Modeling of the near to far acoustic fields of an echolocating bottlenose dolphin and harbor porpoise.

Echolocation signals emitted by odontocetes can be roughly classified into three broad categories: broadband echolocation signals, narrowband high-frequency echolocation signals, and frequency modulated clicks. Previous measurements of broadband echolocation signal propagation in the bottlenose dolphin (Tursiops truncatus) did not find any evidence of focusing as the signals travel from the near-field to far-field. Finite element analysis (FEA) of high-resolution computed tomography scan data was used to examine signal propagation of broadband echolocation signals of dolphins and narrowband echolocation signals of porpoises. The FEA results were used to simulate the propagation of clicks from phonic lips, traveling through the forehead, and finally transmission into the water. Biosonar beam formation in the near-field and far-field, including the amplitude contours for the two species, was determined. The finite element model result for the simulated amplitude contour in the horizontal plane was consistent with prior direct measurement results for Tursiops, validating the model. Furthermore, the simulated far-field transmission beam patterns in both the vertical and horizontal planes were also qualitatively consistent with results measured from live animals. This study indicates that there is no evidence of convergence for either Tursiops or Phocoena as the sound propagates from the near-field to the far-field.

Lack of reproducibility of temporary hearing threshold shifts in a harbor porpoise after exposure to repeated airgun sounds.

Noise-induced temporary hearing threshold shift (TTS) was studied in a harbor porpoise exposed to impulsive sounds of scaled-down airguns while both stationary and free-swimming for up to 90 min. In a previous study, ∼4 dB TTS was elicited in this porpoise, but despite 8 dB higher single-shot and cumulative exposure levels (up to 199 dB re 1 µPa2s) in the present study, the porpoise showed no significant TTS at hearing frequencies 2, 4, or 8 kHz. There were no changes in the study animal's audiogram between the studies or significant differences in the fatiguing sound that could explain the difference, but audible and visual cues in the present study may have allowed the porpoise to predict when the fatiguing sounds would be produced. The discrepancy between the studies may have resulted from self-mitigation by the porpoise. Self-mitigation, resulting in reduced hearing sensitivity, can be achieved via changes in the orientation of the head, or via alteration of the hearing threshold by processes in the ear or central nervous system.

A model and experimental approach to the middle ear transfer function related to hearing in the humpback whale (Megaptera novaeangliae).

At present, there are no direct measures of hearing for any baleen whale (Mysticeti). The most viable alternative to in vivo approaches to simulate the audiogram is through modeling outer, middle, and inner ear functions based on the anatomy and material properties of each component. This paper describes a finite element model of the middle ear for the humpback whale (Megaptera novaeangliae) to calculate the middle ear transfer function (METF) to determine acoustic energy transmission to the cochlea. The model was developed based on high resolution computed tomography imaging and direct anatomical measurements of the middle ear components for this mysticete species. Mechanical properties for the middle ear tissues were determined from experimental measurements and published values. The METF for the humpback whale predicted a better frequency range between approximately 15 Hz and 3 kHz or between 200 Hz and 9 kHz based on two potential stimulation locations. Experimental measures of the ossicular chain, tympanic membrane, and tympanic bone velocities showed frequency response characteristics consistent with the model. The predicted best sensitivity hearing ranges match well with known vocalizations of this species.

Finite element simulation of broadband biosonar signal propagation in the near- and far-field of an echolocating Atlantic bottlenose dolphin (Tursiops truncatus).

Bottlenose dolphins project broadband echolocation signals for detecting and locating prey and predators, and for spatial orientation. There are many unknowns concerning the specifics of biosonar signal production and propagation in the head of dolphins and this manuscript represents an effort to address this topic. A two-dimensional finite element model was constructed using high resolution CT scan data. The model simulated the acoustic processes in the vertical plane of the biosonar signal emitted from the phonic lips and propagated into the water through the animal's head. The acoustic field on the animal's forehead and the farfield transmission beam pattern of the echolocating dolphin were determined. The simulation results and prior acoustic measurements were qualitatively extremely consistent. The role of the main structures on the sound propagation pathway such as the air sacs, melon, and connective tissue was investigated. Furthermore, an investigation of the driving force at the phonic lips for dolphins that emit broadband echolocation signals and porpoises that emit narrowband echolocation signals suggested that the driving force is different for the two types of biosonar. Finally, the results provide a visual understanding of the sound transmission in dolphin's biosonar.

Biosonar signal propagation in the harbor porpoise's (Phocoena phocoena) head: The role of various structures in the formation of the vertical beam.

