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Although many crustaceans produce sounds, their hearing abilities and mechanisms are poorly understood, leaving uncertainties regarding whether or how these animals use sound for acoustic communication. Marine invertebrates lack gas-filled organs required for sound pressure detection, but some of them are known to be sensitive to particle motion. Here, we examined whether the American lobster ( Homarus americanus ) could detect sound and subsequently sought to discern the auditory mechanisms. Acoustic stimuli responses were measured using auditory evoked potential (AEP) methods. Neurophysiological responses were obtained from the brain using tone pips between 80 and 250 Hz, with best sensitivity at 80-120 Hz. There were no significant differences between the auditory thresholds of males and females. Repeated controls (recordings from deceased lobsters, moving electrodes away from the brain and reducing seawater temperature) indicated the evoked potentials' neuronal origin. In addition, AEP responses were similar before and after antennules (including statocysts) were ablated, demonstrating that the statocysts, a long-proposed auditory structure in crustaceans, are not the sensory organs responsible for lobster sound detection. However, AEPs could be eliminated (or highly reduced) after immobilizing hairfans, which cover much of lobster bodies. These results suggest that these external cuticular hairs are likely to be responsible for sound detection, and imply that hearing is mechanistically possible in a wider array of invertebrates than previously considered. Because the lobsters' hearing range encompasses the fundamental frequency of their buzzing sounds, it is likely that they use sound for intraspecific communication, broadening our understanding of the sensory ecology of this commercially vital species. The lobsters' low-frequency acoustic sensitivity also underscores clear concerns about the potential impacts of anthropogenic noise., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Hearing is vital for birds as they rely on acoustic communication with parents, mates, chicks and conspecifics. Amphibious seabirds face many ecological pressures, having to sense cues in air and underwater. Natural noise conditions have helped shape this sensory modality but anthropogenic noise is increasingly impacting seabirds. Surprisingly little is known about their hearing, despite their imperiled status. Understanding sound sensitivity is vital when we seek to manage the impacts of man-made noise. We measured the auditory sensitivity of nine wild Atlantic puffins, Fratercula arctica , in a capture-and-release setting in an effort to define their audiogram and compare these data with the hearing of other birds and natural rookery noise. Auditory sensitivity was tested using auditory evoked potential (AEP) methods. Responses were detected from 0.5 to 6 kHz. Mean thresholds were below 40 dB re. 20 µPa from 0.75 to 3 kHz, indicating that these were the most sensitive auditory frequencies, similar to other seabirds. Thresholds in the 'middle' frequency range 1-2.5 kHz were often down to 10-20 dB re. 20 µPa. The lowest thresholds were typically at 2.5 kHz. These are the first in-air auditory sensitivity data from multiple wild-caught individuals of a deep-diving alcid seabird. The audiogram was comparable to that of other birds of similar size, thereby indicating that puffins have fully functioning aerial hearing despite the constraints of their deep-diving, amphibious lifestyles. There was some variation in thresholds, yet animals generally had sensitive ears, suggesting aerial hearing is an important sensory modality for this taxon., Competing Interests: Competing interestsThe authors declare no competing financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Hearing is a primary sensory modality for birds. For seabirds, auditory data is challenging to obtain and hearing data are limited. Here, we present methods to measure seabird hearing in the field, using two Alcid species: the common murre Uria aalge and the Atlantic puffin Fratercula arctica Tests were conducted in a portable semi-anechoic crate using physiological auditory evoked potential (AEP) methods. The crate and AEP system were easily transportable to northern Iceland field sites, where wild birds were caught, sedated, studied and released. The resulting data demonstrate the feasibility of a field-based application of an established neurophysiology method, acquiring high quality avian hearing data in a relatively quiet setting. Similar field methods could be applied to other seabirds, and other bird species, resulting in reliable hearing data from a large number of individuals with a modest field effort. The results will provide insights into the sound sensitivity of species facing acoustic habitat degradation., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Sound transmission and reception are both vital components to odontocete echolocation and daily life. Here, we combine computed tomography (CT) scanning and finite element modeling to investigate the acoustic propagation of finless porpoise (Neophocaena asiaorientalis sunameri) echolocation pulses. The CT scanning and finite element method wave propagation model results support the well-accepted jaw-hearing pathway hypothesis and suggest an additional alternative auditory pathway composed of structures, mandible (lower jaw) and internal mandibular fat, with different acoustic impedances, which may also conduct sounds to the ear complexes. The internal mandibular fat is attached to the ear complex and encased by the mandibles laterally and anteriorly. The simulations show signals in this pathway initially propagate along the solid mandibles and are transmitted to the acoustically coupled soft tissue of the internal mandibular fat which conducts the stimuli posteriorly as it eventually arrives at ear complexes. While supporting traditional theories, this new bone-tissue conduction pathway might be meaningful to understand the hearing and sound reception processes in a wide variety of odontocetes species.

