Superimposing noise linearizes the responses of primary muscle spindle afferents to sinusoidal muscle stretch

Experiments were conducted in anaesthetized and spinalized cats to measure the extent to which the non-linear response of Ia afferent fibers to sinusoidal muscle stretch as expressed by the peristimulus-time-histograms, PSTHs, can be transformed into a linear one by means of the superposition of random stretch ("mechanical noise"). The gastrocnemius muscles of one hind leg were stretched and the response to sinewave muscle stretch (amplitudes between 0.01 and 4.0 mm, frequencies between 0.1 and 20 Hz) were investigated while band-limited mechanical noise was superimposed on the sinewave stretch. The random stretch upper cut-off frequency was varied between 60 and 300 Hz; the displacements were normally distributed. The noise amplitude sigma, i.e. the standard deviation of the displacement distributions, was varied systematically between 0.002 and 0.4 mm. Mechanical noise was very effective in raising the mean discharge rate. Added to the sinusoidal stretch it prevented the cessation of firing during the release phase of the stretch cycle, or at least reduced the duration of discharge pauses, i.e., a linearization occurred. In general, the larger the noise amplitude, the more the amplitude of the fundamental harmonic component was attenuated and the phase lead reduced. Apart from this rule the particular combination of superimposing small noise (sigma less than 0.02 mm) on small sinewave stretch (A less than 0.02 mm) could enhance the depth of sinusoidal modulation of cycle histograms (compared with responses to pure sinusoids). Linearizing the sinewave response by additional noise allowed the estimation of frequency response characteristics in the otherwise non-linear range of amplitudes (sinewave amplitude 0.5-1.0 mm).(ABSTRACT TRUNCATED AT 250 WORDS)     

Effect of whole-body vibration with different frequencies and intensities on auditory evoked potentials and heart rate in man

Auditory evoked brain potentials (AEP) and electrocardiogram (ECG) were recorded from 9 healthy male subjects during sinusoidal whole-body vibration exposure (WBV) in the longitudinal (+/- az) direction with four frequencies (1 Hz, 2 Hz, 4 Hz, and 8 Hz) and two intensities as well as under non-WBV conditions. The sequences of the different experimental conditions were arranged according to a 9 X 9 Latin Square design. The sound of the electrohydraulic vibrator was masked by a constant noise level. A subtraction technique was used to eliminate vibration-synchronous activity contaminating the electroencephalogram. The AEP amplitude N1-P2 revealed systematic effects of different WBV frequencies and intensities. The amplitude decreased along with an increase in intensity (16 dB) by about 10 per cent. It diminished increasingly with a monotonic trend in the order non-WBV, WBV 8 Hz, WBV 4 Hz, WBV 2 Hz, and WBV 1 Hz. The interbeat-interval histograms computed from the ECG exhibited the highest mean values at MBV of 1 Hz, high intensity, and the lowest ones at WBV of 4 Hz, high intensity. The AEPs are reaffirmed as an informative measure for studying the WBV effect on central nervous information processing, although the modes of action are not yet fully known. Efferent influences on the acoustic input, cross-modality interaction, sensory mismatch, and changes of central nervous activation level are discussed as potential mechanisms.     

Relationship between the efficiency of muscular work during jumping and the energetics of running

The running economy of seventeen athletes was studied during running at a low speed (3.3 m X s-1) on a motor-driven treadmill. The net energetic cost during running expressed in kJ X kg-1 X km-1 was on average 4.06. As expected, a positive relationship was found between the energetic cost and the percentage of fast twitch fibres (r = 0.60, n = 17, p less than 0.01). In addition, the mechanical efficiency during two different series of jumps performed with and without prestretch was measured in thirteen subjects. The effect of prestretch on muscle economy was represented by the ratio between the efficiency of muscular work performed during prestretch jumps and the corresponding value calculated in no prestretch conditions. This ratio demonstrated a statistically significant relationship with energy expenditure during running (r = -0.66, n = 13, P less than 0.01), suggesting that the elastic behaviour of leg extensor muscles is similar in running and jumping if the speeds of muscular contraction during eccentric and concentric work are of similar magnitudes.     

