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C N Christakos

Publications and source records attributed to C N Christakos.

At least 19 recordsLinked to original sources

Analysis of recurrent laryngeal inspiratory discharges in relation to fast rhythms.

1. Inspiratory (I) activities of recurrent laryngeal (RL) motoneurons and efferent nerves were studied by autospectral, interval, and coherence analyses, with emphasis on fast rhythms of two types: medium-frequency oscillations (MFO, usual range 20-50 Hz for nerve autospectral peaks) and high-frequency oscillations (HFO, usual range 50-100 Hz). 2. In decerebrate, paralyzed, and artificially ventilated cats, recordings were taken from 27 isolated single RL fibers (14 cats) and 8 identified RL motoneurons in the medulla (6 cats), together with recordings of phrenic (PHR) and RL whole-nerve activities. In another 50 cats, RL and PHR nerve discharges were recorded simultaneously. 3. The autospectra of RL units showed prominent MFO peaks with frequencies close to that of the RL nerve MFO spectral peak, indicating presence of this type of fast rhythm in the units' discharges. Spectral analysis of RL unit activity in different segments of the I phase showed that the frequency of a unit's MFO was very close to the peak (maintained) firing rate of the unit during the portion of I analyzed. Thus a motoneuron's MFO spectral peak reflected its rhythmic discharge arising from the cell's refractoriness (and possibly with the rate changing in the course of I). 4. The coherences of motoneurons' MFOs to nerve MFOs were very low or 0, indicating that correlations between unitary MFOs of the RL population were rare and/or weak. 5. In those cats (19/20) that had discernible PHR nerve HFO autospectral peaks, about half of the recorded RL motoneurons (16/34) had HFO. For these motoneurons, the unit-nerve HFO coherences were substantial, indicating widespread correlations between unitary HFOs. 6. In a fraction of cats, coherence peaks in the MFO frequency range were observed between bilateral RL nerves, and between RL and PHR nerves, at frequencies that were subharmonics of the HFO frequency. 7. In light of theoretical considerations on the generation of aggregate rhythms from superposition of unitary rhythms, these observations indicate that, similarly, to the case of PHR motoneurons and nerves. 1) RL nerve MFO arises from superposition of uncorrelated, or at most partially correlated, MFOs of RL units, representing the rhythmic discharges of the cells. It is manifested therefore as a spectral deflection with a maximum in the band of peak firing rates of the units. 2) RL nerve HFO arises from correlated, common-frequency HFOs in a subpopulation of RL units, caused by HFO inputs from antecedent medullary I neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Fast rhythms in phrenic motoneuron and nerve discharges.

1. Fast rhythms in discharges of individual phrenic (PHR) motoneurons were studied by spectral and interval analyses; and they were compared, using coherence analysis, with similar rhythms in whole-PHR nerve discharge. The purpose of this study was to ascertain the origin of the two rhythms, manifested as distinct spectral peaks, in PHR motoneuron and nerve discharge: medium-frequency oscillations (MFO, usual range 20-50 Hz); and high-frequency oscillations (HFO, usual range 50-100 Hz). 2. In paralyzed artificially ventilated cats, unit recordings were taken from 1) 26 isolated single PHR fibers (in 8 sodium pentobarbital-anesthetized cats) and 2) 27 identified PHR motoneuron somata in the spinal cord (in 5 decerebrate cats). Simultaneous whole-PHR activity was monophasically recorded from the contralateral PHR nerve for 1 and from both PHR nerves for 2. 3. The signals were subjected to time- and frequency-domain analyses. The latter included a novel application of coherence analysis to the study of population synchrony. 4. The autospectra of all PHR units showed prominent MFO peaks in the frequency range of the nerve MFO spectral peaks, as well as harmonic peaks, indicating the presence of this type of fast rhythm in the units' discharges. Spectral analysis of the augmenting PHR activities in different segments of the inspiratory (I) phase showed that the frequency of unit MFO and of nerve MFO rose during the course of I. Further, cycle-triggered histogram and interval analysis indicated that the frequencies of unit MFO autospectral peaks were very close to the peak firing rates of the units during the portion of I analyzed. Thus unit MFO spectral peaks reflected the rhythmic and augmenting discharges of the motoneurons, and similar nerve MFO peaks reflected the superposition of individual motoneuron discharges. 5. The coherences of motoneurons' MFOs to nerve MFOs were low or zero, indicating that only partial and weak MFO correlations occurred within the PHR motoneuron population. 6. In those cats (n = 11) that had clear PHR nerve HFO spectral peaks, about one-half of the recorded PHR motoneurons had HFO, as indicated by HFO peaks in the unit autospectra and/or the unit-nerve coherences. 7. For motoneurons having HFO, the coherence between unit and nerve HFOs was substantial, particularly when the latter were strong, indicating HFO correlations among a number of PHR motoneurons. 8. In the light of theoretical considerations on the generation of aggregate rhythms from superposition of unitary rhythms, these observations indicate the following.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Changes in frequency content of inspiratory neuron and nerve activities in the course of inspiration.

