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Effects of amplitude modulation on the coding of interaural time differences of low-frequency sounds in the inferior colliculus. I. Response properties.

Most sounds in the natural environment are amplitude-modulated (AM). To determine if AM alters the neuronal sensitivity to interaural time differences (ITDs) in low-frequency sounds, we tested neuronal responses to a binaural beat stimulus with and without modulation. We recorded from single units in the inferior colliculus of the unanesthetized rabbit. We primarily used low frequency ( approximately 25 Hz) modulation that was identical at both ears. We found that modulation could enhance, suppress, or not affect the discharge rate. In extreme cases, a neuron that showed no response to the unmodulated binaural beat did so when modulation was added to both ears. At the other extreme, a neuron that showed sensitivity to the unmodulated binaural beat ceased firing with modulation. Modulation could also affect the frequency range of ITD sensitivity, best ITD, and ITD tuning width. Despite these changes in individual neurons, averaging across all neurons, the peak and width of the population ITD function remained unchanged. Because ITD-sensitive neurons also time-locked to the modulation frequency, the location and sound attributes are processed simultaneously by these neurons.

Acoustic Stimulation↗

Neural rate and timing cues for detection and discrimination of amplitude-modulated tones in the awake rabbit inferior colliculus.

Neural responses to amplitude-modulated (AM) tones in the unanesthetized rabbit inferior colliculus (IC) were studied in an effort to establish explicit relationships between physiological and psychophysical measures of temporal envelope processing. Specifically, responses to variations in modulation depth (m) at the cell's best modulation frequency, with and without modulation maskers, were quantified in terms of average rate and synchronization to the envelope over the entire perceptual dynamic range of depths. Statistically significant variations in the metrics were used to define neural AM detection and discrimination thresholds. Synchrony emerged at modulation depths comparable with psychophysical AM detection sensitivities in some neurons, whereas the lowest rate-based neural thresholds could not account for psychoacoustical thresholds. The majority of rate thresholds (85%) were -10 dB or higher (in 20 log m), and 16% of the population exhibited no systematic dependence of average rate on m. Neural thresholds for AM detection did not decrease systematically at higher SPLs (as observed psychophysically): thresholds remained constant or increased with level for most cells tested at multiple sound-pressure levels (SPLs). At depths higher than the rate-based detection threshold, some rate modulation-depth functions were sufficiently steep with respect to the across-trial variability of the rate to predict depth discrimination thresholds as low as 1 dB (comparable with the psychophysics). Synchrony, on the other hand, did not vary systematically with m in many cells at high modulation depths. A simple computational model was extended to reproduce several features of the modulation frequency and depth dependence of both transient and sustained pure-tone responders.

Acoustic Stimulation↗

Sensitivity to amplitude modulated sounds in the anuran auditory nervous system.

Auditory responses were recorded from single units in the eighth nerve and in the midbrain torus semicircularis of the leopard frog (Rana pipiens). Acoustic stimuli included sinusoidally amplitude-modulated (AM) tones and noise, as well as pure tones. Mean spike rates were measured at various rates of AM, and the degree to which a unit's spikes were restricted to a particular phase of the modulation cycle was described by a synchronization coefficient. The firing rate of eighth-nerve fibers was largely independent of the rate of AM over the modulation range 10 to 150 Hz. Further, the general shape of the spike rate vs. AM-rate function was invariant with either depth of modulation or sound-pressure level (SPL). Although virtually all eighth-nerve fibers exhibited significant synchronization to the envelope of AM, the shape of the synchronization function depended on the unit's best-excitatory frequency (BEF). Fibers with highest BEF's, presumed to innervate the basilar papilla, generally showed greater synchronization as the AM rate was increased (up to 100-150 Hz). Fibers tuned to the low-and midfrequency region, which innervate the amphibian papilla, exhibited low-pass synchronization characteristics. As the depth of modulation was reduced, the degree of synchronization of eighth-nerve fibers decreased. For a given depth of modulation an increase in sound level tended to decrease the degree of synchronization, but significant synchronization could still be observed at stimulus intensities at least 65 dB above threshold. On the basis of the spike rate vs. AM-rate functions, the temporal selectivity of single cells in the torus could be characterized by five response types: AM nonselective (spike rate was largely independent of the AM rate); AM high-pass (activity increased as the AM rate was increased); AM low-pass (response was greatest for slow AM rates and decreased at high rates); AM band-suppression (these neurons responded well to low and high AM rates, but responded weakly to intermediate rates); and AM-tuned (spike rate was greatest over a narrow range of modulation rates). In these measurements the depth of modulation was held constant at 100%. The five response categories are not discrete, but rather reflect representative examples along a continuum with regard to temporal selectivity. The temporal selectivity exhibited by toral units in their firing rates was not evident in their AM-synchronization functions.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Differential response properties to amplitude modulated signals in the dorsal nucleus of the lateral lemniscus of the mustache bat and the roles of GABAergic inhibition.

