Coherence at 16-32 Hz can be caused by short-term synchrony of motor units.

Time- and frequency-domain measures of discharge times for pairs of motor units are used to infer the proportion of common synaptic input received by motor neurons. The physiological mechanisms that can produce the experimentally observed peaks in the cross-correlation histogram and the coherence spectrum are uncertain. The present study used a computational model to impose synchronization on the discharge times of motor units. Randomly selected discharge times of a unit that was being synchronized to a reference unit were aligned with some of the discharge times of the reference unit, provided the original discharge time was within 30 ms of the discharge by the reference unit. All time-domain measures (indexes CIS, E, and k') were sensitive to changes in the level of imposed motor-unit synchronization (P < 0.01). In addition, synchronization caused a peak between 16 and 32 Hz in the coherence spectrum. The shape of the cross-correlogram determined the frequency at which the peak occurred in the coherence spectrum. Further, the magnitude of the coherence peak was highly correlated with the time-domain measures of motor-unit synchronization (r2 > 0.80), with the highest correlation occurring for index E (r2 = 0.98). Thus the peak in the 16- to 32-Hz band of the coherence spectrum can be caused by the time that individual discharges are advanced or delayed to produce synchrony. Although the in vivo processes that adjust the timing of motor-unit discharges are not fully understood, these results suggest that they may not depend entirely on an oscillatory drive by the CNS.

[1]  A Schmied,et al.  Human spinal lateralization assessed from motoneurone synchronization: dependence on handedness and motor unit type. , 1994, The Journal of physiology.

[2]  A M Amjad,et al.  A framework for the analysis of mixed time series/point process data--theory and application to the study of physiological tremor, single motor unit discharges and electromyograms. , 1995, Progress in biophysics and molecular biology.

[3]  R K Powers,et al.  Relationship between simulated common synaptic input and discharge synchrony in cat spinal motoneurons. , 2001, Journal of neurophysiology.

[4]  P H Ellaway,et al.  Cumulative sum technique and its application to the analysis of peristimulus time histograms. , 1978, Electroencephalography and clinical neurophysiology.

[5]  David M. Halliday,et al.  Weak, Stochastic Temporal Correlation of Large-Scale Synaptic Input Is a Major Determinant of Neuronal Bandwidth , 2000, Neural Computation.

[6]  P H Ellaway,et al.  The source and distribution of short-term synchrony between gamma-motoneurones in the cat. , 1985, Quarterly journal of experimental physiology.

[7]  Ramón Huerta,et al.  Regularization mechanisms of spiking-bursting neurons , 2001, Neural Networks.

[8]  J. Wessberg,et al.  Single motor unit activity in relation to pulsatile motor output in human finger movements , 1999, The Journal of physiology.

[9]  Shi Zhou,et al.  Motor unit synchronisation is enhanced during slow lengthening contractions of a hand muscle , 2002, The Journal of physiology.

[10]  R. Enoka,et al.  Motor-unit synchronization increases EMG amplitude and decreases force steadiness of simulated contractions. , 2000, Journal of neurophysiology.

[11]  S. Farmer,et al.  Central nervous pathways underlying synchronization of human motor unit firing studied during voluntary contractions. , 1991, The Journal of physiology.

[12]  Stuart N Baker,et al.  Synchronization in monkey motor cortex during a precision grip task. II. effect of oscillatory activity on corticospinal output. , 2003, Journal of neurophysiology.

[13]  C. Marsden,et al.  Frequency peaks of tremor, muscle vibration and electromyographic activity at 10 Hz, 20 Hz and 40 Hz during human finger muscle contraction may reflect rhythmicities of central neural firing , 1997, Experimental Brain Research.

[14]  J. R. Rosenberg,et al.  The unilateral and bilateral control of motor unit pairs in the first dorsal interosseous and paraspinal muscles in man , 1999, The Journal of physiology.

[15]  John G Semmler,et al.  Motor Unit Synchronization and Neuromuscular Performance , 2002, Exercise and sport sciences reviews.

