Gradients and modulation of K(+) channels optimize temporal accuracy in networks of auditory neurons

Accurate timing of action potentials is required for neurons in auditory brainstem nuclei to encode the frequency and phase of incoming sound stimuli. Many such neurons express "high threshold" Kv3-family channels that are required for firing at high rates (> -200 Hz). Kv3 channels are expressed in gradients along the medial-lateral tonotopic axis of the nuclei. Numerical simulations of auditory brainstem neurons were used to calculate the input-output relations of ensembles of 1-50 neurons, stimulated at rates between 100-1500 Hz. Individual neurons with different levels of potassium currents differ in their ability to follow specific rates of stimulation but all perform poorly when the stimulus rate is greater than the maximal firing rate of the neurons. The temporal accuracy of the combined synaptic output of an ensemble is, however, enhanced by the presence of gradients in Kv3 channel levels over that measured when neurons express uniform levels of channels. Surprisingly, at high rates of stimulation, temporal accuracy is also enhanced by the occurrence of random spontaneous activity, such as is normally observed in the absence of sound stimulation. For any pattern of stimulation, however, greatest accuracy is observed when, in the presence of spontaneous activity, the levels of potassium conductance in all of the neurons is adjusted to that found in the subset of neurons that respond better than their neighbors. This optimization of response by adjusting the K(+) conductance occurs for stimulus patterns containing either single and or multiple frequencies in the phase-locking range. The findings suggest that gradients of channel expression are required for normal auditory processing and that changes in levels of potassium currents across the nuclei, by mechanisms such as protein phosphorylation and rapid changes in channel synthesis, adapt the nuclei to the ongoing auditory environment.     

A disjunctive analysis of neonatal approach stimulus prepotence

Preliminary results of a disjunctive procedure developed to ascertain the relative attractiveness for domestic chicks of auditory and visual stimuli are promising. A detailed account of the procedure and initial results is presented. Seventy-two Canadian Athens random bred chicks were tested at 24 or 36 h posthatch. A repetitive tone (4 per sec, 50 msec duration, 500 Hz) served as the auditory stimulus, and a flickering light (3.5 flashes per sec, 0.8 foot candle) served as the visual stimulus. An age-dependent change in the attractiveness of auditory and visual stimuli obtained with the disjunctive procedure. No change in stimulus preference obtained when the stimuli were presented individually.     

No Counterpart of Visual Perceptual Echoes in the Auditory System

It has been previously demonstrated by our group that a visual stimulus made of dynamically changing luminance evokes an echo or reverberation at ∼10 Hz, lasting up to a second. In this study we aimed to reveal whether similar echoes also exist in the auditory modality. A dynamically changing auditory stimulus equivalent to the visual stimulus was designed and employed in two separate series of experiments, and the presence of reverberations was analyzed based on reverse correlations between stimulus sequences and EEG epochs. The first experiment directly compared visual and auditory stimuli: while previous findings of ∼10 Hz visual echoes were verified, no similar echo was found in the auditory modality regardless of frequency. In the second experiment, we tested if auditory sequences would influence the visual echoes when they were congruent or incongruent with the visual sequences. However, the results in that case similarly did not reveal any auditory echoes, nor any change in the characteristics of visual echoes as a function of audio-visual congruence. The negative findings from these experiments suggest that brain oscillations do not equivalently affect early sensory processes in the visual and auditory modalities, and that alpha (8–13 Hz) oscillations play a special role in vision.     

Relationship of transient and steady-state auditory evoked fields

Transient and steady-state auditory evoked fields (AEFs) to brief tone pips were recorded over the left hemisphere at 7 different stimulus rates (0.125–39 Hz) using a 37-channel biomagnetometer. Previous observations of transient auditory gamma band response (GBR) activity were replicated. Similar rate characteristics and equivalent dipole locations supported the suggestion that the steady-state response (SSR) at about 40 Hz represents the summation of successive overlapping (10 Hz) middle latency responses (MLRs). On the other hand, differences in equivalent dipole locations and habituation effects suggest that the magnetically recorded GBR is a separate phenomenon which occurs primarily at low stimulus rates and is unrelated to either the magnetically recorded MRL or SSR.     