Harbor porpoises (Phocoena phocoena) use narrow band echolocation signals for detecting and locating prey and for spatial orientation. In this study, acoustic impedance values of tissues in the porpoise's head were calculated from computer tomography (CT) scan and the corresponding Hounsfield Units. A two-dimensional finite element model of the acoustic impedance was constructed based on CT scan data to simulate the acoustic propagation through the animal's head. The far field transmission beam pattern in the vertical plane and the waveforms of the receiving points around the forehead were compared with prior measurement results, the simulation results were qualitatively consistent with the measurement results. The role of the main structures in the head such as the air sacs, melon and skull in the acoustic propagation was investigated. The results showed that air sacs and skull are the major components to form the vertical beam. Additionally, both beam patterns and sound pressure of the sound waves through four positions deep inside the melon were demonstrated to show the role of the melon in the biosonar sound propagation processes in the vertical plane.

Discovery of Sound in the Sea: Resources for Educators, Students, the Public, and Policymakers.

There is increasing concern about the effects of underwater sound on marine life. However, the science of sound is challenging. The Discovery of Sound in the Sea (DOSITS) Web site ( http://www.dosits.org ) was designed to provide comprehensive scientific information on underwater sound for the public and educational and media professionals. It covers the physical science of underwater sound and its use by people and marine animals for a range of tasks. Celebrating 10 years of online resources, DOSITS continues to develop new material and improvements, providing the best resource for the most up-to-date information on underwater sound and its potential effects.

Hearing abilities and sound reception of broadband sounds in an adult Risso's dolphin (Grampus griseus).

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.

A comparison of auditory brainstem responses across diving bird species.

There is little biological data available for diving birds because many live in hard-to-study, remote habitats. Only one species of diving bird, the black-footed penguin (Spheniscus demersus), has been studied in respect to auditory capabilities (Wever et al., Proc Natl Acad Sci USA 63:676-680, 1969). We, therefore, measured in-air auditory threshold in ten species of diving birds, using the auditory brainstem response (ABR). The average audiogram obtained for each species followed the U-shape typical of birds and many other animals. All species tested shared a common region of the greatest sensitivity, from 1000 to 3000 Hz, although audiograms differed significantly across species. Thresholds of all duck species tested were more similar to each other than to the two non-duck species tested. The red-throated loon (Gavia stellata) and northern gannet (Morus bassanus) exhibited the highest thresholds while the lowest thresholds belonged to the duck species, specifically the lesser scaup (Aythya affinis) and ruddy duck (Oxyura jamaicensis). Vocalization parameters were also measured for each species, and showed that with the exception of the common eider (Somateria mollisima), the peak frequency, i.e., frequency at the greatest intensity, of all species' vocalizations measured here fell between 1000 and 3000 Hz, matching the bandwidth of the most sensitive hearing range.

Hearing pathways in the Yangtze finless porpoise, Neophocaena asiaeorientalis asiaeorientalis.

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 (11.6 dB). 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'.

Does rotation increase the acoustic field of view? Comparative models based on CT data of a live dolphin versus a dead dolphin.

Rotational behaviour has been observed when dolphins track or detect targets, however, its role in echolocation is unknown. We used computed tomography data of one live and one recently deceased bottlenose dolphin, together with measurements of the acoustic properties of head tissues, to perform acoustic property reconstruction. The anatomical configuration and acoustic properties of the main forehead structures between the live and deceased dolphins were compared. Finite element analysis (FEA) was applied to simulate the generation and propagation of echolocation clicks, to compute their waveforms and spectra in both near- and far-fields, and to derive echolocation beam patterns. Modelling results from both the live and deceased dolphins were in good agreement with click recordings from other, live, echolocating individuals. FEA was also used to estimate the acoustic scene experienced by a dolphin rotating 180° about its longitudinal axis to detect fish in the far-field at elevation angles of -20° to 20°. The results suggest that the rotational behaviour provides a wider insonification area and a wider receiving area. Thus, it may provide compensation for the dolphin's relatively narrow biosonar beam, asymmetries in sound reception, and constraints on the pointing direction that are limited by head movement. The results also have implications for examining the accuracy of FEA in acoustic simulations using recently deceased specimens.

A prediction of the minke whale (Balaenoptera acutorostrata) middle-ear transfer function.

The lack of baleen whale (Cetacea Mysticeti) audiograms impedes the assessment of the impacts of anthropogenic noise on these animals. Estimates of audiograms, which are difficult to obtain behaviorally or electrophysiologically for baleen whales, can be made by simulating the audiogram as a series of components representing the outer, middle, and inner ear (Rosowski, 1991; Ruggero and Temchin, 2002). The middle-ear portion of the system can be represented by the middle-ear transfer function (METF), a measure of the transmission of acoustic energy from the external ear to the cochlea. An anatomically accurate finite element model of the minke whale (Balaenoptera acutorostrata) middle ear was developed to predict the METF for a mysticete species. The elastic moduli of the auditory ossicles were measured by using nanoindentation. Other mechanical properties were estimated from experimental stiffness measurements or from published values. The METF predicted a best frequency range between approximately 30 Hz and 7.5 kHz or between 100 Hz and 25 kHz depending on stimulation location. Parametric analysis found that the most sensitive parameters are the elastic moduli of the glove finger and joints and the Rayleigh damping stiffness coefficient ß. The predicted hearing range matches well with the vocalization range.

