Reversible induction of phantom auditory sensations through simulated unilateral hearing loss

Tinnitus, a phantom auditory sensation, is associated with hearing loss in most cases, but it is unclear if hearing loss causes tinnitus. Phantom auditory sensations can be induced in normal hearing listeners when they experience severe auditory deprivation such as confinement in an anechoic chamber, which can be regarded as somewhat analogous to a profound bilateral hearing loss. As this condition is relatively uncommon among tinnitus patients, induction of phantom sounds by a lesser degree of auditory deprivation could advance our understanding of the mechanisms of tinnitus. In this study, we therefore investigated the reporting of phantom sounds after continuous use of an earplug. 18 healthy volunteers with normal hearing wore a silicone earplug continuously in one ear for 7 days. The attenuation provided by the earplugs simulated a mild high-frequency hearing loss, mean attenuation increased from <10 dB at 0.25 kHz to >30 dB at 3 and 4 kHz. 14 out of 18 participants reported phantom sounds during earplug use. 11 participants presented with stable phantom sounds on day 7 and underwent tinnitus spectrum characterization with the earplug still in place. The spectra showed that the phantom sounds were perceived predominantly as high-pitched, corresponding to the frequency range most affected by the earplug. In all cases, the auditory phantom disappeared when the earplug was removed, indicating a causal relation between auditory deprivation and phantom sounds. This relation matches the predictions of our computational model of tinnitus development, which proposes a possible mechanism by which a stabilization of neuronal activity through homeostatic plasticity in the central auditory system could lead to the development of a neuronal correlate of tinnitus when auditory nerve activity is reduced due to the earplug.     

Hearing sensitivity and amplitude coding in bats are differentially shaped by echolocation calls and social calls

Differences in auditory perception between species are influenced by phylogenetic origin and the perceptual challenges imposed by the natural environment, such as detecting prey- or predator-generated sounds and communication signals. Bats are well suited for comparative studies on auditory perception since they predominantly rely on echolocation to perceive the world, while their social calls and most environmental sounds have low frequencies. We tested if hearing sensitivity and stimulus level coding in bats differ between high and low-frequency ranges by measuring auditory brainstem responses (ABRs) of 86 bats belonging to 11 species. In most species, auditory sensitivity was equally good at both high- and low-frequency ranges, while amplitude was more finely coded for higher frequency ranges. Additionally, we conducted a phylogenetic comparative analysis by combining our ABR data with published data on 27 species. Species-specific peaks in hearing sensitivity correlated with peak frequencies of echolocation calls and pup isolation calls, suggesting that changes in hearing sensitivity evolved in response to frequency changes of echolocation and social calls. Overall, our study provides the most comprehensive comparative assessment of bat hearing capacities to date and highlights the evolutionary pressures acting on their sensory perception.     

Postnatal development of auditory central evoked responses and thalamic cellular properties

During development, the sense of hearing changes rapidly with age, especially around hearing onset. During this period, auditory structures are highly sensitive to alterations of the acoustic environment, such as hearing loss or background noise. This sensitivity includes auditory temporal processing, which is important for processing complex sounds, and for acquiring reading and language skills. Developmental changes can be observed at multiple levels of brain organization—from behavioral responses to cellular responses, and at every auditory nucleus. Neuronal properties and sound processing change dramatically in auditory cortex neurons after hearing onset. However, development of its primary source, the auditory thalamus, or medial geniculate body (MGB), has not been well studied over this critical time window. Furthermore, to understand how temporal processing develops, it is important to determine the relative maturation of temporal processing not only in the MGB, but also in its inputs. Cellular properties of rat MGB neurons were studied using in vitro whole‐cell patch‐clamp recordings, at ages postnatal day (P) 7–9; P15–17, and P22–32. Auditory evoked potentials were measured in P14–17 and P22–32 rats. MGB action potentials became about five times faster, and the ability to generate spike trains increased with age, particularly at frequencies of 50 Hz and higher. Evoked potential responses, including auditory brainstem responses (ABR), middle latency responses (MLR), and amplitude modulation following responses, showed increased amplitudes with age, and ABRs and MLRs additionally showed decreased latencies with age. Overall, temporal processing at subthalamic nuclei is concurrently maturing with MGB cellular properties. © 2013 Wiley Periodicals, Inc. Develop Neurobiol 74: 541–555, 2014     

