ABSTRACT
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.
Lizards have highly directional ears, owing to strong acoustical coupling of the eardrums and almost perfect sound transmission from the contralateral ear. To investigate the neural processing of ...this remarkable tympanic directionality, we combined biophysical measurements of eardrum motion in the Tokay gecko with neurophysiological recordings from the auditory nerve. Laser vibrometry shows that their ear is a two-input system with approximately unity interaural transmission gain at the peak frequency (∼ 1.6 kHz). Median interaural delays are 260 μs, almost three times larger than predicted from gecko head size, suggesting interaural transmission may be boosted by resonances in the large, open mouth cavity (Vossen et al. 2010). Auditory nerve recordings are sensitive to both interaural time differences (ITD) and interaural level differences (ILD), reflecting the acoustical interactions of direct and indirect sound components at the eardrum. Best ITD and click delays match interaural transmission delays, with a range of 200-500 μs. Inserting a mold in the mouth cavity blocks ITD and ILD sensitivity. Thus the neural response accurately reflects tympanic directionality, and most neurons in the auditory pathway should be directional.
Turtles, like other amphibious animals, face a trade-off between terrestrial and aquatic hearing. We used laser vibrometry and auditory brainstem responses to measure their sensitivity to vibration ...stimuli and to airborne versus underwater sound. Turtles are most sensitive to sound underwater, and their sensitivity depends on the large middle ear, which has a compliant tympanic disc attached to the columella. Behind the disc, the middle ear is a large air-filled cavity with a volume of approximately 0.5 ml and a resonance frequency of approximately 500 Hz underwater. Laser vibrometry measurements underwater showed peak vibrations at 500–600 Hz with a maximum of 300 µm s−1 Pa−1, approximately 100 times more than the surrounding water. In air, the auditory brainstem response audiogram showed a best sensitivity to sound of 300–500 Hz. Audiograms before and after removing the skin covering reveal that the cartilaginous tympanic disc shows unchanged sensitivity, indicating that the tympanic disc, and not the overlying skin, is the key sound receiver. If air and water thresholds are compared in terms of sound intensity, thresholds in water are approximately 20–30 dB lower than in air. Therefore, this tympanic ear is specialized for underwater hearing, most probably because sound-induced pulsations of the air in the middle ear cavity drive the tympanic disc.
In most anuran amphibians, acoustic communication is of prime importance for mate localization and selection. The tympanic middle ear increases auditory sensitivity and directionality and is ...therefore expected to be favoured by natural selection. However, especially within the family of true toads (Bufonidae) there is a tendency for species to lose parts of the middle ear apparatus and consequently have a reduced sensitivity to high-frequency sounds (above 1 kHz). Part of the explanation for this may be that development of the middle ear is especially slow in bufonids, and thus the middle ear would be more likely to be lost or non-functional in paedomorphic species. However, a timeline of development of the middle ear has not been established previously. The goal of the present study was to investigate middle ear development in a toad species that has a well-known natural history and acoustic communication behaviour. We made a detailed study of anatomy and biophysics of the middle ear with measurements of auditory sensitivity across age in post-metamorphic natterjack toads (Epidalea calamita). The tadpoles and toadlets were raised in the laboratory, so their exact age was known, their auditory sensitivity was measured using auditory brainstem responses, and middle ear development and function were assessed by anatomical studies and laser vibrometry. We found that the developmental stage of the middle ear depends on the size of the toad rather than its age. The middle ear was functional at the earliest at a snout-vent length of 40 mm, which for these toads was around 500 days post-metamorphosis, close to the time of first reproduction. The functional, adult-like middle ear was shown to have 30 dB increased sensitivity to the dominant frequency of the mating call compared with sensitivities measured in newly metamorphosed individuals.
The eardrums of all terrestrial vertebrates (tetrapods) are connected through Eustachian tubes or interaural canals. In some of the animals, these connections create pressure-gradient directionality, ...an enhanced directionality by interaction of sound arriving at both sides of the eardrum and strongly dependent on interaural transmission attenuation.
Even though the tympanic middle ear has originated independently in the major tetrapod groups, in each group the ancestral condition probably was that the two middle ears were exposed in the mouth cavity with relatively high interaural transmission. Recent vertebrates form a continuum from perfect interaural transmission (0 dB in a certain frequency band) and pronounced eardrum directionality (30–40 dB) in the lizards, over somewhat attenuated transmission and limited directionality in birds and frogs, to the strongly attenuated interaural transmission and functionally isolated pressure receiver ears in the mammals.
Since some of the binaural interaction already takes place at the eardrum in animals with strongly coupled ears, producing enhanced interaural time and level differences, the subsequent neural processing may be simpler. In robotic simulations of lizards, simple binaural subtraction (EI cells, found in brainstem nuclei of both frogs and lizards) produces strongly lateralized responses that are sufficient for steering the animal robustly to sound sources.
► Acoustical coupling between the ears in frogs, lizards and birds produce strong directional cues (pressure-gradient receivers). ► The directionality depends on the interaural transmission gain that can approach unity (lizards). ► The coupled middle ear likely is the
ancestral condition of all tetrapod ears. ► Neural directional processing may be simpler in animals with strong acoustical coupling.
