Infrasound tones at sensation threshold level elicit measurable Frequency-Following responses.

Even barely detectable levels of infrasound are often reported to cause annoyance and complaints. We carefully measured the individual sensation threshold of a pure tone and recorded immediately after the brain's frequency-following response (FFR) at this intensity using the same stimulator. In contrast to 87-Hz tones, 8-Hz tones elicit an FFR already at sensation threshold. Control stimuli with trains of 1-kHz tone pips having the repetition rate of the infrasound tone frequency and sensation threshold intensities evoked no significant FFR. Thus, slow periodicity, causing synchronous activation of auditory nuclei, is not explaining the FFR to low-level infrasound alone.

Dependence of binaural gain for infrasound on interaural phase difference.

Increasing complaints about infrasound have generated interest in understanding its perception, including binaural effects. This study investigated the level difference between monaural and binaural presentation required for detection and equal loudness (binaural gain) for pure tones with frequencies of 8, 32, and 400 Hz and an 8 Hz sinusoidally amplitude-modulated tone with diotic 400 Hz carrier. Monaural stimuli were compared to binaural stimuli with interaural phase differences (IPDs) of 0°, 90°, and 180° in two experiments: absolute threshold measurements and loudness matching at 40 phons. The latter was repeated with transposed tones (400 Hz carrier multiplied by a half-wave-rectified 8 Hz sinusoid). When expressed as differences in sound pressure level, similar binaural gain was found across all stimulus types under the diotic condition. Confirming previous studies, the gain was larger at supra-threshold levels (40 phons) than at threshold. However, when the loudness-matching results were expressed as binaural gain with respect to the loudness level, they became 17.5, 11.2, and 5.8 phons for the 8, 32, and 400 Hz stimuli, respectively. Results for the 8 Hz pure tone and the transposed stimulus were IPD dependent.

Dependency of threshold and loudness on sound duration at low and infrasonic frequencies.

Many environmental sounds contain significant energy in the infrasonic and low-frequency (ISLF) ranges that have been associated with cases of annoyance and noise complaints. This study assessed the effect of sound duration on audibility and loudness of ISLF sounds. A first experiment evaluated detection thresholds for tones of 4, 16, and 32 Hz with durations up to 4000 ms. Furthermore, equal-loudness-level contours (ELCs) were obtained as function of duration up to 2000 ms. Tones of 1000 Hz were also included here. Results displayed the known pattern of general sound level decrease with increasing duration up to several hundred milliseconds. ELCs stabilized slightly earlier than thresholds, but after 1000 ms, levels remained roughly constant for both measures except for 4-Hz tones, where the decrease continued up to the longest durations tested. As 4-Hz cycles are perceptually resolved as separate pressure pulses, the authors hypothesized their duration dependence would resemble that of pulse trains. Hence, a second experiment evaluated pulse-train thresholds (1000-Hz carrier) for durations up to 4000 ms. For both pulse repetition rates of 4 and 32 Hz, threshold stabilized after 1000 ms as for tones ≥16 Hz, suggesting the continuing threshold decrease for a 4-Hz tone is specific to infrasound.

A common periodic representation of interaural time differences in mammalian cortex.

Binaural hearing, the ability to detect small differences in the timing and level of sounds at the two ears, underpins the ability to localize sound sources along the horizontal plane, and is important for decoding complex spatial listening environments into separate objects - a critical factor in 'cocktail-party listening'. For human listeners, the most important spatial cue is the interaural time difference (ITD). Despite many decades of neurophysiological investigations of ITD sensitivity in small mammals, and computational models aimed at accounting for human perception, a lack of concordance between these studies has hampered our understanding of how the human brain represents and processes ITDs. Further, neural coding of spatial cues might depend on factors such as head-size or hearing range, which differ considerably between humans and commonly used experimental animals. Here, using magnetoencephalography (MEG) in human listeners, and electro-corticography (ECoG) recordings in guinea pig-a small mammal representative of a range of animals in which ITD coding has been assessed at the level of single-neuron recordings-we tested whether processing of ITDs in human auditory cortex accords with a frequency-dependent periodic code of ITD reported in small mammals, or whether alternative or additional processing stages implemented in psychoacoustic models of human binaural hearing must be assumed. Our data were well accounted for by a model consisting of periodically tuned ITD-detectors, and were highly consistent across the two species. The results suggest that the representation of ITD in human auditory cortex is similar to that found in other mammalian species, a representation in which neural responses to ITD are determined by phase differences relative to sound frequency rather than, for instance, the range of ITDs permitted by head size or the absolute magnitude or direction of ITD.

