Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution.

The skull of 'Ligulalepis' from the Early Devonian of Australia (AM-F101607) has significantly expanded our knowledge of early osteichthyan anatomy, but its phylogenetic position has remained uncertain. We herein describe a second skull of 'Ligulalepis' and present micro-CT data on both specimens to reveal novel anatomical features, including cranial endocasts. Several features previously considered to link 'Ligulalepis' with actinopterygians are now considered generalized osteichthyan characters or of uncertain polarity. The presence of a lateral cranial canal is shown to be variable in its development between specimens. Other notable new features include the presence of a pineal foramen, the some detail of skull roof sutures, the shape of the nasal capsules, a placoderm-like hypophysial vein, and a chondrichthyan-like labyrinth system. New phylogenetic analyses place 'Ligulalepis' as a stem osteichthyan, specifically as the sister taxon to 'psarolepids' plus crown osteichthyans. The precise position of 'psarolepids' differs between parsimony and Bayesian analyses.

Fast Click Rate Electrocochleography and Auditory Brainstem Response in Normal-Hearing Adults Using Continuous Loop Averaging Deconvolution.

OBJECTIVES: Using the continuous loop averaging deconvolution (CLAD) technique for conventional electrocochleography (ECochG) and auditory brainstem response (ABR) recordings, the effects of testing at high stimulus rates may have the potential to diagnose disorders of the inner ear and auditory nerve. First, a body of normative data using the CLAD technique must be established. DESIGN: Extratympanic click ECochG and ABR to seven stimulus rates using CLAD were measured simultaneously from a tympanic membrane electrode and surface electrodes on the forehead and mastoid of 42 healthy individuals. RESULTS: Results showed that the compound action potential (AP) of the ECochG and waves I, III, and V of the ABR decreased in amplitude and increased in latency as stimulus rate was increased from standard 7.1 clicks/s up to 507.81 clicks/s, with sharp reduction in AP amplitude at 97.66 clicks/s and reaching asymptote at 292.97 clicks/s. The summating potential (SP) of the ECochG, however, stayed relatively stable, resulting in increased SP/AP ratios with increasing rate. The SP/AP amplitude ratio showed more stability than AP amplitude findings, thus it is recommended for use in evaluation of cochlear and neural response. CONCLUSIONS: Results of both amplitude and latency data from this normative neural adaptation function of the auditory pathway serves as guide for improving diagnostic utility of both ECochG and ABR using CLAD as a reliable technique in distinguishing inner ear and auditory nerve disorders.

Acoustic input impedance of the avian inner ear measured in ostrich (Struthio camelus).

In both mammals and birds, the mechanical behavior of the middle ear structures is affected by the mechanical impedance of the inner ear. In this study, the aim was to quantify the acoustic impedance of the avian inner ear in the ostrich, which allows us to determine the effect on columellar vibrations and middle ear power flow in future studies. To determine the inner ear impedance, vibrations of the columella were measured for both the quasi-static and acoustic stimulus frequencies. In the frequency range of 0.3-4 kHz, we used electromagnetic stimulation of the ossicle and a laser Doppler vibrometer to measure the vibration response. At low frequencies, harmonic displacements were imposed on the columella using piezo stimulation and the resulting force response was measured with a force sensor. From these measurement data, the acoustic impedance of the inner ear could be determined. A simple RLC model in series of the impedance measurements resulted in a stiffness reactance of KIE = 0.20·10(12) Pa/m³, an inertial impedance of MIE = 0.652·10(6) Pa s(2)/m³, and a resistance of RIE = 1.57·10(9) Pa s/m. We found that values of the inner ear impedance in the ostrich are one to two orders in magnitude smaller than what is found in mammal ears.

Changes in cochlear function related to acoustic stimulation of cervical vestibular evoked myogenic potential stimulation.