[16]  R. Enoka,et al.  Motor unit physiology: Some unresolved issues , 2001, Muscle & nerve.

[17]  R. Enoka,et al.  Motor-unit synchronization alters spike-triggered average force in simulated contractions. , 2002, Journal of neurophysiology.

[18]  J A Stephens,et al.  Correlation between the discharges of motor units recorded from the same and from different finger muscles in man. , 1991, The Journal of physiology.

[19]  V. Jousmäki,et al.  Task‐dependent modulation of 15‐30 Hz coherence between rectified EMGs from human hand and forearm muscles , 1999, The Journal of physiology.

[20]  S. Mori,et al.  Discharge patterns of soleus motor units with associated changes in force exerted by foot during quiet stance in man. , 1973, Journal of neurophysiology.

[21]  R. Hari,et al.  Cortical control of human motoneuron firing during isometric contraction. , 1997, Journal of neurophysiology.

[22]  J. R. Rosenberg,et al.  An extended difference of coherence test for comparing and combining several independent coherence estimates: theory and application to the study of motor units and physiological tremor , 1997, Journal of Neuroscience Methods.

[23]  T. Sears,et al.  The synaptic connexions to intercostal motoneurones as revealed by the average common excitation potential. , 1978, The Journal of physiology.

[24]  J. Stephens,et al.  Synchronization of motor unit activity during voluntary contraction in man. , 1990, The Journal of physiology.

[25]  D. Winter,et al.  Models of recruitment and rate coding organization in motor-unit pools. , 1993, Journal of neurophysiology.

[26]  T. Sears,et al.  Cross-Correlation Analyses of Motoneuron Inputs in a Coordinated Motor Act , 1991 .

[27]  R. C. Oldfield The assessment and analysis of handedness: the Edinburgh inventory. , 1971, Neuropsychologia.

[28]  Stuart N Baker,et al.  The effect of diazepam on motor cortical oscillations and corticomuscular coherence studied in man , 2003, The Journal of physiology.

[29]  R. Lemon,et al.  Human Cortical Muscle Coherence Is Directly Related to Specific Motor Parameters , 2000, The Journal of Neuroscience.

[30]  E. Fetz,et al.  Short-term synchronization of motor units in human extensor digitorum communis muscle: relation to contractile properties and voluntary control , 2004, Experimental Brain Research.

[31]  J. R. Rosenberg,et al.  Using electroencephalography to study functional coupling between cortical activity and electromyograms during voluntary contractions in humans , 1998, Neuroscience Letters.

[32]  R K Powers,et al.  Relationship between the time course of the afterhyperpolarization and discharge variability in cat spinal motoneurones , 2000, The Journal of physiology.

[33]  Gary Kamen,et al.  Motor unit synchronization in young and elderly adults , 2000, European Journal of Applied Physiology.

[34]  Håkan Johansson,et al.  Modern Techniques in Neuroscience Research , 1999, Springer Berlin Heidelberg.

[35]  T S Miles,et al.  Synchronization of motor units in human masseter during a prolonged isometric contraction. , 1990, The Journal of physiology.

[36]  R N Lemon,et al.  Synchronization in monkey motor cortex during a precision grip task. I. Task-dependent modulation in single-unit synchrony. , 2001, Journal of neurophysiology.

[37]  Allen I. Selverston,et al.  Modeling observed chaotic oscillations in bursting neurons: the role of calcium dynamics and IP3 , 2000, Biological Cybernetics.

[38]  M. Hepp-Reymond,et al.  EMG activation patterns during force production in precision grip , 2004, Experimental Brain Research.

[39]  M. Hepp-Reymond,et al.  EMG activation patterns during force production in precision grip , 2004, Experimental Brain Research.

[40]  R. Enoka,et al.  Quantification of the factors that influence discharge correlation in model motor neurons. , 2004, Journal of neurophysiology.

[41]  E. Fetz,et al.  Effects of synchrony between primate corticomotoneuronal cells on post-spike facilitation of muscles and motor units , 1989, Neuroscience Letters.