Influence of envelope waveform on ITD sensitivity of neurons in the auditory midbrain.

Interaural time differences (ITDs) conveyed by the modulated envelopes of high-frequency sounds can serve as a cue for localizing a sound source. Klein-Hennig et al. (J Acoust Soc Am 129: 3856, 2011) demonstrated the envelope attack (the rate at which stimulus energy in the envelope increases) and the duration of the pause (the interval between successive envelope pulses) as important factors affecting sensitivity to envelope ITDs in human listeners. Modulated sounds with rapid attacks and long pauses produce the lowest ITD discrimination thresholds. The duration of the envelope's sustained component (sustain) and the rate at which stimulus energy falls at the offset of the envelope (decay) are only minor factors. We assessed the responses of 71 single neurons, recorded from the midbrains of 15 urethane-anesthetized tri-colored guinea pigs, to envelope shapes in which the four envelope components, i.e., attack, sustain, decay, and pause, were systematically varied. We confirmed the importance of the attack and pause components in generating ITD-sensitive responses. Analysis of neural firing rates demonstrated more neurons (49/71) show ITD sensitivity in response to "damped" stimuli (fast attack and slow decay) compared with "ramped" stimuli (slow attack and fast decay) (14/71). Furthermore, the lowest threshold for the damped stimulus (91 µs) was lower by a factor of 4 than that for the temporally reversed ramped envelope shape (407 µs). The data confirm the importance of fast attacks and optimal pause durations in generating sensitivity to ITDs conveyed in the modulated envelopes of high-frequency sounds and are incompatible with models of ITD processing based on the integration of sound energy over time.NEW & NOTEWORTHY Using single-neuron electrophysiology, we show that the precise shape of a sound's "energy envelope" is a critical factor in determining how well midbrain neurons are able to convey information about auditory spatial cues. Consistent with human behavioral performance, sounds with rapidly rising energy and relatively long intervals between energy bursts are best at conveying spatial information. The data suggest specific sound energy patterns that might best be applied to hearing devices to aid spatial listening.

Objective Measures of Neural Processing of Interaural Time Differences.

We assessed neural sensitivity to interaural time differences (ITDs) conveyed in the temporal fine structure (TFS) of low-frequency sounds and ITDs conveyed in the temporal envelope of amplitude-modulated (AM'ed) high-frequency sounds. Using electroencephalography (EEG), we recorded brain activity to sounds in which the interaural phase difference (IPD) of the TFS (or the modulated temporal envelope) was repeatedly switched between leading in one ear or the other. When the amplitude of the tones is modulated equally in the two ears at 41 Hz, the interaural phase modulation (IPM) evokes an IPM following-response (IPM-FR) in the EEG signal. For low-frequency signals, IPM-FRs were reliably obtained, and largest for an IPM rate of 6.8 Hz and when IPD switches (around 0°) were in the range 45-90°. IPDs conveyed in envelope of high-frequency tones also generated IPM-FRs; response maxima occurred for IPDs switched between 0° and 180° IPD. This is consistent with the interpretation that distinct binaural mechanisms generate the IPM-FR at low and high frequencies, and with the reported physiological responses of medial superior olive (MSO) and lateral superior olive (LSO) neurons in other mammals. Low-frequency binaural neurons in the MSO are considered maximally activated by IPDs in the range 45-90°, consistent with their reception of excitatory inputs from both ears. High-frequency neurons in the LSO receive excitatory and inhibitory input from the two ears receptively--as such maximum activity occurs when the sounds at the two ears are presented out of phase.

Emphasis of spatial cues in the temporal fine structure during the rising segments of amplitude-modulated sounds.

The ability to locate the direction of a target sound in a background of competing sources is critical to the survival of many species and important for human communication. Nevertheless, brain mechanisms that provide for such accurate localization abilities remain poorly understood. In particular, it remains unclear how the auditory brain is able to extract reliable spatial information directly from the source when competing sounds and reflections dominate all but the earliest moments of the sound wave reaching each ear. We developed a stimulus mimicking the mutual relationship of sound amplitude and binaural cues, characteristic to reverberant speech. This stimulus, named amplitude modulated binaural beat, allows for a parametric and isolated change of modulation frequency and phase relations. Employing magnetoencephalography and psychoacoustics it is demonstrated that the auditory brain uses binaural information in the stimulus fine structure only during the rising portion of each modulation cycle, rendering spatial information recoverable in an otherwise unlocalizable sound. The data suggest that amplitude modulation provides a means of "glimpsing" low-frequency spatial cues in a manner that benefits listening in noisy or reverberant environments.