A method for the interpretation of signals in afferent nerves from muscle spindles

A method is proposed for the interpretation of the signals in sensory nerve fibres. It is applicable to systems in which the transfer function between the input to a sense organ and the action potential firing frequency is known. In the present case, the chelonian muscle spindle is considered since its output to ramphold-relase and sinusoidal stretches can be rather accurately simulated.     

Role of knee ligaments in proprioception and regulation of muscle stiffness

Physiologic evidence for the sensory role of the knee joint ligaments are reviewed. The cruciate and collateral ligaments accomodate morphologically different sensory nerve endings with different capabilities of providing the central nervous system (CNS) with information not only about noxious and chemical stimuli but also about mechanical events, e.g., movement- and position-related stretches of the ligaments. Available data show that low-threshold joint/ligament receptor (i.e., mechanoreceptor) afferents evoke only weak and rare effects in skeletomotor neurons (α-motoneurons), whereas they frequently and powerfully influence fusimotor neurons (γ-motoneurons). The effects on the γ-muscle-spindle system in the muscles around the knee are so potent that even stretches of the cruciate ligaments at relatively moderate loads (not noxious) may induce major changes in responses of the muscle spindle afferents. As the activity in the primary muscle spindle afferents modifies stiffness in the muscles, the cruciate ligament receptors may, through the γ-muscle-spindle system, participate in regulation and preparatory adjustment of the stiffness of the muscles around the knee joint and thereby of knee joint stiffness. Thus, the sensory system of the cruciate ligaments is able to contribute significantly to the functional stability of the knee joint. The possible role of (ligamentous) joint receptors in genesis and spread of muscular tension in occupational muscle pain and in chronic musculoskeletal pain syndromes is also discussed.     

Model of the firing frequency of the chelonian muscle spindle

A model is proposed for the frequency of firing of the chelonian muscle spindle in response to mechanical stretches. First an attempt is made to fit the response to a first and second order model and, after incorporating the encoding properties of the spindle, a simulation is obtained which predicts in terms of the instantaneous frequency of firing the response of the spindle to ramp and sinusoidal mechanical stretches.     

On the Transmission of Information through Sensory Neurons

Information about muscle length is transmitted to the cerebellum from muscle spindle receptors through the dorsal spinocerebellar tract (DSCT). The “transinformation” about muscle length in single DSCT fibers was calculated from steady-state spike trains by two different methods, assuming that the decoding mechanisms use a frequency code. By the first method, the number of distinguishable muscle lengths (and thus the transiformation) was determined from the rate of convergence of the mean frequency of firing (with increasing number of intervals). The observation time necessary to estimate the mean frequency of the impulse train with a certain accuracy was independent of the stretch level, even though the number of intervals necessary to make this estimate was different at high and low levels of stretch. By the second method an input frequency-output frequency matrix was calculated. The transinformations and the rate of transinformation was then calculated from this matrix. There was an acceptable agreement in the estimates of transinformation by the two methods. The rates of transinformation are significantly increased by the particular time structure of the discharge patterns of the nerve cells. Consequently, the loss of information due to the synaptic coupling is appreciably reduced.     

The effects of recurrent inhibition on the cross-correlated firing patterns of motoneurones (and their relation to signal transmission in the spinal cord-muscle channel)

Cross-correlation histograms (CCH) were computed for discharge sequences of pairs of motoneurones which were excited by sinusoidal muscle stretches. These CCH's were compared before and after opening of the recurrent inhibitory loop by Renshaw cell blocking agents. Periodic patterns in the CCH's indicative of specifically timed phase relations between discharges of different motoneurones were enhanced after Renshaw cell blockage. This was confirmed by power spectra computed for the CCH's. They contained power peaks about 50Hz which tended to increase after depression of recurrent inhibition. The correlation was thus due predominantly to line current interference which seemed to act as a common entrainment input at the spinal level. It is concluded that Renshaw cells de-correlate discharge patterns of different motoneurones of the same pool by injecting uncorrelated signals into them. This de-correlation is an important prerequisite for distortion suppression of signal transmission in a multi-channel system, like that of stretch reflex, and for its linearization.     

The response of fast and slow nuclear bag fibres and nuclear chain fibres in isolated cat muscle spindles to fusimotor stimulation, and the effect of intrafusal contraction on the sensory endings.

