, process, document, measure and display laboratory data. The examples are from our neurophysiology laboratory, where investigators study the mammalian spinal cord and peripheral neuromuscular system. Typical calibration problems, some workable solutions, and computer programs (described in pseudocode) are presented.

Calibration

Computer-aided extraction of the features of the EMG of single motor units.

A software-based system is presented for feature extraction of compound, action-potential (EMG) recordings from single motor units. It simplifies and automates the measurement and analysis of several parameters of the action potential: peak-to-peak amplitude, total duration, peak-to-peak duration, and total area. The software is based on a simple algorithm that first finds the baseline (isoelectric line; including a noise level) of each single EMG potential (waveform) and then searches for the minimum and maximum values in the array of data points representing it. The algorithm searches in both directions starting from the minimum and maximum data points (the waveform peaks) to find the beginning and ending points of the waveform. Using the indices (i.e., array-point numbers) of the four data points provided by the algorithm, the desired features are extracted and/or calculated and saved in a standard-format spreadsheet. The algorithm has a potentially widespread usefulness in a broad array of electrophysiological studies.

Action Potentials

Triggering module for waveform digitization.

A full circuit description is provided for a triggering module used to assist a small laboratory computer in digitizing muscle force- and EMG waveforms. During the stimulation of individual motor units using a standard fatigue test, a train of 13 pulses are delivered at a rate of 40 pps either intracellularly to a motor neuron, or extracellularly to functionally isolated single motor axons from among divided ventral-root nerve filaments. Trains are delivered at a rate of 1/s for the duration of the test, which may range from 120 to 3600 s. Both the force and EMG profiles undergo changes during such tests and the quantification of parameters associated with their waveforms are of interest to neurobiologists. The triggering module allows a typical small laboratory computer to capture user-selected waveforms and thereby reduces the programming problems, timing constraints, storage requirements and analysis time associated with obtaining these parameters. The versatile circuit may be easily adapted to solve similar data-acquisition problems. The method was implemented on an Apple Macintosh II computer but can also be applied to other systems equipped with appropriate software and a data-acquisition card.

Animals

Effects of chloralose-urethane anesthesia on single-axon reciprocal Ia IPSPs in the cat.

Reciprocal Ia inhibitory postsynaptic potentials (IPSPs) generated by single afferents have been recorded with signal averaging in unanesthetized ischemic-decapitate cats for comparison with measurements previously obtained from preparations anesthetized with a mixture of chloralose and urethane. The results are similar to those which we obtained recently for single-axon recurrent IPSPs. Together, the studies show that chloralose-urethane anesthesia has a depressant effect on two widely studied circuits in the mammalian spinal cord.

Anesthesia

Association between biochemical and physiological properties in single motor units.

Motor units from the cat tibialis posterior muscle were examined for an association between physiological and biochemical properties. Functionally isolated motor units were categorized on the basis of their physiological properties. This was followed by quantitative microbiochemical analysis of single muscle fibers from each unit, identified in cross sections using the glycogen-depletion method. The activities of malate dehydrogenase and beta-hydroxyacyl-CoA dehydrogenase distinguished between fatigable (type FF) and fatigue-resistant (types FR and S) units. The activities of both lactate dehydrogenase and adenylokinase were higher in fast- than in slow-contracting units. Cluster analyses, based on both physiological and biochemical properties or on biochemical properties alone, produced groupings identical to types FF, FR, and S. The association between physiological and biochemical properties substantiates the idea that biochemically distinct groups of motor units correspond to physiologically identifiable groups.

3-Hydroxyacyl CoA Dehydrogenases

Measurement of axonal conduction velocity in single mammalian motor axons.