Basic auditory processes involved in the analysis of speech sounds

This paper reviews the basic aspects of auditory processing that play a role in the perception of speech. The frequency selectivity of the auditory system, as measured using masking experiments, is described and used to derive the internal representation of the spectrum (the excitation pattern) of speech sounds. The perception of timbre and distinctions in quality between vowels are related to both static and dynamic aspects of the spectra of sounds. The perception of pitch and its role in speech perception are described. Measures of the temporal resolution of the auditory system are described and a model of temporal resolution based on a sliding temporal integrator is outlined. The combined effects of frequency and temporal resolution can be modelled by calculation of the spectro-temporal excitation pattern, which gives good insight into the internal representation of speech sounds. For speech presented in quiet, the resolution of the auditory system in frequency and time usually markedly exceeds the resolution necessary for the identification or discrimination of speech sounds, which partly accounts for the robust nature of speech perception. However, for people with impaired hearing, speech perception is often much less robust.     

Sound production, hearing and possible interception under ambient noise conditions in the topmouth minnow Pseudorasbora parva

Sounds were produced by the topmouth minnow Pseudorasbora parva , a common Eurasian cyprinid, during feeding but not during intraspecific interactions. Feeding sounds were short broadband pulses with main energies between 100 and 800 Hz. They varied in their characteristics (number of single sounds per feeding sequence, sound duration and period, and sound pressure level) depending on the food type (chironomid larvae, Tubifex worms and flake food). The loudest sounds were emitted when food was taken up at the water surface, most probably reflecting 'suctorial' feeding. Auditory sensitivities were determined between 100 and 4000 Hz utilizing the auditory evoked potentials recording technique. Under laboratory conditions and in the presence of natural ambient noise recorded in Lake Neusiedl in eastern Austria, best hearing sensitivities were between 300 and 800 Hz (57 dB re 1 μPa v . 72 dB in the presence of ambient noise). Threshold-to-noise ratios were positively correlated to the sound frequency. The correlation between sound spectra and auditory thresholds revealed that P. parva can detect conspecific sounds up to 40 cm distance under ambient noise conditions. Thus, feeding sounds could serve as an auditory cue for the presence of food during foraging.     

Object localization in cluttered acoustical environments

In nature, sounds from objects of interest arrive at the ears accompanied by sound waves from other actively emitting objects and by reflections off of nearby surfaces. Despite the fact that all of these waveforms sum at the eardrums, humans with normal hearing effortlessly segregate one sound source from another. Our laboratory is investigating the neural basis of this perceptual feat, often called the "cocktail party effect", using the barn owl as an animal model. The barn owl, renowned for its ability to localize sounds and its spatiotopic representation of auditory space, is an established model for spatial hearing. Here, we briefly review the neural basis of sound-localization of a single sound source in an anechoic environment and then generalize the ideas developed therein to cases in which there are multiple, concomitant sound sources and acoustical reflection.     

Ontogenetic development of weberian ossicles and hearing abilities in the African bullhead catfish

Background

The Weberian apparatus of otophysine fishes facilitates sound transmission from the swimbladder to the inner ear to increase hearing sensitivity. It has been of great interest to biologists since the 19^th century. No studies, however, are available on the development of the Weberian ossicles and its effect on the development of hearing in catfishes.

