Influence of amplitude cancellation on the simulated surface electromyogram.

The purpose of the study was to quantify the influence of selected motor unit properties and patterns of activity on amplitude cancellation in the simulated surface electromyogram (EMG). The study involved computer simulations of a motor unit population with physiologically defined recruitment and rate coding characteristics that activated muscle fibers whose potentials were recorded on the skin over the muscle. Amplitude cancellation was quantified as the percent difference in signal amplitude when motor unit potentials were summed before and after rectification. The simulations involved varying the level of activation for the motor unit population, the recording configuration, the upper limit of motor unit recruitment, peak discharge rates, the amount of motor unit synchronization, muscle fiber length, the thickness of the subcutaneous tissue, and the motor unit properties that change with advancing age. The results confirmed a previous experimental report (Day SJ and Hulliger M, J Neurophysiol 86: 2144-2158, 2001) that amplitude cancellation in the surface EMG can reach 62% at maximal activation. A decrease in the range of amplitudes of the motor unit potentials, as can occur during fatiguing contractions, increased amplitude cancellation up to approximately 85%. Differences in the amount of amplitude cancellation were observed across all simulated conditions, and resulted in substantial changes in the absolute magnitude of the EMG signal. The most profound factors influencing amplitude cancellation were the number of active motor units and the duration of the action potentials. The effects of amplitude cancellation were minimal (<5%) when the EMG amplitude was normalized to maximal values, with the exception of variations in peak discharge rate and recruitment range, which resulted in differences up to 17% in the normalized EMG signal across conditions. These results indicate the amount of amplitude cancellation that can occur in various experimental conditions and its influence on absolute and relative measures of EMG amplitude.     

The influence of contraction type,prior performance of a maximal voluntary contraction and measurement duration on fine-wire EMG amplitude

We aimed to investigate the impact of time on fine-wire (fw) electromyography (EMG) signal amplitude, and to determine whether any attenuation is confounded by task type. Twenty healthy participants were instrumented with fw and surface (s) EMG electrodes at the biceps brachii bilaterally. Participants held a weight statically with one arm and with the other arm either repeated the same task following a maximum voluntary contraction (MVC) or repeated dynamic elbow flexion/extension contractions. Each task was repeated for 30 s every five minutes over two hours. EMG amplitude was smoothed and normalized to time = 0. Stable median power frequency of the s-EMG ruled out the confounding influence of fatigue. Repeated-measures ANCOVAs determined the effect of electrode type and time (covariate) on EMG amplitude and the confounding impact of task type. During the isometric protocol, fw-EMG amplitude reduced over time (p = 0.002), while s-EMG amplitude (p = 0.895) and MPF (p > 0.05) did not change. Fw-EMG amplitude attenuated faster during the dynamic than the isometric protocol (p = 0.008) and there was evidence that the MVC preceding the isometric protocol impacted the rate of decline (p = 0.001). We conclude that systematic signal attenuation of fw-EMG occurs over time and is more pronounced during dynamic tasks.     

A comparison of three noise reduction procedures applied to bird vocal signals

ABSTRACT.   Recordings of avian vocal signals in natural habitats include ambient noise. Often this background noise corrupts across all frequencies and is of substantial amplitude. Reducing this ambient noise to prepare vocal signals for playback stimuli or to remove habitat-specific noise signatures prior to analyzing a signal's acoustic characteristics can be useful. We conducted experimental evaluations of three noise reduction procedures to determine their effectiveness. We embedded two bird vocalizations ("clean" signals) in four kinds of natural noise, resulting in eight noise-signal combinations. We then applied three noise reduction procedures (Noise Profile, Band Pass, and Noise Estimate) to each of the embedded signals and compared the recovered signals to the original (clean) signals. Noise Profile filtering was effective in reducing noise and returning fairly high-quality signals from even severe levels of masking noise. The other two noise reduction procedures did not perform as well. For the two most corrupting maskers, however, Noise Profile filtering also altered the signal properties by reducing signal amplitude at those frequencies containing high levels of noise. Apart from this loss of amplitude, the quantitative features of the filtered signals were similar to those of the original model sounds. We conclude that Noise Profile filtering produces good results for cases where noise is approximately constant over the signal duration and the signal intensity exceeds noise intensity over the frequencies of interest.     