We studied the phase-locking of 89 neurons in the dorsal nucleus of the lateral lemniscus (DNLL) of the mustache bat to sinusoidally amplitude modulated (SAM) signals and the influence that GABAergic inhibition had on their response properties. Response properties were determined with tone bursts at each neuron's best frequency and then with a series of SAM signals that had modulation frequencies ranging from 50-100 to 800 Hz in 100-Hz steps. DNLL neurons were divided into two principal types: sustained neurons (55%), which responded throughout the duration of the tone burst, and onset neurons (45%), which responded only at the beginning of the tone burst. Sustained and onset neurons responded differently to SAM signals. Sustained neurons responded with phase-locked discharges to modulation frequencies < or = 400-800 Hz. In contrast, 70% of the onset neurons phase-locked only to low modulation frequencies of 100-300 Hz, whereas 30% of the onset neurons did not phase-lock to any modulation frequency. Signal intensity differentially affected the phase-locking of sustained and onset neurons. Sustained neurons exhibited tight phase-locking only at low intensities, 10-30 dB above threshold. Onset neurons, in contrast, maintained strong phase-locking even at relatively high intensities. Blocking GABAergic inhibition with bicuculline had different effects on the phase-locking of sustained and onset neurons. In sustained neurons, there was an overall decline in phase-locking at all modulation frequencies. In contrast, 70% of the onset neurons phase-locked to much higher modulation frequencies than they did when inhibition was intact. The other 30% of onset neurons phase-locked to SAM signals, although they fired only with an onset response to the same signals before inhibition was blocked. In both cases, blocking GABAergic inhibition transformed their responses to SAM signals into patterns that were more like those of sustained neurons. We also propose mechanisms that could explain the differential effects of GABAergic inhibition on onset neurons that locked to low modulation frequencies and on onset neurons that did not lock to any SAM signals before inhibition was blocked. The key features of the proposed mechanisms are the absolute latencies and temporal synchrony of the excitatory and inhibitory inputs.

Acoustic Stimulation↗

EPSP amplitude modulation at the rat Ia-alpha motoneuron synapse: effects of GABAB receptor agonists and antagonists.

The object of this study was to examine the relationship between excitatory postsynaptic potential (EPSP) amplitude, posttetanic potentiation, and EPSP amplitude modulation at synapses made by group Ia afferents on motoneurons in the rat. These relationships were evaluated in cells in untreated rats and in cells in rats treated with the gamma-aminobutyric acid-B (GABAB) receptor agonist baclofen and antagonist CGP-35348, which were used to manipulate Ca2+ entry into presynaptic terminals and consequently probability of transmitter release from them. There was no evidence for postsynaptic action of these drugs from measurement of their effects on motoneuron properties. During high-frequency stimulation (32 shock bursts at 167 Hz), EPSP amplitude either decreased (negative modulation) or increased (positive modulation) in response to successive stimuli at different connections. In untreated rats this frequency-dependent amplitude modulation behavior was inversely but weakly correlated with EPSP amplitude measured at low frequency. Intravenous (iv) administration of the GABAB agonist, baclofen, produced a marked and progressive decrease in EPSP amplitude measured at low frequency coincident with a change in frequency-dependent EPSP amplitude modulation toward more positive values (synaptic facilitation). In contrast, an increase in EPSP amplitude occurred after iv administration of the GABAB antagonist CGP-35348 that was accompanied by a negative shift in EPSP amplitude modulation during high-frequency stimulation. The negative shift in EPSP amplitude modulation (synaptic depression) after CGP-35348 application was much smaller than the positive shift induced by baclofen when normalized to the change in EPSP amplitude. Posttetanic potentiation decreased after baclofen but did not increase after CGP-35348. The relationship between modulation and EPSP amplitude was much steeper after GABAB receptor manipulation in either direction than that observed in the population of motoneurons in untreated preparations. This suggests that in the rat differences in probability of release play at most a small role in determining EPSP amplitude across the motoneuron pool.