1. The mechanical behaviour of intrafusal muscle fibres during fusimotor stimulation and passive stretch was observed directly in muscle spindles isolated from the cat tenuissimus muscle. 2. Mammalian intrafusal muscle fibres are of three functional types. Most spindles contain one slow nuclear bag fibre, one fast nuclear bag fibre, and four or five nuclear chain fibres. 3. Contraction in slow nuclear bag fibres is characterized by a long latency and very slow initial velocity, whereas the latency for the other intrafusal fibres is short and the inital velocity rapid. The mean time for maximum contraction (at 75 Hz to 100 Hz) and relaxation is significantly longer for slow nuclear bag fibres (0-8s) than for other intrafusal fibres (0-5 s). The contraction time of fast nuclear bag fibres is sometimes longer than that of nuclear chain fibres but the mean values are not significantly different; a difference in the time to attain 90% contraction is more obvious. 4. At low stimulation frequencies (10 Hz) contraction in slow nuclear bag fibres and in most fast nuclear bag fibres is smooth whereas nuclear chain fibres exhibit marked oscillations. Single stimuli elicit small local twitches in nuclear chain fibres and occasionally in fast nuclear bag fibres but produce no visible effect in slow nuclear bag fibres. 5. Maximum contraction of slow and fast nuclear bag fibres at body temperature is attained at a stimulation frequency of 75 Hz to 100 Hz, whereas a frequency of 150 Hz or more is required for maximum contraction of nuclear chain fibres. At 50 Hz at body temperature contraction in nuclear bag fibres is at least half the maximum, whereas in many spindles nuclear chain fibres show only a very small contraction at this frequency. 6. Contraction in slow nuclear bag fibres occurs at one or two discrete foci, most of which lie in the intracapsular region beyond the end of the fluid space. Weak contraction extends the primary sensory spiral by a small amount (2%-8%) at a low velocity (5%-10%s-1). When the fibre is passively stretched the spiral opens and then creeps back to about 75% of the extension at the end of the stretch due to yielding in the poles of fibre; creep is complete in 0-5s to 2-5s. 7. Contraction in fast nuclear bag fibres also occurs at one or two discrete foci, most of which lie in the intracapsular region beyond the end of the fluid space. Shortening of sarcomeres at the foci and extension of the sensory spiral are, however, up to eight times greater (up to 25%) than in slow nuclear bag fibres, and the velocity of stretch of the spiral is three to eight times greater (25%-40%s-1). Fast nuclear bag fibres exhibit little or no creep following passive stretch. 8. Contraction in the nuclear chain fibre bundle is localized to the intracapsular region, centered on a point in the intracapsular region between 0-9 mm and 1-6 mm from the spindle equator. Maximal contraction stretches primary and secondary sensory endings by 15% to 20%, at 30% to 40% s-1...     

The Frequency Response, Coherence, and Information Capacity of Two Neuronal Models

Two neuronal models are analyzed in which subthreshold inputs are integrated either without loss (perfect integrator) or with a decay which follows an exponential time course (leaky integrator). Linear frequency response functions for these models are compared using sinusoids, Poisson-distributed impulses, or gaussian white noise as inputs. The responses of both models show the nonlinear behavior characteristic of a rectifier for sinusoidal inputs of sufficient amplitude. The leaky integrator shows another nonlinearity in which responses become phase locked to cyclic stimuli. Addition of white noise reduces the distortions due to phase locking. Both models also show selective attenuation of high-frequency components with white noise inputs. Input, output, and cross-spectra are computed using inputs having a broad frequency spectrum. Measures of the coherence and information transmission between the input and output of the models are also derived. Steady inputs, which produce a constant “carrier” rate, and intrinsic sources, which produce variability in the discharge of neurons, may either increase or decrease coherence; however, information transmission using inputs with a broad spectrum is generally increased by steady inputs and reduced by intrinsic variability.     

Evaluation and statistical judgement of neural responses to sinusoidal stimulation in cases with superimposed drift and noise

Time histograms of neural responses evoked by sinusoidal stimulation often contain a slow drifting and an irregular noise which disturb Fourier analysis of these responses. Section 2 of this paper evaluates the extent to which a linear drift influences the Fourier analysis, and develops a combined Fourier and linear regression analysis for detecting and correcting for such a linear drift. Usefulness of this correcting method is demonstrated for the time histograms of actual eye movements and Purkinje cell discharges evoked by sinusoidal rotation of rabbits in the horizontal plane. In Sect. 3, the analysis of variance is adopted for estimating the probability of the random occurrence of the response curve extracted by Fourier analysis from noise. This method proved to be useful for avoiding false judgements as to whether the response curve was meaningful, particularly when the response was small relative to the contaminating noise.     