Neuromagnetic evidence for early auditory restoration of fundamental pitch

Background

Understanding the time course of how listeners reconstruct a missing fundamental component in an auditory stimulus remains elusive. We report MEG evidence that the missing fundamental component of a complex auditory stimulus is recovered in auditory cortex within 100 ms post stimulus onset.

Methodology

Two outside tones of four-tone complex stimuli were held constant (1200 Hz and 2400 Hz), while two inside tones were systematically modulated (between 1300 Hz and 2300 Hz), such that the restored fundamental (also knows as “virtual pitch”) changed from 100 Hz to 600 Hz. Constructing the auditory stimuli in this manner controls for a number of spectral properties known to modulate the neuromagnetic signal. The tone complex stimuli only diverged on the value of the missing fundamental component.

Principal Findings

We compared the M100 latencies of these tone complexes to the M100 latencies elicited by their respective pure tone (spectral pitch) counterparts. The M100 latencies for the tone complexes matched their pure sinusoid counterparts, while also replicating the M100 temporal latency response curve found in previous studies.

Conclusions

Our findings suggest that listeners are reconstructing the inferred pitch by roughly 100 ms after stimulus onset and are consistent with previous electrophysiological research suggesting that the inferential pitch is perceived in early auditory cortex.     

Preattentive auditory information processing under exposure to the 902 MHz GSM mobile phone electromagnetic field: A mismatch negativity (MMN) study

Previous studies on the effects of the mobile phone electromagnetic field (EMF) on various event‐related potential (ERP) components have yielded inconsistent and even contradictory results, and often failed in replication. The mismatch negativity (MMN) is an auditory ERP component elicited by infrequent (deviant) stimuli differing in some physical features from the repetitive frequent (standard) stimuli in a sound sequence. The MMN provides a sensitive measure for cortical auditory stimulus feature discrimination, regardless of attention and other contaminating factors. In this study, MMN responses to duration, intensity, frequency, and gap changes were recorded in healthy young adults (n = 17), using a multifeature paradigm including several types of auditory change in the same stimulus sequence, while a GSM mobile phone was placed on either ear with the EMF (902 MHz pulsed at 217 Hz; SAR_1g = 1.14 W/kg, SAR_10g = 0.82 W/kg, peak value = 1.21 W/kg, measured with an SAM phantom) on or off. An MMN was elicited by all deviant types, while its amplitude and latency showed no significant differences due to EMF exposure for any deviant types. In the present study, we found no conclusive evidence that acute exposure to GSM mobile phone EMF affects cortical auditory change detection processing reflected by the MMN. Bioelectromagnetics 30:241–248, 2009. © 2009 Wiley‐Liss, Inc.     

Effects of auditory frequency-shifts on zero and below-zero SCR habituation

The purpose of the present experiment was to test a procedure for the measurement of effects of zero- and below-zero habituation (BZH) on responding to frequency shifts in auditory stimuli. The present procedure avoided some of the drawbacks of other procedures, that is, long duration, ambiguity in the definition of BZH, and inadequate control procedures. Two groups received 18 stimulus presentations each; group 1 received first a tone of 1000 Hz 12 times, then 1400 Hz (test stimulus 1) 3 times, and 1850 Hz (test stimulus 2) 3 times. Group 2 received the same stimuli, but 3, 12, and 3 times, respectively. The procedure had the advantages of short duration of the experiment, zero habituation and BZH were operationally defined (as 3 and 12 stimulus presentations, respectively), and there were adequate control conditions since a control group that did not receive stimulus change on the relevant trial was employed. The results showed no effects of stimulus shifts on responding to the test stimuli. Group 2 responded significantly less than group 1 across trials, though, and this may be explained by an inhibitory process elicited by changes in weak stimulation during the habituation process.     