Afferent Pathways↗

GABAergic and glycinergic inhibition sharpens tuning for frequency modulations in the inferior colliculus of the big brown bat.

Discrimination of amplitude and frequency modulated sounds is an important task of auditory processing. Experiments have shown that tuning of neurons to sinusoidally frequency- and amplitude-modulated (SFM and SAM, respectively) sounds becomes successively narrower going from lower to higher auditory brain stem nuclei. In the inferior colliculus (IC), many neurons are sharply tuned to the modulation frequency of SFM sounds. The purpose of this study was to determine whether GABAergic or glycinergic inhibition is involved in shaping the tuning for the modulation frequency of SFM sounds in IC neurons of the big brown bat (Eptesicus fuscus). We recorded the response of 56 single units in the central nucleus of the IC to SFM stimuli before and during the application of the gamma-aminobutyric acid-A (GABAA) receptor antagonist bicuculline or the glycine receptor antagonist strychnine. To evaluate tuning to the modulation frequency, the normalized spike count (normalized according to the maximal response for each condition tested) was plotted versus the modulation frequency and the upper and lower 50% cutoff points were determined. Bicuculline increased the upper cutoff in 46% of the neurons by >/=25%. The lower cutoff decreased in 48% of the neurons tested. In some neurons (approximately 30%), a sharpening of the tuning by bicuculline was observed. Strychnine induced an increase of the upper cutoff in almost half of the neurons. Compared with bicuculline these changes were smaller. The lower cutoff decreased in 50% of the neurons with strychnine. The synchronization coefficient (SC) was calculated and compared for three modulation frequencies (50, 100, and 200 Hz) between predrug and drug condition. For all neurons, synchronization decreased (n = 36) or did not change (n = 26) during drug application. This was mainly an effect of the prolonged discharge in response to each cycle. Under predrug conditions, many neurons exhibited selectivity to the direction of the FM, hence they only responded once to each cycle. In a minority of neurons, direction selectivity was abolished by drug application. The main finding was that neuronal inhibition sharpens tuning to the modulation frequency in the majority of neurons. In general, changes induced by bicuculline or strychnine were comparable.

Acoustic Stimulation↗

Temporal dynamics of convergent modulation at a crustacean neuromuscular junction.

At least 10 different substances modulate the amplitude of nerve-evoked contractions of the gastric mill 4 (gm4) muscle of the crab, Cancer borealis. Serotonin, dopamine, octopamine, proctolin, red pigment concentrating hormone, crustacean cardioactive peptide, TNRNFLRFamide, and SDRNFLRFamide increased and -allatostatin-3 and histamine decreased the amplitude of nerve-evoked contractions. Modulator efficacy was frequency dependent; TNRNFLRFamide, proctolin, and allatostatin-3 were more effective when the motor neuron was stimulated at 10 Hz than at 40 Hz, whereas the reverse was true for dopamine and serotonin. The modulators that were most effective at high stimulus frequencies produced a significant decrease in muscle relaxation time; those that were most effective at low stimulus frequencies produced modest increases in relaxation time. Thus modulator actions that appear redundant when examined only at one stimulus frequency are differentiated when a range of stimulus dynamics is studied. The effects of TNRNFLRFamide, serotonin, proctolin, dopamine, and -allatostatin-3 on the amplitude and facilitation of nerve-evoked excitatory junctional potentials (EJPs) in the gm4 and gastric mill 6 (gm6) muscles were compared. The EJPs in gm4 have a large initial amplitude and show relatively little facilitation, whereas the EJPs in gm6 have a small initial amplitude and show considerable facilitation. Modulators that enhanced contractions also enhanced EJP amplitude; -allatostatin-3 reduced EJP amplitude. The effects of these modulators on EJP amplitude were modest and showed no significant frequency dependence. This suggests that the frequency dependence of modulator action on contraction results from effects on excitation-contraction coupling. The modulators affected facilitation at these junctions in a manner consistent with a change in release probability. They produced a change in facilitation that is inversely related to their action on EJP amplitude.