Documenting hearing abilities is vital to understanding a species' acoustic ecology and for predicting the impacts of increasing anthropogenic noise. Cetaceans use sound for essential biological functions such as foraging, navigation and communication; hearing is considered to be their primary sensory modality. Yet, we know little regarding the hearing of most, if not all, cetacean populations, which limits our understanding of their sensory ecology, population level variability and the potential impacts of increasing anthropogenic noise. We obtained audiograms (5.6-150 kHz) of 26 wild beluga whales to measure hearing thresholds during capture-release events in Bristol Bay, AK, USA, using auditory evoked potential methods. The goal was to establish the baseline population audiogram, incidences of hearing loss and general variability in wild beluga whales. In general, belugas showed sensitive hearing with low thresholds (<80 dB) from 16 to 100 kHz, and most individuals (76%) responded to at least 120 kHz. Despite belugas often showing sensitive hearing, thresholds were usually above or approached the low ambient noise levels measured in the area, suggesting that a quiet environment may be associated with hearing sensitivity and that hearing thresholds in the most sensitive animals may have been masked. Although this is just one wild population, the success of the method suggests that it should be applied to other populations and species to better assess potential differences. Bristol Bay beluga audiograms showed substantial (30-70 dB) variation among individuals; this variation increased at higher frequencies. Differences among individual belugas reflect that testing multiple individuals of a population is necessary to best describe maximum sensitivity and population variance. The results of this study quadruple the number of individual beluga whales for which audiograms have been conducted and provide the first auditory data for a population of healthy wild odontocetes., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

A critical concern with respect to marine animal acoustics is the issue of hearing "sensitivity," as it is widely used as a criterion for the onset of noise-induced effects. Important aspects of research on sensitivity to sound by marine animals include: uncertainties regarding how well these species detect and respond to different sounds; the masking effects of man-made sounds on the detection of biologically important sounds; the question how internal state, motivation, context, and previous experience affect their behavioral responses; and the long-term and cumulative effects of sound exposure. If we are to better understand the sensitivity of marine animals to sound we must concentrate research on these questions. In order to assess population level and ecological community impacts new approaches can possibly be adopted from other disciplines and applied to marine fauna.

We measured the hearing abilities of seven wild beluga whales (Delphinapterus leucas) during a collection-and-release experiment in Bristol Bay, AK. Here we summarize the methods and initial data from one animal and discuss the implications of this experiment. Audiograms were collected from 4 to 150 kHz. The animal with the lowest threshold heard best at 80 kHz and demonstrated overall good hearing from 22 to 110 kHz. The robustness of the methodology and data suggest that the auditory evoked potential audiograms can be incorporated into future collection-and-release health assessments. Such methods may provide high-quality results for multiple animals, facilitating population-level audiograms and hearing measures in new species.

While odontocetes do not have an external pinna that guides sound to the middle ear, they are considered to receive sound through specialized regions of the head and lower jaw. Yet odontocetes differ in the shape of the lower jaw suggesting that hearing pathways may vary between species, potentially influencing hearing directionality and noise impacts. This work measured the audiogram and received sensitivity of a Risso's dolphin (Grampus griseus) in an effort to comparatively examine how this species receives sound. Jaw hearing thresholds were lowest (most sensitive) at two locations along the anterior, midline region of the lower jaw (the lower jaw tip and anterior part of the throat). Responses were similarly low along a more posterior region of the lower mandible, considered the area of best hearing in bottlenose dolphins. Left- and right-side differences were also noted suggesting possible left-right asymmetries in sound reception or differences in ear sensitivities. The results indicate best hearing pathways may vary between the Risso's dolphin and other odontocetes measured. This animal received sound well, supporting a proposed throat pathway. For Risso's dolphins in particular, good ventral hearing would support their acoustic ecology by facilitating echo-detection from their proposed downward oriented echolocation beam.