Reduction of information redundancy in the ascending auditory pathway

Information processing by a sensory system is reflected in the changes in stimulus representation along its successive processing stages. We measured information content and stimulus-induced redundancy in the neural responses to a set of natural sounds in three successive stations of the auditory pathway-inferior colliculus (IC), auditory thalamus (MGB), and primary auditory cortex (A1). Information about stimulus identity was somewhat reduced in single A1 and MGB neurons relative to single IC neurons, when information is measured using spike counts, latency, or temporal spiking patterns. However, most of this difference was due to differences in firing rates. On the other hand, IC neurons were substantially more redundant than A1 and MGB neurons. IC redundancy was largely related to frequency selectivity. Redundancy reduction may be a generic organization principle of neural systems, allowing for easier readout of the identity of complex stimuli in A1 relative to IC.     

Electrolocation in the presence of jamming signals: electroreceptor physiology

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

Effects of ganglion blocking agents on post-train facilitation in the rabbit superior cervical ganglion

The effect of ganglion blocking agents, hexamethonium and tubocurarine, on post-train facilitation and ganglionic transmission was studied and compared in isolated superior cervical ganglion of the rabbit, using electrophysiological technique--the conditioning-testing methodology. The preganglionic nerve trunk was stimulated, with either a single unconditioned stimulus (UR)-or a train of conditioning stimuli at 10 or 30 Hz, followed by a post-train test stimulus (PTR). The transmitted postganglionic, compound action potential (PCAP) was recorded following single and trains of stimuli, in the presence and absence of ganglion blocking drugs, hexamethonium (1-100 microM) and tubocurarine (1-100 microM). Hexamethonium and tubocurarine produced concentration-dependent reduction in the amplitude of the transmitted PCAP, increased post-train facilitation values and proportionately reduced those of the subliminal fringe (SF). The mean IC50 values (concentration to produce 50% block of PCAP) of hexamethonium and tubocurarine-induced blockade of the single unconditioned response were 15 +/- 1 microM and 26 +/- 2 microM (n = 6, P less than 0.01) respectively. A dose-ratio (tubocurarine)/hexamethonium) of 1.7 was obtained.     

Correlation between auditory evoked responses in the thalamus and species-specific call characteristics

This evoked potential study of the bullfrog's auditory thalamic area (an auditory responsive region in the posterior dorsal thalamus) shows that complex processing, distinct from that reported in lower auditory regions, occurs in this center. An acoustic stimulus consisting of two tones, one which stimulates either the low-frequency or the mid-frequency sensitive population of auditory nerve fibers from the amphibian papilla and the other the high-frequency sensitive population of fibers from the basilar papilla, evoked a maximal response. The amplitude of the response to the simultaneous stimulation of the two auditory organs was, in some locations, much larger than the linear sum of the responses to the individual tones presented separately. Bimodal spectral stimuli that had relatively long rise-times (greater than or equal to 100 ms) evoked much larger responses than similar sounds with short rise-times. The optimal rise-times were close to those occurring in the bullfrog's mating call. The response was dependent on the waveform periodicity and harmonic content, with a fundamental frequency of 200 Hz producing a larger response than those with fundamentals of 50, 100 or 300 Hz. Six of the natural calls in the bullfrog's vocal repertoire were tested and the mating call and warning call were found to evoke the best responses. Each of these calls stimulate the two auditory organs simultaneously. The evoked response had a long refractory period which could not be altered by lesioning the efferent telencephalic pathways. The type of spectral and temporal information extracted by the auditory thalamic area suggests that this center is involved in processing complex sounds and likely plays an important role in the bullfrog's detection of some of its vocal signals.     