Characterization of muscle spindle afferents in the adult mouse using an in vitro muscle-nerve preparation

We utilized an in vitro adult mouse extensor digitorum longus (EDL) nerve-attached preparation to characterize the responses of muscle spindle afferents to ramp-and-hold stretch and sinusoidal vibratory stimuli. Responses were measured at both room (24°C) and muscle body temperature (34°C). Muscle spindle afferent static firing frequencies increased linearly in response to increasing stretch lengths to accurately encode the magnitude of muscle stretch (tested at 2.5%, 5% and 7.5% of resting length [Lo]). Peak firing frequency increased with ramp speeds (20% Lo/sec, 40% Lo/sec, and 60% Lo/sec). As a population, muscle spindle afferents could entrain 1:1 to sinusoidal vibrations throughout the frequency (10-100 Hz) and amplitude ranges tested (5-100 μm). Most units preferentially entrained to vibration frequencies close to their baseline steady-state firing frequencies. Cooling the muscle to 24°C decreased baseline firing frequency and units correspondingly entrained to slower frequency vibrations. The ramp component of stretch generated dynamic firing responses. These responses and related measures of dynamic sensitivity were not able to categorize units as primary (group Ia) or secondary (group II) even when tested with more extreme length changes (10% Lo). We conclude that the population of spindle afferents combines to encode stretch in a smoothly graded manner over the physiological range of lengths and speeds tested. Overall, spindle afferent response properties were comparable to those seen in other species, supporting subsequent use of the mouse genetic model system for studies on spindle function and dysfunction in an isolated muscle-nerve preparation.     

Electrolocation in the presence of jamming signals: behavior

Electrolocation behavior of Apteronotus leptorhynchus was studied by monitoring the animal's ability to maintain orientation to a variety of moving electrolocation targets. The primary goal of this study was to determine the relative effectiveness of various types of electrical 'jamming signals' in disrupting electrolocation performance. 1. Two measures of electrolocation performance were used: The latency between the electrolocation target motion and the fish's following response, and the minimum distance separating the fish from the target during the target movement sequence. Latency increased and minimum fish-target distance decreased as target size was decreased, and when large diameter ceramic targets were used as control stimuli the fish were less able to avoid, and frequently collided with, these 'electrically transparent' objects. 2. Four types of jamming signals were used in attempts to mask the electrosensory input used for electrolocation. Broad-band noise and sinusoidal signals, different in frequency by a few Hz from the animal's personal electric organ discharge (DF stimuli), were used to jam the tuberous electroreceptors. Five Hz and 50 Hz sinusoidal signals were used to jam the low-frequency or ampullary receptor system. Both the noise and the DF stimuli were effective in reducing electrolocation performance, and the threshold intensity for behavior deterioration was about three-fold lower for DF stimuli as compared to the noise. The rate of change of response deterioration as a function of increasing jamming intensity was, however, not different for these two types of stimuli. Neither the 50 Hz nor the 5 Hz jamming signals caused electrolocation deterioration when presented alone. However, 5 Hz jamming, when added to either the noise or DF jamming, did result in significant increments in response deterioration. This suggests that the ampullary receptors can provide supplementary information for electrolocation. 3. Electrolocation performance deterioration was also studied with various difference frequencies between an animal's EOD and the sinusoidal jamming stimulus. Increasing DF results in decreased electrolocation deterioration, but even at the highest DF frequencies used (128 Hz) significant response degradation was observed. 4. The apparent differences in the effectiveness of noise and DF stimulation in reducing electrolocation performance are largely accounted for by the differential effects of the tuberous electroreceptor filter characteristics on these two types of signals.     