In decerebrate paralyzed cats, the spectra and coherences of inspiratory (I) nerve activities and of medullary I neuron discharges were compared between different stages of I. The correlated high-frequency oscillations (HFOs) in the activities had common time courses of frequency and strength, which were influenced by lung afferent input; whereas the time courses for the uncorrelated medium-frequency oscillations (MFOs) depended on individual activity patterns. These results indicate that HFOs are characteristic of the common I pattern generator, whereas MFOs are specific to individual activities.

Action Potentials

Fast rhythms in the discharges of medullary inspiratory neurons.

The discharges of 44 medullary inspiratory (I) neurons in decerebrate paralyzed cats were studied using interval and spectral analysis. Most neurons had a rhythm in their discharge. In 31 the rhythm was at the frequency of, and coherent to, the high-frequency oscillations (HFOs) of I nerves, and in 7 the rhythm was in the range of medium-frequency oscillations (MFOs), with no coherence to nerve MFOs. Thus, correlated HFOs are characteristic of the I system at all levels, whereas MFOs are uncommon in medullary neurons and seem to be unrelated to general mechanisms.

Action Potentials

Intracellular potentials and discharge patterns of expiratory neurons in the caudal ventral respiratory group: influence of phasic pulmonary afferent input.

In decerebrate paralyzed cats, the membrane potential (MP) patterns of 12 augmenting expiratory (E) neurons in the caudal ventral respiratory group, and phrenic and recurrent laryngeal activities, were compared for inspiratory (I) phases with and without lung inflation. No-inflation produced, in the MPs of E neurons, larger hyperpolarization during I and during early E (associated with increased early-E laryngeal activity), suggesting an increase of inhibitory inputs from I neurons and early-E neurons, respectively.

Action Potentials

High-frequency and medium-frequency components of different inspiratory nerve discharges and their modification by various inputs.

In decerebrate paralyzed cats, spectral analysis was performed on simultaneous recordings of efferent inspiratory nerves (phrenic, recurrent laryngeal, hypoglossal). Spectral peaks were present both in the high-frequency (HFO) range (50-100 Hz) and the medium-frequency (MFO) range (20-50 Hz). Different activities were coherent only in the HFO range, indicating that the HFOs arise in a common inspiratory pattern generator that drives the different motoneuron populations, whereas the MFOs are specific to different systems.

Animals

Frequency response of spinal Renshaw cells activated by stochastic motor axon stimulation.

In anaesthetized or decerebrate cats, motor axons in lumbosacral ventral roots or hindlimb muscle nerves were stimulated with random trains of brief electrical pulses, and Renshaw cell spike sequences were recorded. Spectral analysis was used to determine the range of linear operation of Renshaw cells, via coherence computations, and to calculate their frequency-dependent gains and phases. The analysis showed that the dynamic behaviour of Renshaw cells was different for different strengths of their synaptic input from motor axons and for different mean stimulus rates. In general, the changes in dynamics associated with variation of these two input parameters followed a common trend. This can be related to the average response of Renshaw cells per stimulus, as assessed by peri-stimulus time histograms. For axons having a strong excitatory effect on a Renshaw cell (as judged from the size of early peri-stimulus time histogram peaks), and for low mean stimulus rates (10-23 pulses per second), the linear range of signal transmission (assessed by coherence computation) was usually very broad (from zero sometimes up to over 100 Hz, but mostly up to 50-100 Hz). Following an initial elevation in the range 2-15 Hz, the gain showed first a rapid decrease with frequency, down to a value which at 30-50 Hz could be a tenth of the gain at lower frequencies (2-15 Hz); it then continued to decline slowly. Otherwise the linear range was narrower and/or the coherence was generally lower; the gain was lower and showed little decline with frequency. The phase curves of Renshaw cells generally showed a low-frequency phase lead (up to roughly 10 Hz) and an increasing phase lag thereabove that was generated in part by the conduction delay. The results show that Renshaw cells can follow, particularly sensitively, inputs in a frequency range encompassing the steady firing rates of many alpha-motoneurons. This range of high gain also covers that of a component of physiological tremor (ca. 6-12 Hz), a basic mechanism of which is probably related to unfused contractions of newly recruited motor units firing in this range. It can therefore be expected that recurrent inhibition via Renshaw cells is especially powerful in this physiologically important range of alpha-motoneuron firing.