Response of frog midbrain neurons to tones amplitude-modulated by pseudorandom noise

Spike response in torus semicircularis units to the effects of uninterrupted characteristic frequency tones amplitude-modulated by pseudorandom noise were investigated during experiments on immobilizedRana ridibunda. Period histograms of modulating waveform of 512 msec duration (both modulating polarities) were produced for 32 units. Almost all neurons investigated responded exclusively to the positive half of the modulating signal. Difference histograms obtained by calculating period histograms for different polarities of the envelope faithfully reproduced the dynamics of signal amplitude in four units. The remainder responded only to envelope maxima, without reproducing amplitude dynamics among these; over half the units represented only some of the envelope maxima, moreover. Certain cells were found which retained their specific pattern of response to pseudorandom noise over a wide range of carrier intensities.N. N. Andreev Acoustical Institute, Moscow. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 227–235, March–April, 1990.     

Acoustic myography as an indicator of force during sustained contractions of a small hand muscle

To test the hypothesis that muscle sound amplitudes would remain constant during sustained submaximal isometric contractions, we recorded acoustic myograms from the abductor digiti minimi muscle in 12 subjects at 15, 25, 50, and 75% of a maximum voluntary contraction (MVC). Muscle sounds were detected with an omni-directional electret microphone encased in closed-cell foam and attached to the skin over the muscle. Acoustic amplitudes from the middle and end of the sustained contractions were compared with the amplitudes from the beginning of contractions to determine whether acoustic amplitudes varied in magnitude as force remained constant. Physiological tremor was eliminated from the acoustic signal by use of a Fourier truncation at 14 Hz. The amplitudes of the acoustic signal at a contraction intensity of 75% MVC remained constant, reflecting force production over time. At 50% MVC, the root-mean-square amplitude decreased from the beginning to the end of the contraction (P less than 0.05). Acoustic amplitudes increased over time at 15 and 25% MVC and were significantly higher at the end of the contractions than at the beginning (P less than 0.05). Alterations in the acoustic amplitude, which reflect changes in the lateral vibrations of the muscle, may be indicative of the different recruitment strategies used to maintain force during sustained isometric contractions.     

The transmission of advertisement calls in Central American frogs

Acoustic communication signals change over distance due to lossof amplitude and fidelity, and it is assumed that signal degradationinfluences the receiver's ability to detect and decode signals.The degree of degradation depends on the signal's structureand the environment through which it transmits. We broadcastthe advertisement calls of 22 species of Central American frogsat two heights within forested and open environments in Panama. Werecorded these calls at five distances from the source and estimatedsignal degradation with a cross-correlation analysis, a measurethat combines the effects of decrement in signal amplitude andfidelity. Calls degraded less when broadcast higher above theground compared to on the ground, and less in open habitat comparedto forested habitat; there was an additional interaction betweenheight and environment. Furthermore, calls with low dominant frequenciesexperienced less degradation than calls with high dominant frequencies.There was no evidence, however, that the calls of these frogs haveevolved to maximize habitat-specific transmission.     

The vocal repertoires of the bush dog (Speothos venaticus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus)

The vocalizations of the bush dog (Speothos venaticus), crab-eating fox (Cerdocyon thous), and maned wolf (Chrysocyon brachyurus) were studied in a captive setting. The three species' repertoires were similar and included whines, long-distance calls, and growls. The whines and growls varied in amplitude, duration, and repetition rate; these variations and the frequencies of occurrence probably were influenced by both social organization and habitat characteristics. The bush dog, which lives in family groups and inhabits rain forests, had an elaborate whine-scream continuum that at close distances conveyed a wide range of moods. On the other hand, the maned wolf, which is solitary and frequents open pampas, used amplitude variations of the growl to signal hostile intentions over short, medium, and long distances.The long-distance vocalizations of the three species were the least variable in structure; the bush dog and crab-eating fox used the call to promote contact with separated pairmates or family members. Based on recordings during development, these two calls appeared to arise from the repetitive whine, a variant of the basic whine syllable. The maned wolf also had a contact-promoting howl, apparently an extended whine of high amplitude, but it was rarely recorded during the present study. More prominent was the maned wolf roar-bark, which functioned to space individuals over large distances, and appeared to be a growl of high amplitude.     

DNA analysis and sorting of rat testis cells using two-parameter flow cytometry

By use of two-parameter flow cytometry of rat testis cell suspensions stained with mithramycin for DNA (the peak amplitude of the fluorescence signal versus total fluorescence intensity integrated over time), eight cell compartments could be distinguished without pre-enrichment of the samples. Cells in these compartments were identified by sorting and subsequent microscopic examination.     