The effect of the helicotrema on low-frequency loudness perception.

Below approximately 40 Hz, the cochlear travelling wave reaches the apex, and differential pressure is shunted through the helicotrema, reducing hearing sensitivity. Just above this corner frequency, a resonance feature is often observed in objectively measured middle-ear-transfer functions (METFs). This study inquires whether overall and fine structure characteristics of the METF are also perceptually evident. Equal-loudness-level contours (ELCs) were measured between 20 and 160 Hz for 14 subjects in a purpose-built test chamber. In addition, the inverse shapes of their METFs were obtained by adjusting the intensity of a low-frequency suppressor tone to maintain an equal suppression depth of otoacoustic emissions for various suppressor tone frequencies (20-250 Hz). For 11 subjects, the METFs showed a resonance. Six of them had coinciding features in both ears, and also in their ELC. For two subjects only the right-ear METF was obtainable, and in one case it was consistent with the ELC. One other subject showed a consistent lack of the feature in their ELC and in both METFs. Although three subjects displayed clear inconsistencies between both measures, the similarity between inverse METF and ELC for most subjects shows that the helicotrema has a marked impact on low-frequency sound perception.

Emphasis of spatial cues in the temporal fine structure during the rising segments of amplitude-modulated sounds II: single-neuron recordings.

Recently, with the use of an amplitude-modulated binaural beat (AMBB), in which sound amplitude and interaural-phase difference (IPD) were modulated with a fixed mutual relationship (Dietz et al. 2013b), we demonstrated that the human auditory system uses interaural timing differences in the temporal fine structure of modulated sounds only during the rising portion of each modulation cycle. However, the degree to which peripheral or central mechanisms contribute to the observed strong dominance of the rising slope remains to be determined. Here, by recording responses of single neurons in the medial superior olive (MSO) of anesthetized gerbils and in the inferior colliculus (IC) of anesthetized guinea pigs to AMBBs, we report a correlation between the position within the amplitude-modulation (AM) cycle generating the maximum response rate and the position at which the instantaneous IPD dominates the total neural response. The IPD during the rising segment dominates the total response in 78% of MSO neurons and 69% of IC neurons, with responses of the remaining neurons predominantly coding the IPD around the modulation maximum. The observed diversity of dominance regions within the AM cycle, especially in the IC, and its comparison with the human behavioral data suggest that only the subpopulation of neurons with rising slope dominance codes the sound-source location in complex listening conditions. A comparison of two models to account for the data suggests that emphasis on IPDs during the rising slope of the AM cycle depends on adaptation processes occurring before binaural interaction.

A model that predicts the binaural advantage to speech intelligibility from the mixed target and interferer signals.

A model is presented that predicts the binaural advantage to speech intelligibility by analyzing the right and left recordings at the two ears containing mixed target and interferer signals. This auditory-inspired model implements an equalization-cancellation stage to predict the binaural unmasking (BU) component, in conjunction with a modulation-frequency estimation block to estimate the "better ear" effect (BE) component of the binaural advantage. The model's performance was compared to experimental data obtained under anechoic and reverberant conditions using a single speech-shaped noise interferer paradigm. The internal BU and BE components were compared to those of the speech intelligibility model recently proposed by Lavandier et al. [J. Acoust. Soc. Am. 131, 218-231 (2012)], which requires separate inputs for target and interferer. The data indicate that the proposed model provides comparably good predictions from a mixed-signals input under both anechoic and reverberant conditions.

The influence of the envelope waveform on binaural tuning of neurons in the inferior colliculus and its relation to binaural perception.

Recently, Klein-Hennig et al. (J Acoust Soc Am 129:3856­3872, 2011) suggested a design for envelope waveforms that allows for independent setting of the duration of the four segments of an envelope cycle ­ pause, attack, sustain, and decay. These authors conducted psychoacoustic experiments to determine the threshold interaural time differences (ITDs) for different waveforms and revealed that a steep attack flank and at least 4 ms of pause duration prior to the attack are optimal for discrimination performance, whilst sustained and decay durations were of only minor influence. The current study tests the sharpness of rate-ITD-functions recorded in the inferior colliculus of guinea pigs in response to a similar set of waveforms, examining their relationship to the psychoacoustic data. Particular focus is applied to temporally asymmetric envelope waveforms: a long 15-ms attack and a short 1.5-ms decay envelope and the temporally inverted envelope with a short 1.5-ms attack and a long 15-ms decay.

A simple electrical lumped-element model simulates intra-cochlear sound pressures and cochlear impedance below 2 kHz.