Sound is a primary sensory cue for most marine mammals, and this is especially true for cetaceans. To passively and actively acquire information about their environment, cetaceans have some of the most derived ears of all mammals, capable of sophisticated, sensitive hearing and auditory processing. These capabilities have developed for survival in an underwater world where sound travels five times faster than in air, and where light is quickly attenuated and often limited at depth, at night, and in murky waters. Cetacean auditory evolution has capitalized on the ubiquity of sound cues and the efficiency of underwater acoustic communication. The sense of hearing is central to cetacean sensory ecology, enabling vital behaviours such as locating prey, detecting predators, identifying conspecifics, and navigating. Increasing levels of anthropogenic ocean noise appears to influence many of these activities. Here, we describe the historical progress of investigations on cetacean hearing, with a particular focus on odontocetes and recent advancements. While this broad topic has been studied for several centuries, new technologies in the past two decades have been leveraged to improve our understanding of a wide range of taxa, including some of the most elusive species. This chapter addresses topics including how sounds are received, what sounds are detected, hearing mechanisms for complex acoustic scenes, recent anatomical and physiological studies, the potential impacts of noise, and mysticete hearing. We conclude by identifying emerging research topics and areas which require greater focus., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

There is increasing concern that human-produced ocean noise is adversely affecting marine mammals, as several recent cetacean mass strandings may have been caused by animals' interactions with naval 'mid-frequency' sonar. However, it has yet to be empirically demonstrated how sonar could induce these strandings or cause physiological effects. In controlled experimental studies, we show that mid-frequency sonar can induce temporary hearing loss in a bottlenose dolphin (Tursiops truncatus). Mild-behavioural alterations were also associated with the exposures. The auditory effects were induced only by repeated exposures to intense sonar pings with total sound exposure levels of 214 dB re: 1 microPa(2) s. Data support an increasing energy model to predict temporary noise-induced hearing loss and indicate that odontocete noise exposure effects bear trends similar to terrestrial mammals. Thus, sonar can induce physiological and behavioural effects in at least one species of odontocete; however, exposures must be of prolonged, high sound exposures levels to generate these effects.

Noise levels in the ocean are increasing and are expected to affect marine mammals. To examine the auditory effects of noise on odontocetes, a bottlenose dolphin (Tursiops truncatus) was exposed to octave-band noise (4-8 kHz) of varying durations (<2-30 min) and sound pressures (130-178 dB re 1 microPa). Temporary threshold shift (TTS) occurrence was quantified in an effort to (i) determine the sound exposure levels (SELs) (dB re 1 microPa(2) s) that induce TTS and (ii) develop a model to predict TTS onset. Hearing thresholds were measured using auditory evoked potentials. If SEL was kept constant, significant shifts were induced by longer duration exposures but not for shorter exposures. Higher SELs were required to induce shifts in shorter duration exposures. The results did not support an equal-energy model to predict TTS onset. Rather, a logarithmic algorithm, which increased in sound energy as exposure duration decreased, was a better predictor of TTS. Recovery to baseline hearing thresholds was also logarithmic (approximately -1.8 dB/doubling of time) but indicated variability including faster recovery rates after greater shifts and longer recoveries necessary after longer duration exposures. The data reflected the complexity of TTS in mammals that should be taken into account when predicting odontocete TTS.

This is the first report of an underwater audiogram from a dolphin in a capture-and-release scenario. Two bow-riding white-beaked dolphins Lagenorhynchus albirostris (a female and a male) were captured using the hoop-net technique in Faxaflói Bay, Iceland. The dolphins were transferred to a stretcher and hoisted into a plastic research tank on board a small fishing vessel. Two underwater transducers were used to cover the frequency range from 16 to 215 kHz. Two human EEG electrodes mounted in suction cups, one placed near the blow hole and the other on the dorsal fin, picked up bioelectrical responses to acoustic stimuli. Responses to about 1000 sinusoidal amplitude modulated stimuli for each amplitude/frequency combination were averaged and analyzed using a fast Fourier transform to obtain an evoked auditory response. Threshold was defined as the zero crossing of the response using linear regression. Two threshold frequencies at 50 kHz and 64 kHz were obtained from the female. An audiogram ranging from 16 to 181 kHz was obtained from an adult male and showed the typical ;U' shaped curve for odontocetes. The thresholds for both white-beaks were comparable and demonstrated the most sensitive high frequency hearing of any known dolphin and were as sensitive as the harbor porpoise.