Principles behind the multifarious control of signal transduction. ERK phosphorylation and kinase/phosphatase control

General and simple principles are identified that govern signal transduction. The effects of kinase and phosphatase inhibition on a MAP kinase pathway are first examined in silico. Quantitative measures for the control of signal amplitude, duration and integral strength are introduced. We then identify and prove new principles, such that total control on signal amplitude and on final signal strength must amount to zero, and total control on signal duration and on integral signal intensity must equal -1. Collectively, kinases control amplitudes more than duration, whereas phosphatases tend to control both. We illustrate and validate these principles experimentally: (a) a kinase inhibitor affects the amplitude of EGF-induced ERK phosphorylation much more than its duration and (b) a phosphatase inhibitor influences both signal duration and signal amplitude, in particular long after EGF administration. Implications for the cellular decision between growth and differentiation are discussed.     

A portable miniature solid-state heart rate recorder for monitoring daily physical activity

A compact, rugged, solid-state heart rate recorder (63 X 94 X 22 mm, 195 g) to measure normal daily heart rates over periods of up to 33 h is described. With an interval switch a selection can be a mean heart rate over 30, 60 or 120 sec. The ECG amplifier filter is equipped with an automatic gain control to eliminate R-wave amplitude changes during physical activity. A continuous acoustic check on correct electrode attachment and sufficient signal amplitude is possible by means of an earphone. Data may be stored on punch tape, tape recorder, or processed on-line, or plotted directly one a line writer to obtain a heart rate profile over the measured time. In 45 studies with adults and children the heart rate recorder has proven to be a reliable instrument.     

Temporal processing and adaptation in the songbird auditory forebrain

Songbird auditory neurons must encode the dynamics of natural sounds at many volumes. We investigated how neural coding depends on the distribution of stimulus intensities. Using reverse-correlation, we modeled responses to amplitude-modulated sounds as the output of a linear filter and a nonlinear gain function, then asked how filters and nonlinearities depend on the stimulus mean and variance. Filter shape depended strongly on mean amplitude (volume): at low mean, most neurons integrated sound over many milliseconds, while at high mean, neurons responded more to local changes in amplitude. Increasing the variance (contrast) of amplitude modulations had less effect on filter shape but decreased the gain of firing in most cells. Both filter and gain changes occurred rapidly after a change in statistics, suggesting that they represent nonlinearities in processing. These changes may permit neurons to signal effectively over a wider dynamic range and are reminiscent of findings in other sensory systems.     

The intracellular pupil mechanism and photoreceptor signal: noise ratios in the fly Lucilia cuprina

The function of the intracellular pupil mechanism is examined by comparing the responses of photoreceptors in normal flies with those from white-eyed flies that lack the pupil. In white-eyed flies the response to an intensity increment of fixed contrast decreases at high background intensities. There is a smaller decrease in noise amplitude so that the signal:noise ratio falls. The intensity dependence of the photoreceptor signal:noise ratio fits a simple model in which activated photopigment molecules compete for 3 X 10(4) transduction units. The signal:noise ratio decreases at high intensities because the transduction units are saturated. This model is supported by a noise analysis, which provides three estimates of the number of events generating photoreceptor responses. In white-eyed flies the event number saturates at high background intensities, suggesting that a maximum of 2 X 10(4) events can be simultaneously active. Wild-type flies do not exhibit saturation effects over the range of intensities studied. The signal:noise ratio rises with intensity to reach a stable asymptote, close to the maximum observed for white-eyed flies. Pupil attenuation is calculated from measurements of signal:noise ratio in white-eyed and wild-type flies. The pupil is progressively activated over a two log unit intensity range and when fully closed attenuates the effective intensity by 99%. The threshold of this pupil effect coincides with the threshold of pupil activation measured optically. We conclude that the intracellular pupil attenuates the light flux to prevent receptor saturation and to extend the range of intensities at which fly photoreceptors operate close to their maximum signal:noise ratio. This upper limit is determined by the number of transduction units generating a cell's response.     

MAPK cascade possesses decoupled controllability of signal amplification and duration

The three important characteristics of the output signal of mitogen activated protein kinase (MAPK) cascade are time delay between stimulus and response, amplitude gain, and duration of the output signal. In this study, we performed a sensitivity analysis on the computational model of epidermal growth factor receptor (EGFR) activated MAPK cascade developed by Schoeberl and co-workers (1) to identify the sensitive steps of the pathway affecting these characteristics. We show that the signaling network is sensitive in a decoupled manner, which provides the ability to control its output amplitude and duration one at a time. Signal duration is found sensitive only to the phosphatase reactions at the MEK level. In contrast, signal amplitude is found most sensitive to the phosphatase reactions at the ERK level. Time delay is found to be a robust characteristic of the system.     

