Infrasonic hearing in birds: a review of audiometry and hypothesized structure-function relationships.

The perception of airborne infrasound (sounds below 20 Hz, inaudible to humans except at very high levels) has been documented in a handful of mammals and birds. While animals that produce vocalizations with infrasonic components (e.g. elephants) present conspicuous examples of potential use of infrasound in the context of communication, the extent to which airborne infrasound perception exists among terrestrial animals is unclear. Given that most infrasound in the environment arises from geophysical sources, many of which could be ecologically relevant, communication might not be the only use of infrasound by animals. Therefore, infrasound perception could be more common than currently realized. At least three bird species, each of which do not communicate using infrasound, are capable of detecting infrasound, but the associated auditory mechanisms are not well understood. Here we combine an evaluation of hearing measurements with anatomical observations to propose and evaluate hypotheses supporting avian infrasound detection. Environmental infrasound is mixed with non-acoustic pressure fluctuations that also occur at infrasonic frequencies. The ear can detect such non-acoustic pressure perturbations and therefore, distinguishing responses to infrasound from responses to non-acoustic perturbations presents a great challenge. Our review shows that infrasound could stimulate the ear through the middle ear (tympanic) route and by extratympanic routes bypassing the middle ear. While vibration velocities of the middle ear decline towards infrasonic frequencies, whole-body vibrations - which are normally much lower amplitude than that those of the middle ear in the 'audible' range (i.e. >20 Hz) - do not exhibit a similar decline and therefore may reach vibration magnitudes comparable to the middle ear at infrasonic frequencies. Low stiffness in the middle and inner ear is expected to aid infrasound transmission. In the middle ear, this could be achieved by large air cavities in the skull connected to the middle ear and low stiffness of middle ear structures; in the inner ear, the stiffness of round windows and cochlear partitions are key factors. Within the inner ear, the sizes of the helicotrema and cochlear aqueduct are expected to play important roles in shunting low-frequency vibrations away from low-frequency hair-cell sensors in the cochlea. The basilar papilla, the auditory organ in birds, responds to infrasound in some species, and in pigeons, infrasonic-sensitive neurons were traced back to the apical, abneural end of the basilar papilla. Vestibular organs and the paratympanic organ, a hair cell organ outside of the inner ear, are additional untested candidates for infrasound detection in birds. In summary, this review brings together evidence to create a hypothetical framework for infrasonic hearing mechanisms in birds and other animals.

Impacts of broadband sound on silver (Hypophthalmichthys molitrix) and bighead (H. nobilis) carp hearing thresholds determined using auditory evoked potential audiometry.

Invasive silver (Hypophthalmichthys molitrix) and bighead (H. nobilis) carp, collectively referred to as bigheaded carps, threaten aquatic ecosystems of the Upper Midwestern USA. Due to the extensive ecological impacts associated with these species, prevention of their further range expansion is the aim for fisheries management. Recent behavioral studies indicate bigheaded carps are deterred by acoustic barriers and exhibit negative phonotaxis in response to anthropogenic sound sources (≥ 150 dB re 1 µPa). However, the impact of long-term exposure to these sounds on the hearing capabilities of bigheaded carps has not been well documented. In this study, the auditory evoked potential (AEP) technique was used to determine auditory thresholds among bigheaded carps before and after exposure to high intensity (155.7 ± 4.7 dB re 1 µPa SPLrms; - 8.0 ± 4.7 dB re 1 ms-2 PALrms; mean ± SD) broadband sound. Fish were exposed to sound for 30 min or 24 h and AEP measurements were taken at three time points: immediately after exposure, 48 h, or 96 h later. Results indicate that silver and bighead carp experience temporary threshold shifts (TTSs) in frequency detection following sound exposure with the magnitude and length of TTS correlated with exposure duration. The findings from this study will be used to increase the long-term efficacy of acoustical deterrent measures aimed at preventing further range expansion of bigheaded carps.

Frequency-specific electric response audiometry (ERA) and its clinical application in the diagnosis of hearing defects in the dog.

Reference values were established for frequency-specific electric response audiometry (ERA) in dogs on the basis of the results of ERA examinations of 200 animals with normal hearing. Air-conducting acoustic tubes with foam stoppers were used in the determination of the following: the latencies of waves I, III and V; interpeak latencies (IPL) I-III, III-V and I-V; amplitudes I and V; and the amplitude difference I-V. A frequency-specific stimulus (tone pip) was used for frequency-specific examination (1 to 4 kHz) over the entire frequency range indicated. These reference values were then used for the clinical examination of 50 dogs with hearing defects. A frequency-specific ERA was conducted and the results evaluated. These findings made it possible to draw objective conclusions about the degree, type and site of the hearing defects. Frequency-specific electric response audiometry was shown to be an important diagnostic tool for the detection of partial high- and low-frequency hearing loss and for the characterisation of hearing defects of otological, otoneurological and neurological origin.