Evaluation of cervical evoked myogenic potentials (c-VEMP) is commonly applied in clinical investigations of patients with suspected neurotological symptoms. Short intense acoustic stimulation of peak levels close to 130 dB SPL is required to elicit the responses. A recent publication on bilateral significant sensorineural hearing loss related to extensive VEMP stimulation motivates evaluations of immediate effects on hearing acuity related to the intense acoustic stimulation required to elicit c-VEMP responses. The aim of the current study was to investigate changes in DPOAE-levels and hearing thresholds in relation to c-VEMP testing in humans. More specifically, the current focus is on immediate changes in hearing thresholds and changes in DPOAE-levels at frequencies 0.5 octaves above the acoustic stimulation when applying shorter tone bursts than previously used. Hearing acuity before and immediately after exposure to c-VEMP stimulation was examined in 24 patients with normal hearing referred for neurotologic testing. The stimulation consisted of 192 tonebursts of 6 ms and was presented at 500 Hz and 130 dB peSPL. Békésy thresholds at 0.125-8 kHz and DPOAE I/O growth functions with stimulation at 0.75 and 3 kHz were used to assess c-VEMP related changes in hearing status. No significant deterioration in Békésy thresholds was detected. Significant reduction in DPOAE levels at 0.75 (0.5-1.35 dB) and 3 kHz (1.6-2.1 dB) was observed after c-VEMP stimulation without concomitant changes in cochlear compression. The results indicated that there was no immediate audiometric loss related to c-VEMP stimulation in the current group of patients. The significant reduction of DPOAE levels at a wider frequency range than previously described after the c-VEMP test could be related to the stimulation with shorter tone bursts. The results show that c-VEMP stimulation causes reduction in DPOAE-levels at several frequencies that corresponds to half the reductions in DPOAE levels reported after exposure to the maximally allowed occupational noise for an 8 h working day. Consequently, extended stimuli intensity or stimulation repetition with c-VEMP testing should be avoided to reduce the risk for noise-induced cochlear injury.

Attentional modulation of the inner ear: a combined otoacoustic emission and EEG study.

Attending to a single stimulus in a complex multisensory environment requires the ability to select relevant information while ignoring distracting input. The underlying mechanism and involved neuronal levels of this attentional gain control are still a matter of debate. Here, we investigated the influence of intermodal attention on different levels of auditory processing in humans. It is known that the activity of the cochlear amplifier can be modulated by efferent neurons of the medial olivocochlear complex. We used distortion product otoacoustic emission (DPOAE) measurements to monitor cochlear activity during an intermodal cueing paradigm. Simultaneously, central auditory processing was assessed by electroencephalography (EEG) with a steady-state paradigm targeting early cortical responses and analysis of alpha oscillations reflecting higher cognitive control of attentional modulation. We found effects of selective attention at all measured levels of the auditory processing: DPOAE levels differed significantly between periods of visual and auditory attention, showing a reduction during visual attention, but no change during auditory attention. Primary auditory cortex activity, as measured by the auditory steady-state response (ASSR), differed between conditions, with higher ASSRs during auditory than visual attention. Furthermore, the analysis of cortical oscillatory activity revealed increased alpha power over occipitoparietal and frontal regions during auditory compared with visual attention, putatively reflecting suppression of visual processing. In conclusion, this study showed both enhanced processing of attended acoustic stimuli in early sensory cortex and reduced processing of distracting input, both at higher cortical levels and at the most peripheral level of the hearing system, the cochlea.

Evolution of vertebrate mechanosensory hair cells and inner ears: toward identifying stimuli that select mutation driven altered morphologies.

Among the major distance senses of vertebrates, the ear is unique in its complex morphological changes during evolution. Conceivably, these changes enable the ear to adapt toward sensing various physically well-characterized stimuli. This review develops a scenario that integrates sensory cell with organ evolution. We propose that molecular and cellular evolution of the vertebrate hair cells occurred prior to the formation of the vertebrate ear. We previously proposed that the genes driving hair cell differentiation were aggregated in the otic region through developmental re-patterning that generated a unique vertebrate embryonic structure, the otic placode. In agreement with the presence of graviceptive receptors in many vertebrate outgroups, it is likely that the vertebrate ear originally functioned as a simple gravity-sensing organ. Based on the rare occurrence of angular acceleration receptors in vertebrate outgroups, we further propose that the canal system evolved with a more sophisticated ear morphogenesis. This evolving morphogenesis obviously turned the initial otocyst into a complex set of canals and recesses, harboring multiple sensory epithelia each adapted to the acquisition of a specific aspect of a given physical stimulus. As support for this evolutionary progression, we provide several details of the molecular basis of ear development.

An analysis of the acoustic input impedance of the ear.