While there has been recent concern about the effects of sound on marine mammals, including polar bears, there are no data available measuring the hearing of any bear. The in-air hearing of three polar bears was measured using evoked auditory potentials obtained while tone pips were played to three individually anaesthetized bears at the Kolmården Djurpark. Hearing was tested in half-octave steps from 1 to 22.5 kHz. Measurements were not obtainable at 1 kHz and best sensitivity was found in the range from 11.2-22.5 kHz. Considering the tone pips were short and background noise measurements were available, absolute measurements were estimated based on an assumed mammalian integration time of 300 ms. These data show sensitive hearing in the polar bear over a wide frequency range and should cause those concerned with the introduction of anthropogenic noise into the polar bear's environment to operate with caution.

An infant Risso's dolphin (Grampus griseus) was rescued from the beach in Southern Portugal, and an audiogram was measured using auditory evoked potentials (AEP) and envelope following response (EFR) techniques for frequencies from 4 to 150 kHz. The stimuli used were custom sinusoidally amplitude-modulated (SAM) tone-bursts, and the AEP responses were collected, averaged and analyzed to quantify the animal's physiological response and, thereby, hearing thresholds. The infant animal showed a wide range of best sensitivity, with the lowest threshold of 49.5 dB re. 1 microPa at 90 kHz. The audiogram showed a typical mammalian union or logical sum-shape with a gradual, low-frequency slope of 16.4 dB octave-1 and a sharp high-frequency increase of 95 dB octave-1. When compared with an audiogram of an older Risso's dolphin obtained using behavioral methods, the threshold values at upper frequencies were much lower for this infant animal, and this infant heard higher frequencies. These results redefine the hearing capabilities of Risso's dolphins by demonstrating very high-frequency sensitivity.

Human activity is contributing increasing noise to marine ecosystems. Recent studies have examined the effects of boat noise on marine fishes, but there is limited understanding of the prevalence of this type of sound source. This investigation tracks vessel noise on three reefs in the US Virgin Islands National Park over four months in 2013. Ambient noise levels ranged from 106 to 129 dB rms re 1 μPa (100 Hz–20 kHz). Boat noise occurred in 6–12% of samples. In the presence of boat noise, ambient noise in a low-frequency band (100–1000 Hz) increased by >7 dB above baseline levels and sound levels were significantly higher. The frequency with the most acoustic energy shifted to a significantly lower frequency when boat noise was present during the day. These results indicate the abundance of boat noise and its overlap with reef organism sound production, raising concern for the communication abilities of these animals. [ABSTRACT FROM AUTHOR]

Sound is a widely available and vital cue in aquatic environments, yet most bioacoustic research has focused on marine vertebrates, leaving sound detection in invertebrates poorly understood. Cephalopods are an ecologically key taxon that likely use sound and may be impacted by increasing anthropogenic ocean noise, but little is known regarding their behavioral responses or adaptations to sound stimuli. These experiments identify the acoustic range and levels that elicit a wide range of secondary defense behaviors such as inking, jetting and rapid coloration change. Secondarily, it was found that cuttlefish habituate to certain sound stimuli. The present study examined the behavioral responses of 22 cuttlefish (Sepia officinalis) to pure-tone pips ranging from 80 to 1000 Hz with sound pressure levels of 85-188 dB re. 1 µPa rms and particle accelerations of 0-17.1 ms-2. Cuttlefish escape responses (inking, jetting) were observed between frequencies of 80 and 300 Hz and at sound levels above 140dB re. 1 pPa rms and 0.01 ms-2 (0.74ms-2 for inking responses). Body patterning changes and fin movements were observed at all frequencies and sound levels. Response intensity was dependent upon stimulus amplitude and frequency, suggesting that cuttlefish also possess loudness perception with a maximum sensitivity around 150 Hz. Cuttlefish habituated to repeated 200 Hz tone pips, at two sound intensities. Total response inhibition was not reached, however, and a basal response remained present in most animals. The graded responses provide a loudness sensitivity curve and suggest an ecological function for sound use in cephalopods. [ABSTRACT FROM AUTHOR]