Methodology/Principal Findings

We investigated the development of the Weberian apparatus and auditory sensitivity in the catfish Lophiobagrus cyclurus. Specimens from 11.3 mm to 85.5 mm in standard length were studied. Morphology was assessed using sectioning, histology, and X-ray computed tomography, along with 3D reconstruction. Hearing thresholds were measured utilizing the auditory evoked potentials recording technique. Weberian ossicles and interossicular ligaments were fully developed in all stages investigated except in the smallest size group. In the smallest catfish, the intercalarium and the interossicular ligaments were still missing and the tripus was not yet fully developed. Smallest juveniles revealed lowest auditory sensitivity and were unable to detect frequencies higher than 2 or 3 kHz; sensitivity increased in larger specimens by up to 40 dB, and frequency detection up to 6 kHz. In the size groups capable of perceiving frequencies up to 6 kHz, larger individuals had better hearing abilities at low frequencies (0.05–2 kHz), whereas smaller individuals showed better hearing at the highest frequencies (4–6 kHz).

Conclusions/Significance

Our data indicate that the ability of otophysine fish to detect sounds at low levels and high frequencies largely depends on the development of the Weberian apparatus. A significant increase in auditory sensitivity was observed as soon as all Weberian ossicles and interossicular ligaments are present and the chain for transmitting sounds from the swimbladder to the inner ear is complete. This contrasts with findings in another otophysine, the zebrafish, where no threshold changes have been observed.     

Dynamics of Vocalization-Induced Modulation of Auditory Cortical Activity at Mid-utterance

Background

Recent research has addressed the suppression of cortical sensory responses to altered auditory feedback that occurs at utterance onset regarding speech. However, there is reason to assume that the mechanisms underlying sensorimotor processing at mid-utterance are different than those involved in sensorimotor control at utterance onset. The present study attempted to examine the dynamics of event-related potentials (ERPs) to different acoustic versions of auditory feedback at mid-utterance.

Methodology/Principal findings

Subjects produced a vowel sound while hearing their pitch-shifted voice (100 cents), a sum of their vocalization and pure tones, or a sum of their vocalization and white noise at mid-utterance via headphones. Subjects also passively listened to playback of what they heard during active vocalization. Cortical ERPs were recorded in response to different acoustic versions of feedback changes during both active vocalization and passive listening. The results showed that, relative to passive listening, active vocalization yielded enhanced P2 responses to the 100 cents pitch shifts, whereas suppression effects of P2 responses were observed when voice auditory feedback was distorted by pure tones or white noise.

Conclusion/Significance

The present findings, for the first time, demonstrate a dynamic modulation of cortical activity as a function of the quality of acoustic feedback at mid-utterance, suggesting that auditory cortical responses can be enhanced or suppressed to distinguish self-produced speech from externally-produced sounds.     

Event-related potentials in an auditory oddball situation in the rat

Evoked potentials were recorded from the auditory cortex of both freely moving and anesthetized rats when deviant sounds were presented in a homogenous series of standard sounds (oddball condition). A component of the evoked response to deviant sounds, the mismatch negativity (MMN), may underlie the ability to discriminate acoustic differences, a fundamental aspect of auditory perception. Whereas most MMN studies in animals have been done using simple sounds, this study involved a more complex set of sounds (synthesized vowels). The freely moving rats had previously undergone behavioral training in which they learned to respond differentially to these sounds. Although we found little evidence in this preparation for the typical, epidurally recorded, MMN response, a significant difference between deviant and standard evoked potentials was noted for the freely moving animals in the 100-200 ms range following stimulus onset. No such difference was found in the anesthetized animals.     

Auditory brain-stem responses evoked by electrical stimulation of the cochlear nucleus in human subjects

When auditory nerve function is lost due to surgical removal of bilateral acoustic tumors, a sense of hearing may be restored by means of an auditory brain-stem implant (ABI), which electrically stimulates the auditory pathway at the level of the cochlear nucleus. Placement of the stimulating electrodes during surgical implantation may be aided by electrically evoked auditory brain-stem responses (EABRs) recorded intra-operatively. To establish preliminary standards for human EABRs evoked by electrical stimulation of the cochlear nucleus, short-latency evoked potentials were recorded from 6 ABI patients who were either already implanted or undergoing implantation surgery. Neural responses were distinguished from stimulus artifact and equipment artifact by their properties during stimulus polarity reversal and amplitude variation. Other properties contributed to further identification of the evoked potentials as auditory responses (EABRs). The response waveforms generally had 2 or 3 waves. The peak latencies of these waves (approximately 0.3, 1.3, and 2.2 msec) and the brain-stem localization of the region from which they could be elicited are consistent with auditory brain-stem origin.     