Properties of auditory brainstem responses evoked by intra-operative electrical stimulation of the cochlear nucleus in human subjects

Electrically evoked auditory brainstem response (EABR) testing can aid placement of the stimulating electrodes during surgical implantation of an auditory brainstem implant (ABI). To facilitate efficient testing, this study of EABR properties examined the effects of various stimulating and recording parameters on the magnitude and clarity of the EABRs obtained from 9 successive ABI patients during intra-operative monitoring. Both stimulus polarities elicited EABRs; the response waveforms were similar and no significant differences between the latencies were found. Stimulus-response relationships displayed thresholds and non-linear growth, characteristic of neural activity, and provided a stimulus amplitude that elicited readily detectable EABRs in all subjects. The stimulus rate could be increased without degrading the EABRs, but usually 50 Hz was used with a 10 ms sampling sweep so that muscle responses, which occurred later than EABRs, could be detected. When 3 stimulating electrodes in a line were tested, the pair with the largest separation consistently provided the largest response. A recording filter passband of 10–3000 Hz was useful for attenuating interference signals because there is negligible energy in the EABR at frequencies above 3 kHz, but there is some energy below 100 Hz.     

Functional organization of the electroreceptive midbrain in an elasmobranch (Platyrhinoidis triseriata)

The response properties of 322 single units in the electroreceptive midbrain (lateral mesencephalic nucleus, LMN) of the thornback ray, Platyrhinoidis triseriata, were studied using uniform and local electric fields. Tactile, visual, or auditory stimuli were also presented to test for multimodality. Most LMN electrosensory units (81%) are silent in the absence of stimulation. Those with spontaneous activity fired irregularly at 0.5 to 5 impulses/s, the lower values being more common. Two units had firing rates greater than 10/s. Midbrain electrosensory units are largely phasic, responding with one or a few spikes per stimulus onset or offset or both, but the adaptation characteristics of some neurons are complex. The same neuron can exhibit phasic or phasic-tonic responses, depending upon orientation of the electric field. Tonic units without any initial phasic over-shoot were not recorded. Even the phasic-tonic units adapt to a step stimulus within several seconds. Unit thresholds are generally lower than 0.3 microV/cm, the weakest stimulus delivered, although thresholds as high as 5 microV/cm were recorded, Neuronal responses reach a maximum, with few exceptions, at 100 microV/cm and decrease rapidly at higher intensities. LMN neurons are highly sensitive to stimulus repetition rates: most responded to frequencies of 5 pulses/s or less; none responded to rates greater than 10/s. Three distinct response patterns are recognized. Best frequencies in response to sinusoidal stimuli range from 0.2 Hz (the lowest frequency delivered) to 4 Hz. Responses decrease rapidly at 8 Hz or greater, and no units responded to frequencies greater than 32 Hz. Most LMN neurons have small, well defined excitatory electroreceptive fields (RFs) exhibiting no surround inhibition, at least as detectable by methods employed here. Seventy-eight percent of units recorded had RFs restricted to the ventral surface: of these, 98% were contralateral. The remaining 22% of units had disjunct dorsal and ventral receptive fields. Electrosensory RFs on the ventral surface are somatotopically organized. Anterior, middle, and posterior body surfaces are mapped at the rostral, middle, and caudal levels, respectively, of the contralateral LMN. The lateral, middle, and medial body are mapped at medial, middle, and lateral levels of the nucleus. Moreover, the RFs of all units isolated in a given dorsoventral electrode track are nearly superimposable. About 40% of LMN, measured from the dorsal surface, is devoted to input from ventral electroreceptors located in a small region rostral and lateral to the mouth.(ABSTRACT TRUNCATED AT 400 WORDS)     