Action Potentials

Amplitude reduction of motor unit twitches during repetitive activation is accompanied by relative increase of hyperpolarizing membrane potential trajectories in homonymous alpha-motoneurons.

In anaesthetized cats, medial gastrocnemius motor units (MUs) were electrically stimulated via their ventral-root axons with independent random patterns. Isometric muscle tension and homonymous alpha-motoneuron (MN) membrane-potential fluctuations in response to these stimuli were recorded simultaneously, usually for periods of about 2 min. The tension and membrane potential were averaged with respect to a stimulus train over two disjoint time intervals, one stretching 20-40 s at record beginning, and the other a similar duration at the end of recording. Whereas average MU twitch amplitudes usually decreased between these periods, average membrane potential trajectories did not do so, such that, when normalized to the change in twitch amplitude, the membrane potential trajectories usually increased in size. This suggests that the decline in the mechanical effect of MU activation was accompanied by an increase in the gain of the afferent pathway to homonymous MNs, which was confirmed by gain computations in the frequency domain. This compensation could be a mechanism to maintain the high quality of information about MU contractions transmitted to MNs in the course of MU fatigue.

Animals

The information carried by spindle afferents on motor unit activity as revealed by spectral analysis.

Spectral analysis was used to study the effects of motor unit activity on the discharge patterns of muscle spindle endings. Spindle afferents of hind-limb muscles of the cat were recorded during electrical stimulation of one or more motor units, and, for comparison, while the receptors discharged in the absence of induced extrafusal activity ('background discharge'). The stimulus sequences used were random, but had characteristic frequency components representing an underlying rhythm, similar to those of trains in real alpha-motoneuron output. The computed afferent spectra and coherences between stimulus and afferent trains indicate that the discharge patterns of muscle spindles carry information on the activity of particular subsets of motor units. The spectra also demonstrate a complex interaction of internal spindle (pacemaker) mechanisms and external (modulating) processes which determine the discharge patterns of primary and secondary endings. In addition, they reveal interesting differences between primaries and secondaries, possibly indicative of a particular role for each type of ending in motor control.

Action Potentials

Spindle gain increase during muscle unit fatigue.

In anaesthetized cats, medial gastrocnemius motor units (MUs) were stimulated with random sequences (mean rates between 6 and 12 pps) of electrical pulses delivered to their axons in small ventral root filaments. Muscle tension was recorded under isometric conditions, and spike trains of muscle spindle afferents were recorded from small dorsal root filaments during prolonged MU activation. Time-domain (PSTH) and frequency-domain (gain) computations were performed to study the effects of fatiguing muscle unit contractions on the signal transmission from skeletomotor efferents to spindle afferents. In the course of muscle unit fatigue, during which the gain of the force-producing sub-system decreased, the gain of the sub-system transforming force to afferent discharge increased so that the overall gain between skeletomotor efferents and spindle afferents remained relatively high. This could be a mechanism that preserves a high quality of afferent information on MU contractions.

Animals

Dominance of the short-latency component in perturbation induced electromyographic responses of long-trained monkeys.