[42]  A I Selverston,et al.  Nonlinear behavior of sinusoidally forced pyloric pacemaker neurons. , 2001, Journal of neurophysiology.

[43]  Carlo J De Luca,et al.  Common drive in motor units of a synergistic muscle pair. , 2002, Journal of neurophysiology.

[44]  N. Davey,et al.  Rhythmicity associated with a high degree of short‐term synchrony of motor unit discharge in man , 1993, Experimental physiology.

[45]  R. Person,et al.  Discharge frequency and discharge pattern of human motor units during voluntary contraction of muscle. , 1972, Electroencephalography and clinical neurophysiology.

[46]  J. R. Rosenberg,et al.  The Fourier approach to the identification of functional coupling between neuronal spike trains. , 1989, Progress in biophysics and molecular biology.

[47]  A J Fuglevand,et al.  Estimating the strength of common input to human motoneurons from the cross‐correlogram. , 1992, The Journal of physiology.

[48]  L M Harrison,et al.  Physiological tremor in human subjects with X‐linked Kallmann's syndrome and mirror movements , 2001, The Journal of physiology.

[49]  B. Conway,et al.  Synchronization between motor cortex and spinal motoneuronal pool during the performance of a maintained motor task in man. , 1995, The Journal of physiology.

[50]  C. Marsden,et al.  Physiological and pathological tremors and rhythmic central motor control. , 2000, Brain : a journal of neurology.

[51]  P. Matthews,et al.  Measurement of excitability of tonically firing neurones tested in a variable‐threshold model motoneurone , 2002, The Journal of physiology.

[52]  M Swash,et al.  Changes in motor unit synchronization following central nervous lesions in man. , 1993, The Journal of physiology.

[53]  N Kakuda,et al.  Common modulation of motor unit pairs during slow wrist movement in man , 1999, The Journal of physiology.

[54]  D. B. Preston Spectral Analysis and Time Series , 1983 .

[55]  R. Enoka,et al.  Motor-unit coherence during isometric contractions is greater in a hand muscle of older adults. , 2003, Journal of neurophysiology.

[56]  M. Schubert,et al.  Short term synchronization of human motor units and their responses to transcranial magnetic stimulation. , 1995, The Journal of physiology.

[57]  J. R. Rosenberg,et al.  Load-independent contributions from motor-unit synchronization to human physiological tremor. , 1999, Journal of neurophysiology.

[58]  P. Ellaway,et al.  THE SOURCE AND DISTRIBUTION OF SHORT-TERM SYNCHRONY BETWEEN γ-MOTONEURONES IN THE CAT , 1985 .

[59]  R. Enoka,et al.  Mechanisms that contribute to differences in motor performance between young and old adults. , 2003, Journal of electromyography and kinesiology : official journal of the International Society of Electrophysiological Kinesiology.

[60]  S. Farmer,et al.  Abnormal cortex-muscle interactions in subjects with X-linked Kallmann's syndrome and mirror movements. , 2004, Brain : a journal of neurology.

[61]  R M Enoka,et al.  Motor-unit synchronization is not responsible for larger motor-unit forces in old adults. , 2000, Journal of neurophysiology.

[62]  Roger M Enoka,et al.  Multiple features of motor-unit activity influence force fluctuations during isometric contractions. , 2003, Journal of neurophysiology.

[63]  D. Halliday,et al.  The frequency content of common synaptic inputs to motoneurones studied during voluntary isometric contraction in man. , 1993, The Journal of physiology.

[64]  R N Lemon,et al.  Modulation of synchrony between single motor units during precision grip tasks in humans , 2002, The Journal of physiology.

[65]  J. R. Rosenberg,et al.  Time and Frequency Domain Analysis of Spike Train and Time Series Data , 1999 .

[66]  T. Sears,et al.  Short‐term synchronization of intercostal motoneurone activity. , 1976, The Journal of physiology.

[67]  J. R. Rosenberg,et al.  A review of recent applications of cross-correlation methodologies to human motor unit recording , 1997, Journal of Neuroscience Methods.