The position of testudines in vertebrate phylogeny is being re-evaluated. At present, testudine morphological and molecular data conflict when reconstructing phylogenetic relationships. Complicating ...matters, the ecological niche of stem testudines is ambiguous. To understand how turtles have evolved to hear in different environments, we examined middle ear morphology and scaling in most extant families, as well as some extinct species, using 3-dimensional reconstructions from micro magnetic resonance (MR) and submillimeter computed tomography (CT) scans. All families of testudines exhibited a similar shape of the bony structure of the middle ear cavity, with the tympanic disk located on the rostrolateral edge of the cavity. Sea Turtles have additional soft tissue that fills the middle ear cavity to varying degrees. When the middle ear cavity is modeled as an air-filled sphere of the same volume resonating in an underwater sound field, the calculated resonances for the volumes of the middle ear cavities largely fell within testudine hearing ranges. Although there were some differences in morphology, there were no statistically significant differences in the scaling of the volume of the bony middle ear cavity with head size among groups when categorized by phylogeny and ecology. Because the cavity is predicted to resonate underwater within the testudine hearing range, the data support the hypothesis of an aquatic origin for testudines, and function of the middle ear cavity in underwater sound detection.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Sound is vital for communication and navigation across the animal kingdom and sound communication is unrivaled in accuracy and information richness over long distances both in air and water. The ...source level (SL) of the sound is a key factor in determining the range at which animals can communicate and the range at which echolocators can operate their biosonar. Here we compile, standardize and compare measurements of the loudest animals both in air and water. In air we find a remarkable similarity in the highest SLs produced across the different taxa. Within all taxa we find species that produce sound above 100 dB
peak
re 20 μPa at 1 m, and a few bird and mammal species have SLs as high as 125 dB
peak
re 20 μPa at 1 m. We next used pulsating sphere and piston models to estimate the maximum sound pressures generated in the radiated sound field. These data suggest that the loudest species within all taxa converge upon maximum pressures of 140–150 dB
peak
re 20 μPa in air. In water, the toothed whales produce by far the loudest SLs up to 240 dB
peak
re 1 μPa at 1 m. We discuss possible physical limitations to the production, radiation and propagation of high sound pressures. Furthermore, we discuss physiological limitations to the wide variety of sound generating mechanisms that have evolved in air and water of which many are still not well-understood or even unknown. We propose that in air, non-linear sound propagation forms a limit to producing louder sounds. While non-linear sound propagation may play a role in water as well, both sperm whale and pistol shrimp reach another physical limit of sound production, the cavitation limit in water. Taken together, our data suggests that both in air and water, animals evolved that produce sound so loud that they are pushing against physical rather than physiological limits of sound production, radiation and propagation.
The auditory brainstem response in two lizard species Brittan-Powell, Elizabeth F; Christensen-Dalsgaard, Jakob; Tang, Yezhong ...
The Journal of the Acoustical Society of America,
08/2010, Letnik:
128, Številka:
2
Journal Article
Recenzirano
Odprti dostop
Although lizards have highly sensitive ears, it is difficult to condition them to sound, making standard psychophysical assays of hearing sensitivity impractical. This paper describes non-invasive ...measurements of the auditory brainstem response (ABR) in both Tokay geckos (Gekko gecko; nocturnal animals, known for their loud vocalizations) and the green anole (Anolis carolinensis, diurnal, non-vocal animals). Hearing sensitivity was measured in 5 geckos and 7 anoles. The lizards were sedated with isoflurane, and ABRs were measured at levels of 1 and 3% isoflurane. The typical ABR waveform in response to click stimulation showed one prominent and several smaller peaks occurring within 10 ms of the stimulus onset. ABRs to brief tone bursts revealed that geckos and anoles were most sensitive between 1.6-2 kHz and had similar hearing sensitivity up to about 5 kHz (thresholds typically 20-50 dB SPL). Above 5 kHz, however, anoles were more than 20 dB more sensitive than geckos and showed a wider range of sensitivity (1-7 kHz). Generally, thresholds from ABR audiograms were comparable to those of small birds. Best hearing sensitivity, however, extended over a larger frequency range in lizards than in most bird species.
OBJECTIVES:The objectives of this study were to(1) estimate the hearing status of classical symphony orchestra musicians and (2) investigate the hypothesis that occupational sound exposure of ...symphony orchestra musicians leads to elevated hearing thresholds.
DESIGN:The study population comprised all the musicians from five symphony orchestras. Questionnaires were filled in by 337 subjects, and 212 subjects performed an audiometric test. For a group of 182 musicians (363 ears) the results of the audiometry was analyzed in relation to the individual exposure, which was estimated on the basis of sound measurements and questionnaire data regarding the exposure time. The mean hearing threshold at the frequencies 3, 4, and 6 kHz, corrected for age and sex, was used as outcome.
RESULTS:The musician ears with the highest exposure (29 of 363) had an additional threshold shift of 6.3 dB compared with the 238 ears with lowest exposure. The observed hearing loss of musicians was smaller compared with the noise-induced permanent threshold shift (NIPTS) predicted from ISO1999. A remaining confounding effect of age after ISO7029 age corrections could be observed to explain the difference in observed and predicted NIPTS. However, the observed hearing loss difference between the left and the right ear of musicians was 2.5 dB (95% confidence interval 1.5–3.6), which was similar to the NIPTS predicted from ISO1999. Most of the musicians had better hearing at 3, 4, and 6 kHz for age than expected, however, 29 ears with the highest exposure above 90.4 dBA with a mean exposure time of 41.7 years had significantly elevated hearing thresholds. Trumpet players and the left ear of first violinists had significantly elevated hearing thresholds compared with other musicians.
CONCLUSION:Most of the symphony orchestra musicians had better hearing than expected but they had a work-related risk of developing additional noise-induced hearing loss. The additional NITPS of the left ear compared with the right ear was at the expected level based on the cumulated sound exposure and ISO1999, indicating that performing music may induce hearing loss to the same extent as industrial noise.
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