Brainstem auditory-evoked potential assessment of auditory function and congenital deafness in llamas (Lama glama) and alpacas (L. pacos).

Auditory function of llamas and alpacas was assessed objectively by means of brainstem auditory-evoked response audiometry (BAER) to establish the normal hearing range and to test the hypothesis of a correlation between blue eyes, white coat, and deafness. Sixty-three camelids were available for the study. Thirteen animals had blue irides; 1 animal had 1 blue and 1 pigmented iris. Wave latencies, amplitudes, and interpeak latencies were measured under general anesthetic. Click stimuli (dB [HL]) were delivered by an insert earphone. Four to five positive peaks could be detected; waves I, II, and V were reproducible; wave II appeared infrequently; and wave IV generally merged with wave V to form a complex. Peak latencies decreased and peak amplitudes increased as stimulus intensity increased. A hearing threshold level of 10-20 dB (HL) was proposed as the normal range in llamas and alpacas. None of the animals with pigmentation of coat and iris showed any degree of hearing impairment. Seven of the 10 blue-eyed, pure-white animals were bilaterally deaf and one of them was unilaterally deaf. However, 2 blue-eyed, white animals exhibited normal hearing ability. Three blue-eyed animals with pigmented coat did not show any hearing impairment. All white animals with normal iris pigmentation had normal auditory function; so did the 1 animal with 1 normal and 1 blue iris. The high frequency (78%) of bilaterally deaf animals with pure white coat and blue iris pigmentation supports the hypothesis of a correlation between pigmentation anomalies and congenital deafness in llamas and alpacas.

Audiograms estimated from brainstem tone-evoked potentials in dogs from 10 days to 1.5 months of age.

The objective of this study was to build audiograms from thresholds of brainstem tone-evoked potentials in dogs and to evaluate age-related change of the audiogram in puppies. Results were obtained from 9 Beagle puppies 10-47 days of age. Vertex to mastoid brainstem auditory-evoked potentials in response to 5.1-millisecond Hanning-gated sine waves with frequencies octave-spaced from 0.5 to 32 kHz were recorded. Three dogs were examined at 10, 13, 19, 25, and 45 days. Four other dogs were examined at 16 days. Data from 7 dogs between 42 and 47 days of age were pooled to obtain audiogram reference values in 1.5-month-old puppies. The best auditory threshold lowered from above 60 dB sound pressure level (SPL) to values close to 0 dB SPL between 13 and 25 days of age and then stabilized. The audible frequency range widened, including 32 kHz in all tested dogs from the 19th day. In the 7 1.5-month-old puppies, the mean auditory threshold decreased by 11 dB per octave from 0.5 to 2 kHz. The auditory threshold was lowest and held the same value from 2 to 8 kHz. The mean auditory threshold increased by 20 dB per octave from 8 to 32 kHz. Near threshold, click-evoked potentials test only a small part of the audible frequency range in dogs. Use of tone-evoked potentials may become a powerful tool in investigating dogs with possible partial hearing loss, including during the auditory system maturation period.

The auditory neurobiology of marsupials: a review.

The marsupials, the large group of mammals which develop during fetal life in an externalized pouch, have been given little attention by auditory neurobiologists. In this review the structure of the auditory systems of the handful of marsupials which have been studied is described, the course of auditory development mapped, and the behavioral and electrophysiological manifestations of hearing examined. It is argued that research on the highly accessible developing marsupial will provide information about the development of hearing difficult to obtain from, but applicable to all, mammalian species.

The postnatal development of frequency-place code and tuning characteristics in the auditory midbrain of the phyllostomid bat, Carollia perspicillata.

This report describes the postnatal development of hearing range, auditory sensitivity and tonotopy within the inferior colliculus (IC) of a mammal specialized for ultrasonic hearing. The experimental animal, Carollia perspicillata, has an adult hearing range of 7-110 kHz (characteristic frequencies) but lack any significant overrepresentation of a limited frequency band as known for rhinolophoid bats and Pteronotus. The audiogram of the newborn Carollia includes characteristic frequencies from 8 to 76 kHz, which is about 65% of the adult hearing range. As in adults, low frequencies are represented in the dorsolateral portion of the IC. However, at birth the ventromedial IC is non-responsive to acoustic stimulation up to intensities of 90 dB SPL. During development there is a progressive conversion of non-responsive IC areas into acoustically responsive slabs with characteristic frequencies above 76 kHz along the dorsolateral to ventromedial (low-to-high frequency) IC axis. This development is superimposed by a non-uniform shift of characteristic frequency: a decrease of CFs in dorsolateral regions, and an increase of CFs in ventromedial areas. The results suggest a bidirectional shift of frequency representation along the cochlear tonotopic axis.