Ear canal acoustics was examined using a one-dimensional lossy transmission line with a distributed load impedance to model the ear. The acoustic input impedance of the ear was derived from sound pressure measurements in the ear canal of healthy human ears. A nonlinear least squares fit of the model to data generated estimates for ear canal radius, ear canal length, and quantified the resistance that would produce transmission losses. Derivation of ear canal radius has application to quantifying the impedance mismatch at the eardrum between the ear canal and the middle ear. The length of the ear canal was found, in general, to be longer than the length derived from the one-quarter wavelength standing wave frequency, consistent with the middle ear being mass-controlled at the standing wave frequency. Viscothermal losses in the ear canal, in some cases, may exceed that attributable to a smooth rigid wall. Resistance in the middle ear was found to contribute significantly to the total resistance. In effect, this analysis "reverse engineers" physical parameters of the ear from sound pressure measurements in the ear canal.

Vibratory sources as compound stimuli for the octavolateralis systems: dissection of specific stimulation channels using multiple behavioral approaches.

Underwater vibratory sources simultaneously present acoustic and hydrodynamic disturbances. Because vibratory dipole sources are poor sonic projectors, most researchers have assumed that such sources are of greatest relevance to the lateral line system. Both hydroacoustic principles and empirical studies have shown that dipole sources are also a potent stimulus to the inner ear. Responses to vibratory sources in mottled sculpin (Cottus bairdi) were assessed using unconditioned orienting, differential and nondifferential conditioning. Orienting responses are dominated by lateral line inputs and eliminated by lateral line inactivation. Simple conditioning depends on inputs from other systems and was not affected by lateral line inactivation. Differential conditioning alters behavioral control, and sculpin could be conditioned to ignore substrate-borne vibrations and respond only to hydroacoustic stimulation of the ear. The lateral line and inner ear of mottled sculpin do not necessarily exhibit range fractionation, as both systems operate over a similar distance (within 1.5 body lengths) and respond to many of the same sources. Vibratory dipole sources generate compound stimuli that simultaneously activate multiple octavolateralis systems, and sculpin make use of the channels differentially under different behavioral tasks.

Frequency matching of vocalizations to inner-ear sensitivity along an altitudinal gradient in the coqui frog.

Acoustic communication involves both the generation and the detection of a signal. In the coqui frog (Eleutherodactylus coqui), it is known that the spectral contents of its calls systematically change with altitude above sea level. Here, distortion product otoacoustic emissions are used to assess the frequency range over which the inner ear is sensitive. It is found that both the spectral contents of the calls and the inner-ear sensitivity change in a similar fashion along an altitudinal gradient. As a result, the call frequencies and the auditory tuning are closely matched at all altitudes. We suggest that the animal's body size determines the frequency particulars of the call apparatus and the inner ear.

Modeling the vestibular evoked myogenic potential.

Measuring the vestibular evoked myogenic potential (VEMP) promises to become a routine method for assessing vestibular function, although the technique is not yet standardized. To overcome the problem that the VEMP amplitude depends not only on the inhibition triggered by the acoustic stimulation of the vestibular end organs in the inner ear, but also on the tone of the muscle from which the potential is recorded, the VEMP is often normalized by dividing through a measure of the electromyogram (EMG) activity. The underlying idea is that VEMP amplitude and EMG activity are proportional. But this would imply that the muscle tone is irrelevant for a successful VEMP recording, contradicting experimental evidence. Here, an analytical model is presented that allows to resolve the contradiction. The EMG is modeled as the sum of motor unit action potentials (MUAPs). A brief inhibition can be characterized by its equivalent rectangular duration (ERD), irrespective of the actual time course of the inhibition. The VEMP resembles a polarity-inverted MUAP under such circumstances. Its amplitude is proportional to both the ERD and the MUAP rate. The EMG activity, by contrast, is proportional to the square root of the MUAP rate. Thus, the normalized VEMP still depends on the muscle tone. To avoid confounding effects of the muscle tone, the standard deviation of the EMG could be considered. But the inhibition effect on the standard deviation is small so that the measuring time would have to be much longer than usual today.

Ten-year longitudinal study of the effect of impulse noise exposure from gunshot on inner ear function.

This longitudinal study investigated how chronic gunshot noise exposure affects cochlear and saccular function in police officers who engaged in regular target shooting practice using dual protection (ear plugs plus earmuffs) for >10 years. In 1997, 20 male police officers underwent audiometry before and two weeks after shooting. Twelve of the original subjects were re-examined by audiometry coupled with vestibular evoked myogenic potential (VEMP) test in 2007. Significant deterioration of mean hearing thresholds at frequencies of 500 Hz through 4000 Hz was noted ten years later, affecting both ears. However, only the frequencies of 4000 and 6000 Hz on the left ear revealed significant difference in mean hearing thresholds compared with healthy controls. Abnormal VEMP responses were evident in nine police officers (75%), including absent VEMPs 7 and delayed VEMPs 2. In conclusion, deterioration to hearing may occur after long term exposure to gunshots, even when double hearing protection is used. Further study is in progress regarding how to preserve both cochlear and saccular function during long term gunshot exposure.