The effects of prolonged training of adult monkeys subjected to random, brief perturbations of alternating elbow flexions and extensions were studied over a period of four years. The training was intensive at first, for about one year, and then irregular, with long pauses, during the following three years. As a consequence of the prolonged training with the brief perturbations, the M2 component of the electromyographic (EMG) response of the biceps and triceps muscles became gradually smaller, and finally disappeared. The M1 component, on the other hand, progressively increased in amplitude and continued to do so after the loss of the M2, until it finally dominated the EMG response. The training had similar effects on the response of the biceps muscle to longer perturbations, but, only under certain conditions, did it affect the triceps muscle response. All changes occurred at earlier stages of the training in the flexor than in the extensor muscle. These observations demonstrate a long-term functional plasticity of the sensorimotor system of adult animals and suggest a growing role for fast segmental mechanisms in the reaction to external disturbances as motor learning progresses. Changes at various levels of the stretch reflex system could underlie the enlargement of the M1 component, while the lack of the M2 component should, at least partially, reflect a reduced cortical effect on alpha-motoneurones and/or changes in spinal systems processing afferent information.

Animals

The mathematical basis of population rhythms in nervous and neuromuscular systems.

The mechanism underlying rhythmical aggregate activity of a population of neural or neuromuscular elements is examined in this report. By making use of the spectral properties of stochastic processes (Papoulis, 1965), it is shown that such population rhythms are the inevitable effect of the rhythmical activities of the individual elements, irrespective of the phase relations of the latter. This result applies to both "discrete" signals, such as spike trains, and "continuous" ones, such as membrane potential fluctuations. It has implications regarding the generation of common physiological rhythms and the preservation of rhythms when converging activity of one of the above two types is transformed into activity of the other type.

Action Potentials

Note on the estimation of the correlation function of neural spike trains.

The use of time-bins in the estimation of the correlation function of neural spike trains has a filtering effect on the estimate and results in distortion and aliasing. Prior low-pass filtering of the spike trains, on the other hand, and computation of the correlation function of the emerging waveforms in the standard way result in an estimate that is also a filtered version of the original function but distortion- and alias -free. In addition, the correlation function so computed can be normalized. An analogous definition of the correlation coefficient for the first technique enables the comparison of these various correlation estimates and clarifies their properties.

Animals

The use of frequency domain techniques in the study of signal transmission in skeletal muscle.

Spectral analysis provides a description of the moments of random signals and enables the characterization of the behaviour of systems in terms of input-output relations. The merits of such an approach in the study of signal transmission in skeletal muscle are described in this paper. The representation of neural spike trains as impulse sequences and the subsequent treatment appropriate for this kind of analysis are discussed together with some practical problems. Spectral analysis of muscle afferent signals is applied to data obtained from cat experiments, and the use of the related frequency-domain techniques is demonstrated on a subsystem of the stretch reflex.

Action Potentials

The 'M2' electromyographic response to random perturbations of arm movements is missing in long-trained monkeys.

Adult monkeys who have been under training over a long period of time show a loss of the M2 component in their biceps electromyographic response to brief, random perturbations of an alternating arm movement, the M1 component being apparently enhanced. These changes of the stretch-induced responses indicate a long-term plasticity of the sensorimotor system of monkeys. This result also provides hints for the origin and possible functional significance of the short- and long-latency components of the electromyographic response in subjects with less experience.

Animals

A linear stochastic model of the single motor unit.

The production of force and of the electrical signal by an active motor unit is theoretically described. Neural spikes are modelled using the Dirac delta function. Mechanisms for the generation of random impulse trains and the properties of the corresponding stochastic processes are discussed; the "renewal" model is proposed as the most appropriate. The possibility of using a linear model for the systems that produce force and electrical signal in the unit is examined. It is concluded that the linear assumption is justifiable during steady, constant-strength contractions of muscle. This linear stochastic model of the motor unit is used in two subsequent papers to study the muscle force and the electromyogram.

Animals

A study of the muscle force waveform using a population stochastic model of skeletal muscle.

A population stochastic model based on the differing properties and the independent activation of motor units is used to describe the production of force in the contracting skeletal muscle. Detailed force predictions of the model concerning a hand muscle are obtained by computer simulation. General features of the force signal are established analytically on the basis of the general properties of the neuromuscular system which the population model takes into account. The results show that the asynchronous activity of motor units and the distribution of their filtering and firing properties at various levels of muscle contraction are responsible, at least partially, for the main features of the muscle force waveform, including tremor.

Animals