Hearing in primitive mammals: Monodelphis domestica and Marmosa elegans.

Although opossums of the Family Didelphidae usually serve as a parsimonious starting point for tracing the otological and neurological evolution of modern mammals, audiological data for Didelphid opossums is available only for the North American opossum (Didelphis virginiana) which because of its large size, may be one of the least representative genera of the family. The present report extends the audiological data to two other species of Didelphid opossums, Monodelphis domestica, and Marmosa elegans. At 60 dB SPL, the hearing of Monodelphis extends from 3.6 kHz to 77 kHz, with a range of best sensitivity from 8 to 64 kHz while the hearing of Marmosa extends from 3.8 kHz to 80 kHz, with a range of best sensitivity from 8 to 64 kHz. Neither species was found to be particularly sensitive to tones, with the average lowest threshold near 20 dB SPL for Monodelphis and 33 dB SPL for Marmosa. These results indicate that like the North American opossum both genera are sensitive to high frequencies yet relatively insensitive to sound. Because the hearing of the three genera of Didelphids agree in several respects, it can be concluded that sensitivity to high frequencies almost certainly was present in ancient mammals, probably following quickly after the acquisition of a 3 ossicle middle ear linkage. It is not unlikely that the utility value of high frequency hearing, rather than highly sensitive hearing, may have been a primary source of selective pressure for this morphological transformation.

Ototoxicity in dogs and cats.

Ears are special sense organs whose principal functions are hearing and maintaining equilibrium. Aminoglycoside antibiotics, erythromycin, polymyxin B, and cisplatin can affect either or both of these functions by binding with, injuring, and/or destroying special receptor cells associated with these functions. Severe hearing loss manifests itself as deafness, whereas loss of equilibrium will present as abnormal righting reflexes, nausea, and vomiting. Damage is proportional to levels of these ototoxins in the endolymphatic fluids. Evidence suggests that toxicity may be influenced by endolymphatic calcium concentrations, and levels of cAMP and cGMP are altered in specialized cochlear cells during ototoxicity, suggesting an additional mechanism for ototoxicity. The administration of salicylates and loop diuretics may potentiate the action of ototoxins, especially aminoglycoside antibiotics, probably by increasing the levels of these toxins in the endolymphatic fluid. Although many of these assessments have been made in laboratory animals, applicability may also be expected in small domestic animals, and extreme care should be taken in prescribing potentially ototoxic drugs to small animals. Cochlear damage from ototoxic compounds occurs initially in the cells detecting high-frequency sounds located at the lower basal region. In aging dogs and humans, this sensitivity of receptors in the lower basal region is enhanced. Early auditory damage is detectable by BAER and cochlear microphonic potentials. Vestibular responses can also be detected early as vestibular ocular reflexes and visual-vestibulo-ocular reflexes. Early detection is especially important because early changes can sometimes be reversible. Cavinton (apovincaminic acid) and fosfomycin represent examples of experimental agents being evaluated in laboratory animals for application as potential treatments to limit the ototoxicity associated with various drugs.

Auditory evoked potentials in the Japanese monkey.

Auditory sensitivity based on auditory brain stem response (ABR), whole nerve action potential (AP), and cochlear microphonics (CM) to tone bursts of 0.5-8 kHz were compared with behavioral audiometry in the Japanese monkeys. Although sensitivity loss at 4-6 kHz was observed in these potentials, an increase in sensitivity at 8 kHz was obtained only in the ABR. Thus the sensitivity loss at 4-6 kHz originates at the peripheral system and the increased sensitivity at 8 kHz originates at the central.

Brainstem auditory evoked response in the diagnosis of inner ear injury in the horse.

Brainstem auditory evoked response (BAER) testing was done to evaluate inner ear/VIIIth cranial nerve (CN8) function in the horse. The BAER test consisted of stimulating the auditory system with clicks and recording far-field responses of the brainstem auditory components via cutaneous electrodes and a signal averaging system. The normal response was shown to be a series of waves occurring within the first 10 msec after the stimulus click. Functional loss of the auditory receptor organ (cochlea) or CN8 results in loss of the entire response on the side of the injury. Because of the anatomic relationships of the peripheral auditory and vestibular systems, trauma to one will injure the other. Therefore, auditory testing (BAER tests) may be used to advantage in the diagnosis of peripheral vestibular disease. The BAER test was used in a horse that had signs suggestive of vestibular dysfunction or a brain lesion. The test helped to demonstrate a unilateral inner ear/CN8 lesion and to discount the probability of a more central lesion.