Sound pressure and particle acceleration audiograms in three marine fish species from the Adriatic Sea.

Fishes show great variability in hearing sensitivity, bandwidth, and the appropriate stimulus component for the inner ear (particle motion or pressure). Here, hearing sensitivities in three vocal marine species belonging to different families were described in terms of sound pressure and particle acceleration. In particular, hearing sensitivity to tone bursts of varying frequencies were measured in the red-mouthed goby Gobius cruentatus, the Mediterranean damselfish Chromis chromis, and the brown meagre Sciaena umbra using the non-invasive auditory evoked potential-recording technique. Hearing thresholds were measured in terms of sound pressure level and particle acceleration level in the three Cartesian directions using a newly developed miniature pressure-acceleration sensor. The brown meagre showed the broadest hearing range (up to 3000 Hz) and the best hearing sensitivity, both in terms of sound pressure and particle acceleration. The red-mouthed goby and the damselfish were less sensitive, with upper frequency limits of 700 and 600 Hz, respectively. The low auditory thresholds and the large hearing bandwidth of S. umbra indicate that sound pressure may play a role in S. umbra's hearing, even though pronounced connections between the swim bladder and the inner ears are lacking.

Prospective electrophysiologic findings of round window stimulation in a model of experimentally induced stapes fixation.

HYPOTHESIS: Mechanical stimulation of the round window (RW) with an active middle ear implant (AMEI) with and without experimentally induced stapes fixation (SF) results in equivalent electrophysiologic measures of cochlear microphonic (CM), compound action potential (CAP), and auditory brainstem response (ABR). BACKGROUND: Where normal oval window functionality is mitigated, the RW provides a pathway to mechanically stimulate the inner ear. METHODS: Measurements of the CM, CAP, and ABR were made in 5 ears of 4 chinchillas with acoustic stimulation and with application of the AMEI to the RW with and without experimentally induced SF using pure-tone stimuli (0.25-20 kHz) presented at differing intensities (-20 to 80 dB SPL vs. 0.01 mV to 3.16 V). RESULTS: Morphologies of the CM, CAP, and ABR were similar between acoustic and RW stimulation with and without SF. Stapes fixation increased CM thresholds relative to RW stimulation without fixation by a frequency-dependent 4- to 13-dB mV (mean, 7.9 +/- 3.2 dB mV). Although the thresholds changed with SF, CM sensitivities and amplitude dynamic range were identical to normal. The CAP in all conditions demonstrated equivalent decreasing amplitudes and increasing latency with decreasing intensity (decibel sound pressure level versus decibel millivolt). Stapes fixation increased the CAP thresholds at all frequencies, ranging from 9 to 24 dB mV (mean, 17.7 +/- 4.9 dB mV). Auditory brainstem response waveforms were preserved across experimental conditions. CONCLUSION: Mechanical stimulation of the RW in an animal model of SF generates functionally similar inputs to the cochlea as normal acoustic and RW mechanical inputs but with increased thresholds. With further study, AMEIs may provide a surgical option for correction of otosclerosis and ossicular chain disruption.

The detection of pressure fluctuations, sonic audition, is the dominant mode of dipole-source detection in goldfish (Carassius auratus).

Behavioral detection of a low-frequency (40 Hz) vibratory dipole at source distances of 1.5-24 cm was measured by classically conditioned respiratory suppression in goldfish (Carassius auratus). Detection thresholds were compared across distances and before and after ablation of individual octavolateralis sensory channels. Detection thresholds, expressed in units of pressure (SPL), remained roughly constant as distance between the stimulus source and animal increased. Lateral line inactivation, using CoCl2, had no measurable effect on sensitivity, although some other results can be construed as weak evidence for a small contribution of the lateral line to dipole detection when source distances are

Reflex control of the human inner ear: a half-octave offset in medial efferent feedback that is consistent with an efferent role in the control of masking.