Comparison of Coding Capabilities of Type I and Type II Neurons

We consider the dependence of information transfer by neurons on the Type I vs. Type II classification of their dynamics. Our computational study is based on Type I and II implementations of the Morris-Lecar model. It mainly concerns neurons, such as those in the auditory or electrosensory system, which encode band-limited amplitude modulations of a periodic carrier signal, and which fire at random cycles yet preferred phases of this carrier. We first show that the Morris-Lecar model with additive broadband noise ("synaptic noise") can exhibit such firing patterns with either Type I or II dynamics, with or without amplitude modulations of the carrier. We then compare the encoding of band-limited random amplitude modulations for both dynamical types. The comparison relies on a parameter calibration that closely matches firing rates for both models across a range of parameters. In the absence of synaptic noise, Type I performs slightly better than Type II, and its performance is optimal for perithreshold signals. However, Type II performs well over a slightly larger range of inputs, and this range lies mostly in the subthreshold region. Further, Type II performs marginally better than Type I when synaptic noise, which yields more realistic baseline firing patterns, is present in both models. These results are discussed in terms of the tuning and phase locking properties of the models with deterministic and stochastic inputs.     

Regulation of vocal amplitude by the blue-throated hummingbird, Lampornis clemenciae

Animals that rely on vocal communication must broadcast sound so that a perceptible signal is transmitted over an appropriate distance. We found that male blue-throated hummingbirds modified the amplitude of their vocalizations in response to both naturally occurring and experimenter-controlled changes in ambient noise levels. This phenomenon is known as the Lombard effect and may increase the efficiency of acoustic signalling. This study demonstrates the effect under natural field conditions and documents the first hummingbird species (Apodiformes: Trochilidae) to show this behaviour. We measured sound pressure levels (SPLs) of Serial Chip territorial advertisement calls across a natural range of ambient noise, primarily due to creeks within male territories. We found a significant correlation between the amplitude of Serial Chips and the amplitude of background noise. To test this relationship, we broadcast recordings of creek noise at high and low amplitudes while target individuals were producing Serial Chip vocalizations. We measured vocal SPLs before and during the playback. Individuals responded to changes in playback creek noise by changing the amplitude of Serial Chip production. We also measured transmission properties of Serial Chip calls through natural habitat to calculate the approximate amplitude of vocalizations at the position of the calling bird. We suggest that amplitude regulation of vocalizations contibutes to signal transmission distance along with the established relationships between singing behaviour, acoustic structure and habitat. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.     

Modeling nonlinear errors in surface electromyography due to baseline noise: A new methodology

The surface electromyographic (EMG) signal is often contaminated by some degree of baseline noise. It is customary for scientists to subtract baseline noise from the measured EMG signal prior to further analyses based on the assumption that baseline noise adds linearly to the observed EMG signal. The stochastic nature of both the baseline and EMG signal, however, may invalidate this assumption. Alternately, “true” EMG signals may be either minimally or nonlinearly affected by baseline noise. This information is particularly relevant at low contraction intensities when signal-to-noise ratios (SNR) may be lowest. Thus, the purpose of this simulation study was to investigate the influence of varying levels of baseline noise (approximately 2–40% maximum EMG amplitude) on mean EMG burst amplitude and to assess the best means to account for signal noise. The simulations indicated baseline noise had minimal effects on mean EMG activity for maximum contractions, but increased nonlinearly with increasing noise levels and decreasing signal amplitudes. Thus, the simple baseline noise subtraction resulted in substantial error when estimating mean activity during low intensity EMG bursts. Conversely, correcting EMG signal as a nonlinear function of both baseline and measured signal amplitude provided highly accurate estimates of EMG amplitude. This novel nonlinear error modeling approach has potential implications for EMG signal processing, particularly when assessing co-activation of antagonist muscles or small amplitude contractions where the SNR can be low.     

Crosstalk in surface electromyography of the proximal forearm during gripping tasks.