Animals↗

Computational model of the serotonergic modulation of sensory neurons in Aplysia.

Serotonergic modulation of the sensory neurons that mediate the gill- and tail-withdrawal reflexes of Aplysia is a useful model system for studies of neuronal plasticity that contributes to learning and memory. The effects of serotonin (5-HT) are mediated, in part, via two protein kinases (protein kinase A, PKA, and protein kinase C, PKC), which in turn, modulate at least four membrane currents, including a S ("serotonin-sensitive") K(+) current (I(K, S)), a steeply voltage-dependent K(+) current (I(K-V)), a slow component of the Ca(2+)-activated K(+) current (I(K,Ca-S)), and a L-type Ca(2+) current (I(Ca-L)). The present study investigated how the modulation of these currents altered the spike duration and excitability of sensory neurons and examined the relative contributions of PKA- and PKC-mediated effects to the actions of 5-HT. A Hodgkin-Huxley type model was developed that described the ionic conductances in the somata of sensory neurons. The descriptions of these currents and their modulation were based largely on voltage-clamp data from sensory neurons. Simulations were preformed with the program SNNAP (Simulator for Neural Networks and Action Potentials). The model was sufficient to replicate empirical data that describes the membrane currents, action potential waveform and excitability as well as their modulation by application of 5-HT, increased levels of adenosine cyclic monophosphate or application of active phorbol esters. In the model, modulation of I(K-V) by PKC played a dominate role in 5-HT-induced spike broadening, whereas the concurrent modulation of I(K,S) and I(K,Ca-S) by PKA primarily accounted for 5-HT-induced increases in excitability. Finally, simulations indicated that a PKC-induced increase in excitability resulted from decreases of I(K,S) and I(K,Ca-S), which was likely the indirect result of cross-talk between the PKC and PKA systems. The results provide several predictions that warrant additional experimental investigation and illustrate the importance of considering indirect as well as direct effects of modulatory agents on the modulation of membrane currents.

Action Potentials↗

Angiotensin-mediated renin suppression is altered in non-modulating hypertension.

Plasma renin activity (PRA) suppression by angiotensin II (Ang II), the "short feedback loop," has been shown to be blunted in essential hypertension. Non-modulators comprise 50% of normal- and high-renin essential hypertensive individuals and are defined by their abnormal adrenal and renal responses to infused Ang II. This study sought to determine if the abnormal Ang II suppression of PRA in essential hypertension is specific to non-modulators. We measured PRA suppression during infusion of Ang II in 22 normotensive and 32 hypertensive (16 modulators and 16 non-modulators) subjects. A significant correlation between basal PRA and the fall in PRA in response to Ang II infusion was seen in the normotensives (r = -0.934, p less than 0.001) and modulators (r = -0.831, p less than 0.001) but not in the non-modulators (r = -0.275, p = 0.304). However, after 3 days of converting enzyme inhibition therapy, the correlation between basal PRA and decrement in PRA was significant in the non-modulators (r = -0.762, p less than 0.01). Thus, the abnormal PRA suppression documented previously in hypertensive individuals is a defect specific to non-modulators and is corrected with converting enzyme inhibition therapy.

Adult↗

Mechanisms of frequency and amplitude modulation in ring dove song.