The high sensitivity and frequency selectivity of the mammalian cochlea is due to amplification produced by outer hair cells (OHCs) and controlled by medial olivocochlear (MOC) efferents. Data from animals led to the view that MOC fibers provide frequency-specific inhibitory feedback; however, these studies did not measure intact MOC reflexes. To test whether MOC inhibition is primarily at the frequency that elicits the MOC activity, acoustically elicited MOC effects were quantified in humans by the change in otoacoustic emissions produced by 60-dB SPL tone and half-octave-band noise elicitors at different frequencies relative to a 40-dB SPL, 1-kHz probe tone. On average, all elicitors produced MOC effects that were skewed (elicitor frequencies -1 octave below the probe produced larger effects than those -1 octave above). The largest MOC effects were from elicitors below the probe frequency for contra- and bilateral elicitors but were from elicitors centered at the probe frequency for ipsilateral elicitors. Typically, ipsilateral elicitors produced larger effects than contralateral elicitors and bilateral elicitors produced effects near the ipsi+contra sum. Elicitors at levels down to 30-dB SPL produced similar patterns. Tuning curves (TCs) interpolated from these data were V-shaped with Q10s approximately 2. These are sharper than MOC-fiber TCs found near 1 kHz in cats and guinea pigs. Because cochlear amplification is skewed (more below the best frequency of a cochlear region), these data are consistent with an anti-masking role of MOC efferents that reduces masking by reducing the cochlear amplification seen at 1 kHz.

Molecular identity and functional properties of a novel T-type Ca2+ channel cloned from the sensory epithelia of the mouse inner ear.

The molecular identity of non-Cav1.3 channels in auditory and vestibular hair cells has remained obscure, yet the evidence in support of their roles to promote diverse Ca2+-dependent functions is indisputable. Recently, a transient Cav3.1 current that serves as a functional signature for the development and regeneration of hair cells has been identified in the chicken basilar papilla. The Cav3.1 current promotes spontaneous activity of the developing hair cell, which may be essential for synapse formation. Here, we have isolated and sequenced the full-length complementary DNA of a distinct isoform of Cav3.1 in the mouse inner ear. The channel is derived from alternative splicing of exon14, exon25A, exon34, and exon35. Functional expression of the channel in Xenopus oocytes yielded Ca2+ currents, which have a permeation phenotype consistent with T-type channels. However, unlike most multiion channels, the T-type channel does not exhibit the anomalous mole fraction effect, possibly reflecting comparable permeation properties of divalent cations. The Cav3.1 channel was expressed in sensory and nonsensory epithelia of the inner ear. Moreover, there are profound changes in the expression levels during development. The differential expression of the channel during development and the pharmacology of the inner ear Cav3.1 channel may have contributed to the difficulties associated with identification of the non-Cav1.3 currents.

Imaging the living inner ear using intravital confocal microscopy.

Confocal laser scanning microscopy permits detailed visualization of structures deep within thick fluorescently labeled specimen. This makes it possible to investigate living cells inside intact tissue without prior chemical sample fixation and sectioning. Isolated guinea pig temporal bones have previously been used for confocal experiments in vitro, but tissue deterioration limits their use to a few hours after the death of the animal. In order to preserve the cochlea in an optimal functional and physiological condition, we have developed an in vivo model based on a confocal microscopy approach. Using a ventral surgical approach, the inner ear is exposed in deeply anaesthetized, tracheotomized, living guinea pigs. To label the inner ear structures, scala tympani is perfused via an opening in the basal turn, delivering tissue culture medium with fluorescent vital dyes (RH 795 and calcein AM). An apical opening is made in the bony shell of cochlea to enable visualization using a custom-built objective lens. Intravital confocal microscopy, with preserved blood and nerve supply, may offer an important tool for studying auditory physiology and the pathology of hearing loss. After acoustic overstimulation, shortening and swelling of the sensory hair cells were observed.

Distortion product otoacoustic emission of symphony orchestra musicians before and after rehearsal.

The 2f1-f2 distortion product otoacoustic emission (DPOAE) and hearing levels are obtained for 12 normal-hearing symphony orchestra musicians both before and after their rehearsal. The DPOAE fine structures are determined and analyzed according to the character and prevalence of ripples. Hearing levels, DPOAE levels, and DPOAE fine structures before and after rehearsal are similar, indicating that no or marginal temporary change of the state of hearing were caused by the exposure. The data were further compared to similar data for occupationally nonexposed subjects, one group which was age and gender matched, and other two groups of younger individuals (one group with better hearing levels than the other). The data for the age and gender matched group compared well with the musicians data (and the data for the group of better-hearing younger individuals). In general, the analyses of hearing thresholds and DPOAE data thus lead to the same conclusions concerning the state of hearing.