In deeply anesthetized cats, determinations of motor-axonal conduction velocity (CV) were made using extracellular potentials recorded from single, functionally isolated motor axons innervating the muscle tibialis posterior. Axons were activated by suprathreshold electrical stimulation at the ventral-root level. Action potentials were recorded with 3 bipolar electrodes located on the muscle nerve at the level of the popliteal fossa. The most proximal and distal of the bipolar muscle-nerve electrodes were 16.4-22.0 mm apart. Estimates were made of CV from ventral root to muscle nerve (conventional CV) and between the proximal and the distal pairs of muscle-nerve electrodes (muscle-nerve CV). An evaluation was based on comparison of these CVs, estimates of uncertainties in time and distance measurements and simulations of the effects of recording conditions on CV estimates. The analysis indicated that the uncertainty in the conventional CV measurement of mammalian motor axons is at least +/- 2%. However, variability may be as great as 20% between muscle-nerve CV measurements from different experiments, probably due to such factors as regional variation in CV and differences in recording configuration.

Action Potentials

Uniformity of metabolic enzymes within individual motor units.

Individual muscle fibers of 10 motor units from the tibialis posterior muscle of cat were identified by glycogen depletion techniques, characterized for histochemical type, diameter, and intramuscular locations, and analyzed by quantitative biochemical methods. Four enzymes, representing different energy-yielding pathways, were quantitatively assayed in muscle fibers belonging to motor units selected from each of the three major physiological types. All four enzymes demonstrated identical activities among fibers within a motor unit, while showing up to 11-fold differences among fibers belonging to different motor units. Moreover, fibers within a single motor unit, but of substantially different diameters, were nevertheless homogeneous in specific enzyme activities.

3-Hydroxyacyl CoA Dehydrogenases

Detection of synchrony in the discharge of a population of neurons. I. Development of a synchronization index.

A test for synchronization among the spike trains of muscle afferents or motor units is described which utilizes averages of neurograms and rectified neurograms. Synchronization is quantified by the increase of a synchronization index Is above a theoretical value for asynchrony. The dependence of the Is on signal amplitude and certain experimental conditions and a method of estimating confidence limits for the test are presented.

Afferent Pathways

Detection of synchrony in the discharge of a population of neurons. II. Implementation and sensitivity of a synchronization index.

This report describes the use of a synchronization index (Is; Hamm et al., 1985a) and its sensitivity to various forms and degrees of synchrony between spike trains. The dependence of the Is on signal-to-noise ratio, the number of synchronized spike trains and their degree of synchrony is shown in analog and digital simulations. These simulations and a comparison with peristimulus time histograms under conditions of induced synchrony reveal that the Is is a sensitive measure of synchronization in a population of spike trains.

Afferent Pathways

Synchronous afferent discharge from a passive muscle of the cat: significance for interpreting spike-triggered averages.

Evidence is presented for the existence of synchrony between the spike trains of muscle afferents of the passive cat medial gastrocnemius muscle held at fixed length. Synchrony between the spike trains of a population of muscle afferents was quantified by means of a synchronization index (Is), derived from spike-triggered averages of the muscle-nerve neurogram and the rectified neurogram. A previously used test based solely upon the neurogram average (Watt, Stauffer, Taylor, Reinking & Stuart, 1976) is shown to be invalid. The differences between experimentally derived Is values and theoretical Is values derived for the condition of asynchrony were compared to estimated confidence limits for those differences. This comparison revealed that twenty-two of fifty-three muscle-afferent spike trains whose rectified averages satisfied certain conditions for interpreting the Is were synchronized with the discharge of other afferents. The form of the rectified averages of another eight afferents suggested that these afferents might also have been synchronized. Synchrony in the discharge of muscle afferents was found in three experiments in which the neurogram was recorded from a single nerve branch to medial gastrocnemius, as well as in the data of experiments in which the whole muscle nerve was used. The degree of synchrony was similar for Ia, spindle group II and Ib afferents. The magnitude of the synchrony found in these experiments was judged by comparison to the results of analog simulations and the increase in Is values resulting from the application of small, quick stretches to the medial gastrocnemius muscle. The degree of synchrony found on average was approximately equivalent to that of a single spike occurring once for every four discharges of the reference spike train. Simulations were performed to determine the distortion of monosynaptic excitatory post-synaptic potentials (e.p.s.p.s) obtained by spike-triggered averaging which would be produced by synchrony between the spike trains of Ia and spindle group II afferents of the magnitude found in this study. These simulations indicate that the apparent amplitude would be increased by approximately 4 microV on average. Both the 10-90% rise time and the half-width would increase, the effects being greater for smaller e.p.s.p.s. Consequently, the synchrony found in this study is of most concern in the study of small post-synaptic potentials, such as those due to spindle group II afferents.