Birdsong assumes its complex and specific forms by the modulation of phonation in frequency and time domains. The organization of control mechanisms and intrinsic properties causing such modulation have been studied in songbirds but much less so in non-songbirds, the songs of which are often regarded as relatively simple. We examined mechanisms of frequency and amplitude modulation of phonation in ring doves Streptopelia risoria, which are non-songbirds. Spontaneous coo vocalizations were recorded together with concurrent pressure patterns in two different air sacs and air flow rate in the trachea. The results show that amplitude modulation is the result of the cyclic opening and closure of a valve instead of fluctuations in driving pressure, as is the current explanation. Frequency modulation is more complex than previously recognized and consists of gradual, continuous time-frequency patterns, punctuated by instantaneous frequency jumps. Gradual frequency modulation patterns correspond to pressure variation in the interclavicular air sac but not to pressure variation in the cranial thoracic air sac or air flow rate variation in the trachea. The cause of abrupt jumps in frequency has not been identified but can be explained on the basis of intrinsic properties of the vocal organ. Air sac pressure variation as a mechanism for frequency modulation contrasts with the specialized syringeal musculature of songbirds and may explain why the fundamental frequency in non-songbird vocalizations is generally modulated within a limited frequency range.

Air Sacs↗

Luminance and chromatic modulation sensitivity of macaque ganglion cells and human observers.

We measured the sensitivity of macaque ganglion cells to luminance and chromatic sinusoidal modulation. Phasic ganglion cells of the magnocellular pathway (M-pathway) were the more sensitive to luminance modulation, and tonic ganglion cells of the parvocellular pathway (P-pathway) were more sensitive to chromatic modulation. With decreasing retinal illuminance, phasic ganglion cells' temporal sensitivity to luminance modulation changed in a manner that paralleled psychophysical data. The same was true for tonic cells and chromatic modulation. Taken together, the data suggest strongly that the cells of the M-pathway form the physiological substrate for detection of luminance modulation and the cells of the P-pathway the substrate for detection of chromatic modulation. However, at high light levels, intrusion of a so-called luminance mechanism near 10 Hz in psychophysical detection of chromatic modulation is probably due to responses in the M-pathway, arising primarily from a nonlinearity of cone summation. Both phasic and tonic ganglion cells responded to frequencies higher than can be psychophysically detected. This suggests that central mechanisms, acting as low-pass filters, modify these cells' signals, though the corner frequency is lower for the P-pathway than for the M-pathway. For both cell types, the response phase at different frequencies was consistent with the cells' description as linear filters with a fixed time delay.

Animals↗

Molecular determinants for modulation of persistent sodium current by G-protein betagamma subunits.

Voltage-gated sodium channels are responsible for the upstroke of the action potential in most excitable cells, and their fast inactivation is essential for controlling electrical signaling. In addition, a noninactivating, persistent component of sodium current, I(NaP), has been implicated in integrative functions of neurons including threshold for firing, neuronal bursting, and signal integration. G-protein betagamma subunits increase I(NaP), but the sodium channel subtypes that conduct I(NaP) and the target site(s) on the sodium channel molecule required for modulation by Gbetagamma are poorly defined. Here, we show that I(NaP) conducted by Na(v)1.1 and Na(v)1.2 channels (Na(v)1.1 > Na(v)1.2) is modulated by Gbetagamma; Na(v)1.4 and Na(v)1.5 channels produce smaller I(NaP) that is not regulated by Gbetagamma. These qualitative differences in modulation by Gbetagamma are determined by the transmembrane body of the sodium channels rather than their cytoplasmic C-terminal domains, which have been implicated previously in modulation by Gbetagamma. However, the C-terminal domains determine the quantitative extent of modulation of Na(v)1.2 channels by Gbetagamma. Studies of chimeric and truncated Na(v)1.2 channels identify molecular determinants that affect modulation of I(NaP) located between amino acid residue 1890 and the C terminus at residue 2005. The last 28 amino acid residues of the C terminus are sufficient to support modulation by Gbetagamma when attached to the proximal C-terminal domain. Our results further define the sodium channel subtypes that generate I(NaP) and identify crucial molecular determinants in the C-terminal domain required for modulation by Gbetagamma when attached to the transmembrane body of a responsive sodium channel.