While hearing is the primary sensory modality for odontocetes, there are few data addressing variation within a natural population. This work describes the hearing ranges (4-150 kHz) and sensitivities of seven apparently healthy, wild beluga whales (Delphinapterus leucas) during a population health assessment project that captured and released belugas in Bristol Bay, Alaska. The baseline hearing abilities and subsequent variations were addressed. Hearing was measured using auditory evoked potentials (AEPs). All audiograms showed a typical cetacean U-shape; substantial variation (>30 dB) was found between most and least sensitive thresholds. All animals heard well, up to at least 128 kHz. Two heard up to 150 kHz. Lowest auditory thresholds (35-45 dB) were identified in the range 45-80 kHz. Greatest differences in hearing abilities occurred at both the high end of the auditory range and at frequencies of maximum sensitivity. In general, wild beluga hearing was quite sensitive. Hearing abilities were similar to those of belugas measured in zoological settings, reinforcing the comparative importance of both settings. The relative degree of variability across the wild belugas suggests that audiograms from multiple individuals are needed to properly describe the maximum sensitivity and population variance for odontocetes. Hearing measures were easily incorporated into field-based settings. This detailed examination of hearing abilities in wild Bristol Bay belugas provides a basis for a better understanding of the potential impact of anthropogenic noise on a noise-sensitive species. Such information may help design noise-limiting mitigation measures that could be applied to areas heavily influenced and inhabited by endangered belugas. [ABSTRACT FROM AUTHOR]

How an animal receives sound may influence its use of sound. While 'jaw hearing' is well supported for odontocetes, work examining how sound is received across the head has been limited to a few representative species. The substantial variation in jaw and head morphology among odontocetes suggests variation in sound reception. Here, we address how a divergent subspecies, the Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) hears low-, mid- and high-frequency tones, as well as broadband clicks, comparing sounds presented at different locations across the head. Hearing was measured using auditory evoked potentials (AEPs). Click and tone stimuli (8, 54 and 120 kHz) were presented at nine locations on the head and body using a suction- cup transducer. Threshold differences were compared between frequencies and locations, and referenced to the underlying anatomy using computed tomography (CT) imaging of deceased animals of the same subspecies. The best hearing locations with minimum thresholds were found adjacent to a mandibular fat pad and overlaying the auditory bulla. Mean thresholds were not substantially different at locations from the rostrum tip to the ear (11.6dB). This contrasts with tests with bottlenose dolphins and beluga whales, in which 30-40 dB threshold differences were found across the animals' heads. Response latencies increased with decreasing response amplitudes, which suggests that latency and sensitivity are interrelated when considering sound reception across the odontocete head. The results suggest that there are differences among odontocetes in the anatomy related to receiving sound, and porpoises may have relatively less acoustic 'shadowing'. [ABSTRACT FROM AUTHOR]

Abstract: Odontocetes are believed to receive sounds primarily through the pan bone region of the lower jaw although much variation in jaw morphology exists among species. In order to further examine this jaw hearing hypothesis we tested the head receiving sensitivity and directional hearing of a beluga whale, Delphinapterus leucas. Hearing thresholds were measured using auditory evoked potentials (AEPs). The subject proved to have highly directional hearing for far-field click stimuli similar to that of bottlenose dolphins and more directional than the harbor porpoise. For near-field jawphone stimulation, the beluga''s lowest thresholds were found when click stimuli were presented at the rostrum tip (76 dB re: 1 μPa) although thresholds from the pan bone region stimulation were only 2–3 dB higher. Stimulation at and behind the external auditory meatus were elevated by nearly 20 dB. Stimuli presented at the surface of the melon did not generate detectable AEP responses, although sound levels of up to 142 dB were employed. Latencies of responses were generally shortest for meatal stimulation and increased with distance. Results support a shaded receiver model for odontocete hearing but how received sounds are filtered and shaded may depend on species. We also suggest that odontocete hearing thresholds are not necessarily lowest through the pan bone region. Rather, hearing pathway variations appear to exist among odontocete species and are at least partially dependent on head morphology. [Copyright &y& Elsevier]

Auditory evoked potentials (AEPs) allow researchers to measure the hearing abilities of animals that would be difficult or impossible to train for behavioral measurements of hearing. The hearing abilities of live-stranded cetaceans and wild dolphins can only be made with AEP techniques. In these situations, time with the animal is often restricted to an hour or less, and there is often little control over the acoustic environment in which the tests are performed. AEP measurements may be made while the animals are in air or in shallow pools. For cetaceans in air, sounds are typically presented with a suction cup jawphone. For cetaceans in water, sounds may be presented in a direct field (with the transducer located at some distance from the test subject) or with a jawphone. In each of these situations it is important to understand how thresholds derived from AEP measurements compare with behavioral hearing measurements. Examples of AEP measurements from wild and live-stranded cetaceans are presented to illustrate their usefulness and the constraints under which these measurements must be made. AEP measurements from bottlenose dolphins in air and in water are also compared with their behavioral audiograms. [ABSTRACT FROM AUTHOR]