Electromyographic (EMG) crosstalk was systematically analyzed to evaluate the magnitude of common signal present between electrode pairs around the forearm. Surface EMG was recorded and analyzed from seven electrode pairs placed circumferentially around the proximal forearm in six healthy individuals. The cross-correlation function was used to determine the amount of common signal, which was found to decrease as the distance between electrode pairs increased, but was not significantly altered by forearm posture (pronation, neutral, supination). Overall, approximately 40% common signal was detected between adjacent electrode pairs (3 cm apart), dropping to about 10% at 6 cm spacing and 2.5% at 9 cm. The magnitude of common signal approached 50% between adjacent electrode pairs over the extensor muscles, while over 60% was observed between neighbouring sites on the flexor aspect of the forearm. Although flexor and extensor EMG amplitude was similar, less than 2% common signal was present between flexor and extensor electrode pairs during both pinch and grasp tasks. Maximum grip force production was not affected by forearm rotation for pinch, but reduced 18% from neutral (mid-prone) to pronation during grasp (p=0.01). In spite of differences in grip force, mean muscle activity did not vary between the three forearm postures during maximum pinch or grasp trials. While this study improved our knowledge of crosstalk and electrode spacing issues, further examination of forearm EMG is required to improve understanding of muscle loading, EMG properties and motor control during gripping tasks.     

EMG gait data from indwelling electrodes is attenuated over time and changes independent of any experimental effect

The effect of time on the validity of electromyography (EMG) signals from indwelling fine-wire electrodes has not been explored. This is important because experiments using intramuscular electrodes are often long and biochemical and mechanical factors, may impair measurement accuracy over time. Measures over extended periods might therefore be erroneous. Twelve healthy participants (age = 33 ± 8 years) walked for 50 min at a controlled speed. Fine-wire electrodes were inserted into tibialis anterior and a surface EMG sensor attached near the fine-wire insertion site. EMG signals progressively and significantly decreased with time with the fine-wire electrode, but not the surface electrode. For the fine-wire electrode, after 25 min mean amplitude had reduced by 11% (p < 0.001) and after 50 min by 16% (p < 0.001), and peak amplitude reduced 22% at 20 min (p = 0.006) and 37% at 50 min (p < 0.001). Reduced amplitude with indwelling EMG without concurrent changes in surface EMG signal suggests an important inconsistency in data from fine-wire EMG electrodes. Changes in EMG signal will occur over time independent of the experimental condition and this questions their use in experiments of more than 30 min. These results should impact on experimental study design. They also invite reinterpretation of prior literature and sensor innovation to improve measurement performance.     

Laser flow cytometric light scatter and fluorescence pulse width and pulse rise-time sizing of mammalian cells.

In laser flow cytometry, an increasingly popular technique of analytical cytology, quantitative measurements of interest include cell and nuclear diameters. Electronic circuitry for a new cell sizing technique has been developed which measured the time that signal pulses from either fluorescence or light scatter sensors exceed a preset constant fraction of the peak signal amplitude (pulse width) or the time that it takes a signal to rise between constant fractions of the peak signal amplitude on the rising side of the pulse (pulse rise-time). These pulse width or pulse rise-time measurements were related to cell or nuclear diameters and were used in combination to determine nuclear size to cell size ratios. This method of sizing was found to be independent of fluorescent or light-absorbing stain intensity, linearly related to cell or nuclear diameter, and capable of resolving small diameter differences.     

Breathing cardiovascular variability and baroreflex in mechanically ventilated patients

Heart rate and blood pressure variations during spontaneous ventilation are related to the negative airway pressure during inspiration. Inspiratory airway pressure is positive during mechanical ventilation, suggesting that reversal of the normal baroreflex-mediated pattern of variability may occur. We investigated heart rate and blood pressure variability and baroreflex sensitivity in 17 mechanically ventilated patients. ECG (RR intervals), invasive systolic blood pressure (SBP), and respiratory flow signals were recorded. High-frequency (HF) amplitude of RR and SBP time series and HF phase differences between RR, SBP, and ventilatory signals were continuously computed by Complex DeModulation (CDM). Cross-spectral analysis was used to assess the coherence and the gain functions between RR and SBP, yielding baroreflex sensitivity indices. The HF phase difference between SBP and ventilatory signals was nearly constant in all patients with inversion of SBP variability during the ventilator cycle compared with cycling with negative inspiratory pressure to replicate spontaneous breathing. In 12 patients (group 1), the phase difference between RR and ventilatory signals changed over time and the HF-RR amplitude varied. In the remaining five patients (group 2), RR-ventilatory signal phase and HF-RR amplitude showed little change; however, only one of these patients exhibited a RR-ventilatory signal phase difference mimicking the normal pattern of respiratory sinus arrhythmia. Spectral coherence between RR and SBP was lower in the group with phase difference changes. Positive pressure ventilation exerts mainly a mechanical effect on SBP, whereas its influence on HR variability seems more complex, suggesting a role for neural influences.