A model-based approach for the analysis of neuronal information transmission in multi-input and -output systems

We present a new method to characterize multi-input and output neuronal systems using information theory. To obtain a lower bound of transinformation we take three steps: (1) Estimation of the deterministic response to isolate components carrying stimulus information. The deviation of the original response from the deterministic estimate is defined as noise. (2) Coordinate transformation using PCA yields an uncorrelated representation. (3) Partial transinformation values are calculated independently either by Shannon's formula assuming normality or based on density estimation for arbitrary distributions. We investigate the performance of the algorithms using simulated data and discuss suitable parameter settings. The approach allows to evaluate the degree to which stimulus features are encoded. Its potential is illustrated by analyses of neuronal activity in cat primary visual cortex evoked by electrical retina stimulation.     

Electrolocation in the presence of jamming signals: electroreceptor physiology

Responses of ampullary and tuberous electroreceptor afferents were studied using moving electrolocation targets and electrical modulations of the animal's electric organ discharge as stimuli. The ability of the electroreceptors to encode these stimuli was measured with and without various forms of electrical jamming signals. The goal of this study was to measure the deterioration in electroreceptor responses due to the jamming signals, and to compare these results with the behavioral measures of electrolocation under the same conditions of jamming as described in the preceding report (Bastian 1987). 1. Three types of jamming stimuli were used to interfere with the tuberous electroreceptor afferents' ability to respond to the test stimuli mentioned above: Broad-band noise, high frequency stimuli consisting of a sinusoidal waveform having a frequency maintained at a chosen difference frequency (DF) from the EOD frequency of the fish being studied, and 5 or 50 Hz sinusoidal stimuli. 2. The tuberous receptor afferents' spontaneous frequency was sensitive to continuous presentation of all but the 5 Hz jamming signals. The 4 Hz DF signal caused the largest increase in spontaneous activity, the 50 Hz stimulus was intermediate in effectiveness, and the noise stimulus caused the smallest increase. Estimates of the variability of the ongoing receptor activity were also made, and both the 4 Hz DF and the 50 Hz stimuli reduced the coefficient of variation of the receptor activity, but noise had no significant effect on this parameter. Noise, 4 Hz DF, and 50 Hz jamming signals also reduced the tuberous receptors' responses to a 100 ms EOD amplitude modulation, and the 5 Hz stimulus was again ineffective. 3. Noise and 4 Hz DF jamming were also effective in reducing tuberous receptor afferents' responses to a moving metal electrolocation target. The 4 Hz DF stimulus was most effective in reducing the receptor's ability to encode information about the target. Receptor responses showed about a three-fold larger decrease per 10 dB increase in DF jamming amplitude as compared to similar sized increases in noise amplitude. Threshold target distances were also determined with and without noise and DF jamming, and again, the noise stimulus was less effective in reducing the distance at which electrolocation targets were just detectable. 4. Recordings from ampullary receptor afferents confirmed that the galvanic potentials produced by metal electrolocation targets stimulate these receptors while EOD distortions caused by such objects probably do not.(ABSTRACT TRUNCATED AT 400 WORDS)     

Motor responses ofHalobacterium halobrium to sinusodial light stimuli

Sinusoidal light stimuli have been used to elicit photoresponses inHalobacterium halobium at various periods of the stimulus. Cycle histograms and histograms of the times between consecutive reversals are used to illustrate the responses, and the method used to obtain sinusoidally modulated intense light (with periods up to 1 s) is described.     

Measuring myosin cross-bridge attachment time in activated muscle fibers using stochastic vs. sinusoidal length perturbation analysis

The average time myosin cross bridges remain bound to actin (t(on)) can be measured by sinusoidal length perturbations (sinusoidal analysis) of striated muscle fibers using recently developed analytic methods. This approach allows measurements of t(on) in preparations possessing a physiologically relevant myofilament lattice. In this study, we developed an approach to measure t(on) in 5-10% of the time required for sinusoidal analysis by using stochastic length perturbations (white noise analysis). To compare these methods, we measured the influence of MgATP concentration ([MgATP]) on t(on) in demembranated myocardial strips from mice, sampling muscle behavior from 0.125 to 200 Hz with a 20-s burst of white noise vs. a 300-s series of sinusoids. Both methods detected a similar >300% increase in t(on) as [MgATP] decreased from 5 to 0.25 mM, differing by only 3-14% at any [MgATP]. Additional experiments with Drosophila indirect flight muscle fibers demonstrated that faster cross-bridge cycling kinetics permit further reducing of the perturbation time required to measure t(on). This reduced sampling time allowed strain-dependent measurements of t(on) in flight muscle fibers by combining 10-s bursts of white noise during periods of linear shortening and lengthening. Analyses revealed longer t(on) values during shortening and shorter t(on) values during lengthening. This asymmetry may provide a mechanism that contributes to oscillatory energy transfer between the flight muscles and thoracic cuticle to power flight. This study demonstrates that white noise analysis can detect underlying molecular processes associated with dynamic muscle contraction comparable to sinusoidal analysis, but in a fraction of the time.     