Low-frequency sounds displace large parts of the basilar membrane (BM) and can have a modulating and possibly disturbing effect on hearing at other frequencies. A better understanding of the transfer of such sounds onto the BM is therefore desirable. Lumped-element models have previously been employed to determine the low-frequency acoustic properties of the cochlea. Although helpful in illustrating schematically the role of the helicotrema, BM compliance, and the round window on low-frequency hearing, these models, when applied quantitatively, have not been able to explain experimental data in detail. Building on these models, an extended electrical analog requires just 13 lumped elements to capture, in surprising detail, the physiologically determined frequency-dependence of intra-cochlear pressure and cochlear impedance between 10 Hz and 2 kHz. The model's verification is based on data from cat, guinea pig, and humans, who differ principally in their low-frequency cochlear acoustics. The modeling data suggest that damping within the helicotrema plays a less prominent role than previously assumed. A resonance feature, which is often observed experimentally near 150 Hz in these animals and near 50 Hz in humans, is presumably a phenomenon local to the apex and not the result of a standing wave between stapes and helicotrema.

The Spectral Extent of Phasic Suppression of Loudness and Distortion-Product Otoacoustic Emissions by Infrasound and Low-Frequency Tones.

We investigated the effect of a biasing tone close to 5, 15, or 30 Hz on the response to higher-frequency probe tones, behaviorally, and by measuring distortion-product otoacoustic emissions (DPOAEs). The amplitude of the biasing tone was adjusted for criterion suppression of cubic DPOAE elicited by probe tones presented between 0.7 and 8 kHz, or criterion loudness suppression of a train of tone-pip probes in the range 0.125-8 kHz. For DPOAEs, the biasing-tone level for criterion suppression increased with probe-tone frequency by 8-9 dB/octave, consistent with an apex-to-base gradient of biasing-tone-induced basilar membrane displacement, as we verified by computational simulation. In contrast, the biasing-tone level for criterion loudness suppression increased with probe frequency by only 1-3 dB/octave, reminiscent of previously published data on low-side suppression of auditory nerve responses to characteristic frequency tones. These slopes were independent of biasing-tone frequency, but the biasing-tone sensation level required for criterion suppression was ~ 10 dB lower for the two infrasound biasing tones than for the 30-Hz biasing tone. On average, biasing-tone sensation levels as low as 5 dB were sufficient to modulate the perception of higher frequency sounds. Our results are relevant for recent debates on perceptual effects of environmental noise with very low-frequency content and might offer insight into the mechanism underlying low-side suppression.

Representation of interaural time delay in the human auditory midbrain.

Interaural time difference (ITD) is a critical cue to sound-source localization. Traditional models assume that sounds leading at one ear, and perceived on that side, are processed in the opposite midbrain. Using functional magnetic resonance imaging we demonstrate that as the ITDs of sounds increase, midbrain activity can switch sides, even though perceived location remains on the same side. The data require a new model for human ITD processing.

Ocular and cervical vestibular evoked myogenic potentials elicited by air-conducted, low-frequency sound.

BACKGROUND: Sound is not only detected by the cochlea, but also, at high intensities, by the vestibular system. Acoustic activation of the vestibular system can manifest itself in vestibular evoked myogenic potentials (VEMPs). In a clinical setting, VEMPs are usually evoked with rather high-frequency sound (500 Hz and higher), despite the fact that only a fraction of saccular and utricular hair cells in the striolar region is available for high-frequency stimulation. OBJECTIVE: As a growing proportion of the population complains about low-frequency environmental noise, including reports on vestibular symptoms, the activation of the vestibular system by low-frequency sound deserves better understanding. METHODS: We recorded growth functions of oVEMPs and cVEMPs evoked with air-conducted sound at 120 Hz and below. We estimated VEMP thresholds and tested whether phase changes of the stimulus carrier result in changes of VEMP amplitude and latency. RESULTS: The VEMP response of the otholith organs to low-frequency sound is uniform and not tuned when corrected for middle ear attenuation by A-weighting the stimulus level. Different stimulus carrier phases result in phase-correlated changes of cVEMP latencies and amplitudes. CONCLUSIONS: VEMPs can be evoked with rather low-frequency sound, but high thresholds suggest that they are unlikely to be triggered by environmental sounds.

Masking with interaurally "double-delayed" stimuli: the range of internal delays in the human brain.

Is binaural processing in humans different to that of other mammals? While psychophysical data suggest that the range of internal delays necessary for processing interaural time differences is at least +/-3 ms, physiological data from small mammals indicate a more limited range. This study demonstrates that binaural detection is impeded by reduced interaural coherence in auditory channels remote from the signal frequency, in accordance with the wider critical bandwidths reported for binaural processing. This explains previous psychophysical data without requiring long internal delays. The current psychophysical data support the view that human binaural processing is similar to that of other mammals.