Action Potentials

Analysis of muscle receptor connections by spike-triggered averaging. 1. Spindle primary and tendon organ afferents.

1. The synaptic connections of 44 single identified muscle spindle Ia afferents and of 21 Golgi tendon organ (Ib) afferents from medial gastrocnemius (MG) were studied in 46 cats by the spike-triggered averaging of synaptic noise in 803 motoneurons of various types. 2. The well-known monosynaptic Ia excitatory connections were confirmed and their characteristics examined in 113 cells. The method was used at greater sensitivity than before and revealed that, in addition to the larger EPSPs of the order of 300 muV, there were many below the previously reported lower 17-muV limit. 3. By studying the Ia disynaptic inhibitory pathway with quick stretch-evoked Ia volleys and by spike-triggered averaging (STA), it was shown that the latter method can reveal disynaptic and possibly trisynaptic excitatory connections. This is believed to depend on having continuous activity in the relevant interneurons. 4. Latencies of individual connections showed broad distributions and arguments are advanced for setting working limits to mono- and disynaptic paths for Ia excitation and inhibition. Monosynaptic EPSP latency from cord entry was 0.4-1.1 ms and disynaptic inhibition was 1.2-2.4 ms. It was recognized that the boundaries are not rigid and monosynaptic Ia EPSPs may have latencies up to 1.5 ms. 5. Rise times of disynaptic PSPs were, on average, significantly longer than monosynaptic, but individual disynaptic responses could have values within the monosynaptic range. 6. A small diphasic wave shortly preceding the monosynaptic EPSPs was interpreted as a presynaptic spike. Its timing was consistent with this and, as such, permitted estimates to be made of central conduction time. 7. An early negative wave (latency less than or equal 1.1 ms) of small emplitude was sometimes detected in antagonist motoneurons when triggering from Ia afferents. It was found tha extracellular fields could be detected due to single Ia afferent excitations and efforts were made to see if the early negative wave could be explained by this. In a few cases there was evidence that a very short-latency IPSP might be occuring. This evidence and its implications are discussed with attention to the new factors which have to be considered in using the spike-triggered averaging method at very high sensitivity. 8. Ib effect were di- or trisynaptic. They were excitatory to 18% of synergists and to 28% of antagonists. They were inhibitory to 41% of synergists and to 19% of antagonists. The Ib IPSPs were larger than the EPSPs.

Animals

Analysis of muscle receptor connections by spike-triggered averaging. 2. Spindle group II afferents.

1. The spike-triggered averaging (STA) method has been used to study synaptic connections of nine spindle group II afferents from medial gastrocnemius to 151 motoneurons of leg muscles in the cat. 2. EPSPs were found in 40 cells, predominantly of triceps surae with latency from cord entry ranging from 0.3 to 4.2 ms. Those with latency less than or equal to 1.4 ms were deduced to be monosynaptic in confirmation of Kirkwood and Sears (13). Mean amplitude for MG-LGS cells was 30.1 muV and mean rise time 1.0 ms (compared with 65.4 muV and 1.0 ms for monosynaptic Ia EPSPs from the preceding report (26)). It is argued that monosynaptic latency for spindle group II afferents could be as large as 1.65 ms. 3. The occurrence of a presynaptic spike permitted the division of EPSP latencies into central conduction time and synaptic delay components. Sindle group II central conduction times were significantly longer than those of Ia afferents, while there were no differences in the synaptic delays associated with the two afferent types. 4. EPSPs of longer latency were judged to be di- or trisynaptic. They were smaller and had longer rise times than the monosynaptic effects. Evidence is presented to show that short rise time of an individual PSP does not guarantee that it is monosynaptic. Rise times are different only on a population basis. 5. Inhibitory responses were found with latencies and mean rise times appropriate for di- and trisynaptic connections. Their mean amplitude was 4.6 muV. 6. The distribution of EPSPs and IPSPs was generally consistent with their exerting stretch reflex effects similar to that of Ia afferents and inconsistent with the inclusion of them in the grouping known as "flexor reflex afferents."