Amino Acid Sequence↗

Chemical prevention of colony foundation by Cryptotermes brevis (Isoptera: Kalotermitidae) in attic modules.

Disodium octaborate tetrahydrate (DOT) dust, DOT aqueous solution, imidacloprid dust, and amorphous silica gel dust with synergized 1% pyrethrins were applied on wood surfaces to simulated attic modules. Modules (30 by 30 cm) with and without fiberglass insulation were exposed to dispersal flights of Cryptotermes brevis (Walker) in May and June of 1998 and 1999. Six months after flights, modules were disassembled and inspected for nuptial chamber location and contents. During both years, air and water control treatments contained 22.2+/-9.94 (mean +/- SD) nuptial chambers, 7.5+/-5.7 live imagos, and 2.0+/-1.4 chambers with brood. This survivorship indicated that the attic modules performed well as a colonizing platform for C. brevis. C. brevis dealates preferred constructing nuptial chambers in the crevices at the bases or tops of the modules instead of internal crevices. Modules treated in 1998 and 1999 with DOT or silica dusts contained no live termites, whereas zero of five modules treated with imidacloprid dust in 1998 and two of 20 modules treated with imidacloprid dust in 1999 contained single live incipient colonies. In 1998, 15% DOT solution, applied as a postconstruction treatment, yielded significantly fewer chambers and live termites than controls, but was not as effective as dusts in preventing successful colonization. In 1999, the DOT solution, applied as a construction-phase treatment, was equally as effective in preventing colonization as the dust treatments during that year. Results indicate that dust formulations of DOT, silica gel, and imidacloprid can be used to prevent drywood termite colonization in existing building voids and attics. Where the entire wood framing is exposed to treatment, such as during building construction, aqueous DOT solution can be equally effective as dusts in preventing colonization by C. brevis.

Animals↗

Biological effects of radiofrequency fields: does modulation matter?

This commentary considers modulation as a factor of potential biological importance in assessment of risk of radiofrequency (RF) energy emitted by communications systems and other technologies. Modulation introduces a spread of frequencies into a carrier waveform, but in nearly all cases this spread is small compared to the frequency of the carrier. Consequently, any nonthermal (field-dependent) biological effects related to modulation must result from interaction mechanisms that are fast enough to produce a response at radiofrequencies. Despite considerable speculation, no such mechanisms have been established. While a variety of modulation-dependent biological effects of RF energy have been reported, few such effects have been independently confirmed. Some widely discussed effects, for example a reported modulation-dependent effect of RF fields on the efflux of calcium from brain tissue, remain controversial with no established biological significance. The lack of understanding of the mechanisms underlying such effects prevents any assessment of their significance for communications signals with complex modulation characteristics. Future research should be directed at confirmation and mechanistic understanding of reported biological effects related to modulation. While modulation should be considered in the design of risk studies involving communications-type signals, it should not compromise other aspects of good study design, such as maintaining adequate statistical power and identifying dose-response relationships.

Calcium↗

Emission characteristics of Cu in Dc voltage modulation applied glow discharge optical emission spectrometry.

In order to obtain the depth profile of a thin film, we investigated the emission characteristics of a voltage modulation glow discharge to optimize the modulation parameters (modulation voltage, offset voltage, and modulation frequency). In this study, a phase-sensitive detection method with a lock-in amplifier to the modulation technique led to a higher sensitivity and a larger signal-to-noise ratio in the emission analysis compared to the normal dc amplification method. Upon increasing the maximum voltage, the emission intensity of the Cu atomic line (CuI 239.34 nm) increased linearly at a modulation voltage of 400 V and an offset voltage of 300 V. On the other hand, the emission intensity was gradually reduced when a modulation frequency increased. It is advantageous for surface analysis that the voltage modulation technique gives a lower sputtering rate rather than the conventional dc discharge.

Journal Article↗

MEGO: gene functional module expression based on gene ontology.