P300 in young and elderly subjects: Auditory frequency and intensity effects

Auditory event-related potentials (ERPs) were assessed in young and elderly subjects when stimulus intensity (40 vs. 60 dB SL) and standard/target tone frequency (250/500 Hz and 1000/2000 Hz) were manipulated to study the effects of these variables on the P3(00) and N1, P2 and N2 components. Auditory thresholds for each stimulus type were obtained, and the stimulus intensity was adjusted to effect perceptually equal intensities across conditions for each subject. Younger subjects demonstrated larger P3 amplitudes and shorter latencies than elderly subjects. The low frequency stimuli produced larger P3 amplitude and shorter latencies than the high frequency stimuli. Low intensity stimuli yielded somewhat smaller P3 amplitudes and longer peak latencies than high intensity stimulus tones. Although additional stimulus intensity and frequency effects were obtained for the N1, P2 and N2 components, these generally differed relatively little with subject age. The findings suggest that auditory stimulus parameters contribute to P3 measures, which are different for young compared to elderly subjects.     

Late-night presentation of an auditory stimulus phase delays human circadian rhythms

Although light is considered the primary entrainer of circadian rhythms in humans, nonphotic stimuli, including exercise and melatonin also phase shift the biological clock. Furthermore, in birds and nonhuman mammals, auditory stimuli are effective zeitgebers. This study investigated whether a nonphotic auditory stimulus phase shifts human circadian rhythms. Ten subjects (5 men and 5 women, ages 18-72, mean age +/- SD, 44.7 +/- 21.4 yr) completed two 4-day laboratory sessions in constant dim light (<20 lux). They received two consecutive presentations of either a 2-h auditory or control stimulus from 0100 to 0300 on the second and third nights (presentation order of the stimulus and control was counterbalanced). Core body temperature (CBT) was collected and stored in 2-min bins throughout the study and salivary melatonin was obtained every 30 min from 1900 to 2330 on the baseline and poststimulus/postcontrol nights. Circadian phase of dim light melatonin onset (DLMO) and of CBT minimum, before and after auditory or control presentation was assessed. The auditory stimulus produced significantly larger phase delays of the circadian melatonin (mean +/- SD, -0.89 +/- 0.40 h vs. -0.27 +/- 0.16 h) and CBT (-1.16 +/- 0.69 h vs. -0.44 +/- 0.27 h) rhythms than the control. Phase changes for the two circadian rhythms also positively correlated, indicating direct effects on the biological clock. In addition, the auditory stimulus significantly decreased fatigue compared with the control. This study is the first demonstration of an auditory stimulus phase-shifting circadian rhythms in humans, with shifts similar in size and direction to those of other nonphotic stimuli presented during the early subjective night. This novel stimulus may be a useful countermeasure to facilitate circadian adaptation after transmeridian travel or shift work.     

Characteristics of borderline modulation frequencies in the differentiation of tonal and amplitude-modulated stimuli following the removal of the inferior colliculus in rats

Elaboration of differentiation between sound stimuli was carried out in 15 laboratory rats. After bilateral ablations of auditory inferior colliculi the border frequency of stimulus amplitude modulation was determined for all rats when they still could differentiate between tonal and amplitude-modulated stimuli. Decrease in frequency of modulation by 2 Hz and more from the border frequency caused a complete loss of ability to differentiate. In all rats bilateral inferior colliculi ablations completely disturbed differentiation between tonal and amplitude-modulated signals with modulation frequency below 183-191 Hz (the range of border frequencies). The surgery however did not affect differentiation between tonal and amplitude-modulated signals with the modulation frequencies above 183-191 Hz. The data suggest that the functions of completion of coding of amplitude-modulated stimuli in the auditory system is strictly linked with definite structures.     

Human auditory evoked potentials during natural sleep: The early components

The auditory brain-stem evoked potential (ABEP) was recorded from 9 female subjects during 1 night of natural sleep. Monaural click stimuli were delivered at a rate of either 11, 41 or 81/sec through a hearing-aid device. The intensity was held constant at 70 dB nHL. In other runs, the intensity was lowered to either 50, 30 or 10 dB, the rate of click presentation being held constant at 81/sec. Tympanic temperature was monitored throughout the recording session. The ABEP was unaltered during any stage of sleep regardless of the rate of presentation or stimulus intensity. Distinct peak V responses were recognizable to within 10 dB of the adult threshold in the sleeping subject. It may be concluded that sleep has no virtually on effect on ABEP morphology.     