Auditory processing of complex sounds: an overview.

The past 30 years has seen a remarkable development in our understanding of how the auditory system--particularly the peripheral system--processes complex sounds. Perhaps the most significant has been our understanding of the mechanisms underlying auditory frequency selectivity and their importance for normal and impaired auditory processing. Physiologically vulnerable cochlear filtering can account for many aspects of our normal and impaired psychophysical frequency selectivity with important consequences for the perception of complex sounds. For normal hearing, remarkable mechanisms in the organ of Corti, involving enhancement of mechanical tuning (in mammals probably by feedback of electro-mechanically generated energy from the hair cells), produce exquisite tuning, reflected in the tuning properties of cochlear nerve fibres. Recent comparisons of physiological (cochlear nerve) and psychophysical frequency selectivity in the same species indicate that the ear's overall frequency selectivity can be accounted for by this cochlear filtering, at least in bandwidth terms. Because this cochlear filtering is physiologically vulnerable, it deteriorates in deleterious conditions of the cochlea--hypoxia, disease, drugs, noise overexposure, mechanical disturbance--and is reflected in impaired psychophysical frequency selectivity. This is a fundamental feature of sensorineural hearing loss of cochlear origin, and is of diagnostic value. This cochlear filtering, particularly as reflected in the temporal patterns of cochlear fibres to complex sounds, is remarkably robust over a wide range of stimulus levels. Furthermore, cochlear filtering properties are a prime determinant of the 'place' and 'time' coding of frequency at the cochlear nerve level, both of which appear to be involved in pitch perception. The problem of how the place and time coding of complex sounds is effected over the ear's remarkably wide dynamic range is briefly addressed. In the auditory brainstem, particularly the dorsal cochlear nucleus, are inhibitory mechanisms responsible for enhancing the spectral and temporal contrasts in complex sounds. These mechanisms are now being dissected neuropharmacologically. At the cortical level, mechanisms are evident that are capable of abstracting biologically relevant features of complex sounds. Fundamental studies of how the auditory system encodes and processes complex sounds are vital to promising recent applications in the diagnosis and rehabilitation of the hearing impaired.     

Cortical motion deafness

The extent to which the auditory system, like the visual system, processes spatial stimulus characteristics such as location and motion in separate specialized neuronal modules or in one homogeneously distributed network is unresolved. Here we present a patient with a selective deficit for the perception and discrimination of auditory motion following resection of the right anterior temporal lobe and the right posterior superior temporal gyrus (STG). Analysis of stimulus identity and location within the auditory scene remained intact. In addition, intracranial auditory evoked potentials, recorded preoperatively, revealed motion-specific responses selectively over the resected right posterior STG, and electrical cortical stimulation of this region was experienced by the patient as incoming moving sounds. Collectively, these data present a patient with cortical motion deafness, providing evidence that cortical processing of auditory motion is performed in a specialized module within the posterior STG.     

Ontogenetic development of auditory sensitivity and sound production in the squeaker catfish <Emphasis Type="Italic">Synodontis schoutedeni</Emphasis>

Background  

Surveys of ontogenetic development of hearing and sound production in fish are scarce, and the ontogenetic development of acoustic communication has been investigated in only two fish species so far. Studies on the labyrinth fish Trichopsis vittata and the toadfish Halobatrachus didactylus show that the ability to detect conspecific sounds develops during growth. In otophysine fish, which are characterized by Weberian ossicles and improved hearing sensitivities, the ontogenetic development of sound communication has never been investigated. We analysed the ontogeny of the auditory sensitivity and vocalizations in the mochokid catfish Synodontis schoutedeni. Mochokid catfishes of the genus Synodontis are commonly called squeakers because they produce broadband stridulation sounds during abduction and adduction of pectoral fin spines. Fish from six different size groups - from 22 mm standard length to 126 mm - were studied. Hearing thresholds were measured between 50 Hz and 6 kHz using the auditory evoked potentials recording technique; stridulation sounds were recorded and their sound pressure levels determined. Finally, absolute sound power spectra were compared to auditory sensitivity curves within each size group.     