Audio-tactile cues from an object's fall change estimates of one's body height.

When we drop an object from our hands, we use internal models of both our body height and object-motion to predict when it will hit the floor. What happens if the sensory feedback finally received from the impact conflicts with this prediction? The present study shows that such conflict results in changes in the internal estimates of our body height: When the object people dropped takes longer than expected to hit the floor, they report feeling taller and behave as if their legs were longer. This provides the first evidence of cross-modal recalibration of body-height representations as a function of changes in the distant environment. Crucially, the recalibration results from a mismatch between the predicted and actual outcome of an action, the ball's release and impact, which are causally-related but separated in space and time. These results suggest that implicit models of object-motion can interact with implicit and explicit models of one's body height.

A framework for testing and comparing binaural models.

Auditory research has a rich history of combining experimental evidence with computational simulations of auditory processing in order to deepen our theoretical understanding of how sound is processed in the ears and in the brain. Despite significant progress in the amount of detail and breadth covered by auditory models, for many components of the auditory pathway there are still different model approaches that are often not equivalent but rather in conflict with each other. Similarly, some experimental studies yield conflicting results which has led to controversies. This can be best resolved by a systematic comparison of multiple experimental data sets and model approaches. Binaural processing is a prominent example of how the development of quantitative theories can advance our understanding of the phenomena, but there remain several unresolved questions for which competing model approaches exist. This article discusses a number of current unresolved or disputed issues in binaural modelling, as well as some of the significant challenges in comparing binaural models with each other and with the experimental data. We introduce an auditory model framework, which we believe can become a useful infrastructure for resolving some of the current controversies. It operates models over the same paradigms that are used experimentally. The core of the proposed framework is an interface that connects three components irrespective of their underlying programming language: The experiment software, an auditory pathway model, and task-dependent decision stages called artificial observers that provide the same output format as the test subject.

Low-frequency characteristics of human and guinea pig cochleae.

Previous physiological studies investigating the transfer of low-frequency sound into the cochlea have been invasive. Predictions about the human cochlea are based on anatomical similarities with animal cochleae but no direct comparison has been possible. This paper presents a noninvasive method of observing low frequency cochlear vibration using distortion product otoacoustic emissions (DPOAE) modulated by low-frequency tones. For various frequencies (15-480 Hz), the level was adjusted to maintain an equal DPOAE-modulation depth, interpreted as a constant basilar membrane displacement amplitude. The resulting modulator level curves from four human ears match equal-loudness contours (ISO226:2003) except for an irregularity consisting of a notch and a peak at 45 Hz and 60 Hz, respectively, suggesting a cochlear resonance. This resonator interacts with the middle ear stiffness. The irregularity separates two regions of the middle ear transfer function in humans: A slope of 12 dB/octave below the irregularity suggests mass-controlled impedance resulting from perilymph movement through the helicotrema; a 6-dB/octave slope above the irregularity suggests resistive cochlear impedance and the existence of a traveling wave. The results from four guinea pig ears showed a 6-dB/octave slope on either side of an irregularity around 120 Hz, and agree with published data.

Low-frequency sound exposure causes reversible long-term changes of cochlear transfer characteristics.

Intense, low-frequency sound presented to the mammalian cochlea induces temporary changes of cochlear sensitivity, for which the term 'Bounce' phenomenon has been coined. Typical manifestations are slow oscillations of hearing thresholds or the level of otoacoustic emissions. It has been suggested that these alterations are caused by changes of the mechano-electrical transducer transfer function of outer hair cells (OHCs). Shape estimates of this transfer function can be derived from low-frequency-biased distortion product otoacoustic emissions (DPOAE). Here, we tracked the transfer function estimates before and after triggering a cochlear Bounce. Specifically, cubic DPOAEs, modulated by a low-frequency biasing tone, were followed over time before and after induction of the cochlear Bounce. Most subjects showed slow, biphasic changes of the transfer function estimates after low-frequency sound exposure relative to the preceding control period. Our data show that the operating point changes biphasically on the transfer function with an initial shift away from the inflection point followed by a shift towards the inflection point before returning to baseline values. Changes in transfer function and operating point lasted for about 180 s. Our results are consistent with the hypothesis that intense, low-frequency sound disturbs regulatory mechanisms in OHCs. The homeostatic readjustment of these mechanisms after low-frequency offset is reflected in slow oscillations of the estimated transfer functions.