Existing analysis tools to study the collective properties of gene functional modules cannot return highly homogeneous modules and do not provide quantitative measures of module activity level. By partitioning genes according to multiple gene functional categorization principles and summarizing gene expression values into module expression values, MEGO (module expression based on gene ontology), a standalone microarray data analysis program, is able to extract highly activated gene functional modules that are of much interest to microarray experimenters. With multiple functional categorization principles simultaneously introduced in MEGO, the partition of genes is more delicate, and the collective property of a group of genes is sharpened and easier to capture. The quantitative measures of module activity levels returned by MEGO give users a quick impression of the direction and degree of module regulation. MEGO efficiently determines the answers to frequently asked questions, such as which functional classes have been induced or repressed under a specific experiment and to which levels these functional classes have been affected. MEGO is available free of charge for academic use and may be downloaded from http://www.dxy.cn/mego/MEGOInstall.EXE. Supplementary information can be found on the authors' web page at http://www.dxy.cn/mego/ and at the BioTechniques' web site at http://www. BioTechniques.com/February2005/TuSupplementary.html.

Algorithms↗

Modulation threshold functions for chronically impaired Menière patients.

Detection thresholds for sinusoidally amplitude-modulated broad-band noise were measured as a function of modulation frequency for both ears of 6 chronic Menière patients who suffered unilateral hearing impairments. Modulation thresholds were measured with an adaptive cued-standard forced-choice psychophysical method. Better-ear modulation thresholds were similar to normative data previously reported [Formby, C.: J. acoust. Soc.Am. 78:70-77, 1985], whereas 5 of the 6 patients exhibited deficits in modulation detection with their poorer ears. Modulation thresholds averaged across the patients' poorer ears were similar to the normative thresholds through 60-100 Hz; at higher modulation frequencies, sensitivity declined at approximately twice the normal attenuation rate (i.e., 6 vs. 3 dB per octave). The poorer-ear data can be described by the mathematical representation for a simple low-pass filter with a cutoff frequency of 60 Hz. This pattern of the Menière modulation thresholds is consistent with broadened peripheral tuning due to hydrops.

Adult↗

[Auditory steady-state response to sinusoidally amplitude-modulated tones. Second report: investigation of response in the sleeping state].

The auditory steady state response (SSR) elicited by a sinusoidally amplitude-modulated (SAM) tone may be an important technique in objective audiometry, which provides frequency-specific information. This response is affected the state of arousal of the subject. An effect of sleep on the amplitude of this response has been observed by a number of previous investigators. Since young children must usually be sedated to carry out evoked potential testing, the effects of sleep on this response are a crucial factor limiting clinical applicability. The basic characteristics of SSR elicited by a 500 Hz SAM tone were studied in 21 adults with normal hearing during sleep. The responses while sleeping were compared with the responses in the awake state. The responses to 2048 stimuli were averaged with the analysis time set at 102.4 ms, and responses were judged by passing the waveforms through a 20-100 Hz digital filter. The same technique was performed in 25 infants (2 months-2 years 9 months) evaluated as having normal hearing. The results obtained are summarized as follows: The amplitude of the response while sleeping was approximately half the amplitude in the awake state. Responses with a modulation frequency of 70Hz, however, showed a persistent decrease in 71.0%, suggesting a trend different from that seen at a modulation frequency of less than 70Hz. The effect of modulation frequency on response amplitude was almost the same for the sleeping state and the awake state. Even during sleep, the response amplitude was maximal at modulation frequencies of 20, 30 and 40Hz, and there was no significant difference between the values at 20Hz -40Hz. The latency of the response was measured by the Diamond method, and the results showed that latency is significantly prolonged while sleeping (p < 0.01). The response threshold at the 500Hz SAM tone (modulation rate 40Hz, modulation depth 90%) was 8.5dBnHL when awake and 13.9dBnHL during sleep. The rise in threshold as a result of sleep was no more than about 5dB. In infants, the response amplitude at a modulation frequency of 20Hz was maximal, and amplitude at the 40Hz was significantly smaller (p < 0.01). Latency measured by the Diamond method in infants was 12.8ms. Thus, a difference from the response in adults was confirmed. Comparisons were made between the responses of infants under 1 year of age and infants over 1 year of age to confirm changes in response according to age.

Adolescent↗