Seismic and auditory tuning curves from bullfrog saccular and amphibian papular axons

We present seismic and auditory frequency tuning curves of individual bullfrog, Rana catesbeiana, saccular and amphibian papilla axons that responded to both seismic and auditory stimuli. In this study we found: 1) most saccular axons respond well to auditory stimuli with moderate signal strength (50-70 dB SPL) as well as to seismic stimuli; 2) most amphibian papilla axons respond well to seismic stimuli as well as to auditory stimuli, and their seismic sensitivities are comparable to those of saccular axons (responding to sinusoidal stimuli with peak accelerations in the range 0.001 to 0.1 cm/S2); 3) the responses to both seismic and auditory stimuli from both saccule and amphibian papilla are tuned, i.e. the strength of the response varies with the frequency of the stimulus; and this tuning is clearly not the result of second order resonance; 4) in individual axons the tuning properties for seismic stimuli often are not the same as those for auditory stimuli, a fact that may provide clues about how the stimulus signal energy is transferred to the hair cells in each case.     

P300-like potentials in the normal monkey using classical conditioning and an auditory ‘oddball’ paradigm

Endogenous components of evoked potentials resembling P300 in humans were sequentially studied in 3 cynomolgus monkeys (Macaca fascicularis) using an auditory ‘oddball’ paradigm. The two different auditory stimuli were 500 Hz and 4000 Hz tones, designated as the ‘frequent’ and ‘rare’ stimuli, respectively. The probability of ‘rare’ tone presentation was initially 0.2. We further used probabilities of 0.1, 0.3 and 0.5. The ‘rare’ stimulus was reinforced by electrical stimulation, which followed the onset of the high tone by 700 msec. After 3–5 training sessions, a late positive wave was observed following the ‘rare’ tone. The latency of this P300-like signal was 314±16.2 msec, and teh amplitude 23.6±3.14 μV. The amplitude of this potential was modified by changes in stimulus presentation probability and by withholding reinforcement.     

Effects of Auditory Stimuli in the Horizontal Plane on Audiovisual Integration: An Event-Related Potential Study

This article aims to investigate whether auditory stimuli in the horizontal plane, particularly originating from behind the participant, affect audiovisual integration by using behavioral and event-related potential (ERP) measurements. In this study, visual stimuli were presented directly in front of the participants, auditory stimuli were presented at one location in an equidistant horizontal plane at the front (0°, the fixation point), right (90°), back (180°), or left (270°) of the participants, and audiovisual stimuli that include both visual stimuli and auditory stimuli originating from one of the four locations were simultaneously presented. These stimuli were presented randomly with equal probability; during this time, participants were asked to attend to the visual stimulus and respond promptly only to visual target stimuli (a unimodal visual target stimulus and the visual target of the audiovisual stimulus). A significant facilitation of reaction times and hit rates was obtained following audiovisual stimulation, irrespective of whether the auditory stimuli were presented in the front or back of the participant. However, no significant interactions were found between visual stimuli and auditory stimuli from the right or left. Two main ERP components related to audiovisual integration were found: first, auditory stimuli from the front location produced an ERP reaction over the right temporal area and right occipital area at approximately 160–200 milliseconds; second, auditory stimuli from the back produced a reaction over the parietal and occipital areas at approximately 360–400 milliseconds. Our results confirmed that audiovisual integration was also elicited, even though auditory stimuli were presented behind the participant, but no integration occurred when auditory stimuli were presented in the right or left spaces, suggesting that the human brain might be particularly sensitive to information received from behind than both sides.