Animals

The motor units of cat medial gastrocnemius: problem of their categorisation on the basis of mechanical properties.

The mechanical properties of 126 motor units from medial gastrocnemius muscle have been studied in 12 adult cats. Units with long contraction times (greater than 45 msec) were non fatigable (24 out of 26 units) and small (25 out of 26 units with less than or equal to 0.3% of the parent whole muscle tetanic tension) thus forming a very homogeneous population. In contrast, fast twitch units (contraction time less than or equal to 45 msec) exhibited a very broad range of tetanic tensions and fatigability. Significant correlations were found, however, within the fast contracting population which indicate a tendency for the more fatigable units to develop more tetanic tension and to be faster contracting. These findings are discussed in relation to the problems associated with using the interrelationships between twitch contraction time, tetanic tension and fatigue resistance to classify motor units into subpopulations sharing similar mechanical properties.

Animals

The motor units of cat medial gastrocnemius: electrical and mechanical properties as a function of muscle length.

The effects of changing muscle length on the mechanical properties of 89 motor units from adult cat medial gastrocnemius have been studied in eight experiments. Few differences were found between the effects of length on tetanic tension, twitch tension, twitch-tetanus ratio, twitch contraction time, twitch half relaxation time, rate of force development and electrical activity for fast contracting (twitch contraction time less than or equal to 45 msec) and slowly contracting (greater than 45 msec) units. Those differences that did appear did not persist when these two groups were matched by tetanic tension. It is concluded that the biophysical mechanisms responsible for the changes in mechanical and electrical properties with length must be similar for fast and slow twitch units and not related to potential differences in their muscle fiber type. The effects of changing muscle length on the mechanical properties of the eight whole muscles suggest that changes in force output with length are of minor importance during normal movements as the muscle is found to be electrically active over a relatively narrow range of lengths close to the optimum length for tetanus of the whole muscle. The very shortest muscle lengths at which there is only minimal force development are not used in natural movements, while the declining limb of the length tension curve is at muscle lengths beyond the maximum in situ length.

Animals

The tendon organs of cat medial gastrocnemius: significance of motor unit type and size for the activation of Ib afferents.

1. Histological and histochemical studies suggest that each tendon organ in a mixed mammalian muscle should be particularly responsive to the contraction of a discrete number of motor units (ca. ten to fifteen), each with differing mechanical properties. This report describes physiological experiments that demonstrate this arrangement for the tendon organs of cat medial gastrocnemius. 2. No correlations could be found between the intensity of discharge of a single tendon organ and the contraction strengths of motor units whose contraction excited the receptor. Tendon organs were found to be as responsive to contraction of small slow twitch units as they were to contraction of larger fast twitch units. Taking the data as a whole, the apparent sensitivity of the receptors during motor unit contractions (pps/force recorded at the tendon) was inversely related to the contraction strengths of the motor units. 3. These findings are discussed in relation to recent evidence on the territory of single motor units in medial gastrocnemius and the force producing capabilities of their individual muscle fibres. It is concluded that in general each motor unit, whose contraction excites a given receptor, contributes one muscle fibre to the receptor capsule. Further, it appears that the various excitatory effects of those muscle fibres inserting into a given receptor capsule are not simply related to their relative contraction strengths but also depend on the details of the mechanical coupling between each fibre and the Ib afferent receptor endings. 4. The results of an ensemble analysis show that despite the lack of correlation between the intensity of tendon organ discharge and the force developed at the tendon during contraction of different motor units, a correlation does appear when the responses of several tendon organs and the forces developed by the motor units which excite them are summed progressively. This finding has implications for the recruitment order of motor units in that the profile of the collective Ib response is shown to differ according to whether motor unit forces are summed randomly or in order of increasing contraction strengths.

Action Potentials