Influence of the Background Noise on Recognition of Signals with a Complex Spectrum Structure in the Beluga Whale (<Emphasis Type="Italic">Delphinapterus leucas</Emphasis>)

The frequency resolving power of hearing (FRP) of the beluga whale Delphinapterus leucas was studied as dependent on influence of lasting low-intensity sounds (of the ultrasonic range from –20 to +10 dB). Testing of the spectrum ripple-phase reversal was used in conjunction with a noninvasive recording of auditory evoked potentials. FRP parameters were found to depend nonmonotonically on the intensity of the background noise. The resultant adaptation effects can be explained by the fact that, in response to the high-intensity signals, the auditory system sensitivity is reduced to the level optimal for analysis of these signals.     

Brain activity in an awake chimpanzee in response to the sound of her own name

The brain activity of a fully awake chimpanzee being presented with her name was investigated. Event-related potentials (ERPs) were measured for each of the following auditory stimuli: the vocal sound of the subject''s own name (SON), the vocal sound of a familiar name of another group member, the vocal sound of an unfamiliar name and a non-vocal sound. Some differences in ERP waveforms were detected between kinds of stimuli at latencies at which P3 and Nc components are typically observed in humans. Following stimulus onset, an Nc-like negative shift at approximately 500 ms latency was observed, particularly in response to SON. Such specific ERP patterns suggest that the chimpanzee processes her name differently from other sounds.     

Seeing invisible motion: a human FMRI study

A common view about visual consciousness is that it could arise when and where activity reaches some higher level of processing along the cortical hierarchy. Reports showing that activity in striate cortex can be dissociated from awareness , whereas the latter modulates activity in higher areas , point in this direction. In the specific case of visual motion, a central, "perceptual" role has been assigned to area V5: several human and monkey studies have shown V5 activity to correlate with the motion percept. Here we show that activity in this and other higher cortical areas can be also dissociated from perception and follow the physical stimulus instead. The motion information in a peripheral grating modulated fMRI responses, despite being invisible to human volunteers: under crowding conditions , areas V3A, V5, and parietal cortex still showed increased activity when the grating was moving compared to when it was flickering. We conclude that stimulus-specific activation of higher cortical areas does not necessarily result in awareness of the underlying stimulus.     

Auditory temporal resolution and evoked responses to pulsed sounds for the Yangtze finless porpoises (<Emphasis Type="Italic">Neophocaena phocaenoides asiaeorientalis</Emphasis>)

Temporal cues are important for some forms of auditory processing, such as echolocation. Among odontocetes (toothed whales, dolphins, and porpoises), it has been suggested that porpoises may have temporal processing abilities which differ from other odontocetes because of their relatively narrow auditory filters and longer duration echolocation signals. This study examined auditory temporal resolution in two Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis) using auditory evoked potentials (AEPs) to measure: (a) rate following responses and modulation rate transfer function for 100 kHz centered pulse sounds and (b) hearing thresholds and response amplitudes generated by individual pulses of different durations. The animals followed pulses well at modulation rates up to 1,250 Hz, after which response amplitudes declined until extinguished beyond 2,500 Hz. The subjects had significantly better hearing thresholds for longer, narrower-band pulses similar to porpoise echolocation signals compared to brief, broadband sounds resembling dolphin clicks. Results indicate that the Yangtze finless porpoise follows individual acoustic signals at rates similar to other odontocetes tested. Relatively good sensitivity for longer duration, narrow-band signals suggests that finless porpoise hearing is well suited to detect their unique echolocation signals.     