Fusimotor reflex profiles of individual triceps surae primary muscle spindle afferents assessed with multi-afferent recording technique.

The experiments were performed on 21 cats anaesthetized with alpha-chloralose. The aim of the study was to investigate sets of simultaneously recorded spindle afferents (2-4 in each set) from the triceps surae muscle (GS) with respect to the pattern of fusimotor reflex effects evoked by different types of ipsi- and contralateral reflex stimulation. The afferents' responses to sinusoidal stretching of the GS muscle were determined and the fusimotor reflex effects were assessed by comparing the afferent responses (i.e. the mean rate of firing and the depth of modulation) elicited during reflex stimulation with those evoked in absence of any reflex stimulus. Natural of electrical activations of ipsi- and contralateral muscle, skin and joint receptor afferents were used as reflex stimuli. The spindle afferents were influenced by several modalities and from wide areas, with a majority responding to both ipsi- and contralateral stimuli. A particular reflex stimulus often caused different effects on different afferents, and the various reflex stimuli seldom gave similar effects on a particular afferent. Multivariate analysis revealed that the variation in response profiles among simultaneously recorded afferents were as great as between afferents recorded on different occasions. This suggests that the individualized response prifiles, observed in earlier investigations, represent a very diversified reflex control of the spindle primary afferents, and are not a reflection of changes in the setting of the spinal interneuronal network, occurring during the time interval between the recordings of different units. Also, there was no relation between the conduction velocity of the afferents and the reflex profiles of the afferents, but non-linear relations were found between effects elicited by different types of stimuli. Indications were also found that it may be possible to separate the population of GS muscle spindles into subgroups, according to the fusimotor effects exhibited by activation of various categories of ipsi- and contralateral receptor afferents. It is concluded that one possible way of making the very complex reflex system controlling the muscle spindles intelligible may be a combination of multiple simultaneous recordings of spindle afferents and multivariate analysis.     

Fusimotor reflexes influencing secondary muscle spindle afferents from flexor and extensor muscles in the hind limb of the cat.

The purpose of this study was to investigate secondary muscle spindle afferents from the triceps-plantaris (GS) and posterior biceps and semitendinosus (PBSt) muscles with respect to their fusimotor reflex control from different types of peripheral nerves and receptors. The activity of single secondary muscle spindle afferents was recorded from dissected and cut dorsal root filaments in alpha-chloralose anaesthetized cats. Both single spindle afferents and sets of simultaneously recorded units (2-3) were investigated. The modulation and mean rate of firing of the afferent response to sinusoidal stretching of the GS and PBSts muscle were determined. Control measurements were performed in the absence of any reflex stimulation, while test measurements were made during reflex stimulation. The reflex stimuli consisted of manually performed movements of the contralateral hind limb, muscle stretches, ligament tractions and electrical stimulations of cutaneous afferents. Altogether 21 secondary spindle afferents were investigated and 20 different reflex stimuli were employed. The general responsiveness (i.e. number of significant reflex effects/number of control-test series) was 52.4%, but a considerable variation between different stimuli was found, with the highest (89.9%) for contralateral whole limb extension and the lowest (25.0%) for stretch of the contralateral GS muscle. The size of the response to a given stimulus varied considerably between different afferents, and, in the same afferent, different reflex stimuli produced effects of varying size. Most responses were characterized by an increase in mean rate of discharge combined with a decrease in modulation, indicative of static fusimotor drive (Cussons et al., 1977). Since the secondary muscle spindle afferents are part of a positive feedback loop, projecting back to both static and dynamic fusimotor neurones (Appelberg Et al., 1892 a, 1983 b; Appelberg et al., 1986), it is suggested that the activity in the loop may work like an amplified which, during some circumstances, enhance the effect of other reflex inputs to the system (Johansson et al., 1991 b).