Contralateral Noise Stimulation Delays P300 Latency in School-Aged Children

Background and Objective

The auditory cortex modulates auditory afferents through the olivocochlear system, which innervates the outer hair cells and the afferent neurons under the inner hair cells in the cochlea. Most of the studies that investigated the efferent activity in humans focused on evaluating the suppression of the otoacoustic emissions by stimulating the contralateral ear with noise, which assesses the activation of the medial olivocochlear bundle. The neurophysiology and the mechanisms involving efferent activity on higher regions of the auditory pathway, however, are still unknown. Also, the lack of studies investigating the effects of noise on human auditory cortex, especially in peadiatric population, points to the need for recording the late auditory potentials in noise conditions. Assessing the auditory efferents in schoolaged children is highly important due to some of its attributed functions such as selective attention and signal detection in noise, which are important abilities related to the development of language and academic skills. For this reason, the aim of the present study was to evaluate the effects of noise on P300 responses of children with normal hearing.

Methods

P300 was recorded in 27 children aged from 8 to 14 years with normal hearing in two conditions: with and whitout contralateral white noise stimulation.

Results

P300 latencies were significantly longer at the presence of contralateral noise. No significant changes were observed for the amplitude values.

Conclusion

Contralateral white noise stimulation delayed P300 latency in a group of school-aged children with normal hearing. These results suggest a possible influence of the medial olivocochlear activation on P300 responses under noise condition.     

The hearing abilities of the prawn Palaemon serratus

The mechanism of sound reception and the hearing abilities of the prawn (Palaemon serratus) have been studied using a combination of anatomical, electron microscopic and electrophysiological approaches, revealing that P. serratus is responsive to sounds ranging in frequency from 100 to 3000 Hz. It is the first time that the Auditory Brainstem Response (ABR) recording technique has been used on invertebrates, and the acquisition of hearing ability data from the present study adds valuable information to the inclusion of an entire sub-phylum of animals when assessing the potential impact of anthropogenic underwater sounds on marine organisms. Auditory evoked potentials were acquired from P. serratus, using two subcutaneous electrodes positioned in the carapace close to the supraesophageal ganglion and the statocyst (a small gravistatic organ located below the eyestalk on the peduncle of the bilateral antennules). The morphology of the statocyst receptors and the otic nerve pathways to the brain have also been studied, and reveal that P. serratus possesses an array of sensory hairs projecting from the floor of the statocyst into a mass of sand granules embedded in a gelatinous substance. It is the purpose of this work to show that the statocyst is responsive to sounds propagated through water from an air mounted transducer. The fundamental measure of the hearing ability of any organism possessing the appropriate receptor mechanism is its audiogram, which presents the lowest level of sound that the species can hear as a function of frequency. The statocyst of P. serratus is shown here to be sensitive to the motion of water particles displaced by low-frequency sounds ranging from 100 Hz up to 3000 Hz, with a hearing acuity similar to that of a generalist fish. Also, recorded neural waveforms were found to be similar in both amplitude and shape to those acquired from fish and higher vertebrates, when stimulated with low-frequency sound, and complete ablation of the electrophysiological response was achieved by removal of the statocyst.     

Evoked Potential Audiograms of the Nurse Shark (Ginglymostoma cirratum) and the Yellow Stingray (Urobatis jamaicensis)

The hearing thresholds of the nurse shark, Ginglymostoma cirratum, and the yellow stingray, Urobatis jamaicensis, were measured using auditory evoked potentials (AEP). Stimuli were calibrated using a pressure-velocity probe so that the acoustic field could be completely characterized. The results show similar hearing thresholds for both species and similar hearing thresholds to previously measured audiograms for the lemon shark, Negaprion brevirostris, and the horn shark, Heterodontis francisi. All of these audiograms suggest poor hearing abilities, raising questions about field studies showing attraction of sharks to acoustic signals. By extrapolating the particle acceleration thresholds into estimates of their equivalent far-field sound pressure levels, it appears that these sharks cannot likely detect most of the sounds that have attracted sharks in the field.