Sonar-induced pressure fields in a post-mortem common dolphin.

Potential physical effects of sonar transmissions on marine mammals were investigated by measuring pressure fields induced in a 119-kg, 211-cm-long, young adult male common dolphin (Delphinus delphis) cadaver. The specimen was instrumented with tourmaline acoustic pressure gauges used as receiving sensors. Gauge implantation near critical tissues was guided by intraoperative, high-resolution, computerized tomography (CT) scanning. Instrumented structures included the melon, nares, ear, thoracic wall, lungs, epaxial muscle, and lower abdomen. The specimen was suspended from a frame equipped with a standard 50.8-mm-diameter spherical transducer used as the acoustic source and additional receiving sensors to monitor the transmitted and external, scattered field. Following immersion, the transducer transmitted pulsed sinusoidal signals at 5, 7, and 10 kHz. Quantitative internal pressure fields are reported for all cases except those in which the gauge failed or no received signal was detected. A full necropsy was performed immediately after the experiment to examine instrumented areas and all major organs. No lesions attributable to acoustic transmissions were found, consistent with the low source level and source-receiver distances.

Morphology and unbiased stereology of the lateral superior olive in the short-beaked common dolphin, Delphinus delphis (Cetacea, Delphinidae).

In all mammals, the superior olivary complex (SOC) comprises a group of auditory brainstem nuclei that are important for sound localization. Its principal nuclei, the lateral superior olive (LSO) and the medial superior olive (MSO) process interaural time and intensity differences, which are the main cues for sound localization in the horizontal plane. Toothed whales (odontocetes) rely heavily on hearing and echolocation for foraging, orientation, and communication and localize sound with great acuity. The investigation of the SOC in odontocetes provides insight into adaptations to underwater hearing and echolocation. However, quantitative anatomical data for odontocetes are currently lacking. We quantified the volume, total neuron number, and neuron density of the LSO of six common dolphins (Delphinus delphis) using the Cavalieri principle and the unbiased stereology optical fractionator. Our results show that the LSO in D. delphis has a volume of 150 + (SD = 27) mm3 , which is on average 69 (SEM = 19) times larger than the LSO in human, or 37 (SEM = 11) times larger than the human LSO and MSO combined. The LSO of D. delphis contains 20,876 ± (SD = 3300) neurons. In comparison, data reported for the human brainstem indicate the LSO has only about » that number but about the same number for the LSO and MSO combined (21,100). LSO neurons range from 21 to 25 µm (minor axis) and from 44 to 61 µm (major axis) in transverse sections. The LSO neuron packing density is 1080 ± (SD = 204) neurons/mm3 , roughly half of the LSO neuron density in human. SMI-32-immunohistochemistry was used to visualize projection neurons in the LSO and revealed the presence of principal, marginal, and multipolar neurons in transverse sections. The distinct morphology of the LSO likely reflects the common dolphin's superb sensitivity to ultra-high frequencies and ability to detect and analyze sounds and their location as part of its underwater spatial localization and echolocation tasks.

Specializations of somatosensory innervation in the skin of humpback whales (Megaptera novaeangliae).

Cetacean behavior and life history imply a role for somatosensory detection of critical signals unique to their marine environment. As the sensory anatomy of cetacean glabrous skin has not been fully explored, skin biopsy samples of the flank skin of humpback whales were prepared for general histological and immunohistochemical (IHC) analyses of innervation in this study. Histology revealed an exceptionally thick epidermis interdigitated by numerous, closely spaced long, thin diameter penicillate dermal papillae (PDP). The dermis had a stratified organization including a deep neural plexus (DNP) stratum intermingled with small arteries that was the source of intermingled nerves and arterioles forming a more superficial subepidermal neural plexus (SNP) stratum. The patterns of nerves branching through the DNP and SNP that distribute extensive innervation to arteries and arterioles and to the upper dermis and PDP provide a dense innervation associated through the whole epidermis. Some NF-H+ fibers terminated at the base of the epidermis and as encapsulated endings in dermal papillae similar to Merkel innervation and encapsulated endings seen in terrestrial mammals. However, unlike in all mammalian species assessed to date, an unusual acellular gap was present between the perineural sheaths and the central core of axons in all the cutaneous nerves perhaps as mechanism to prevent high hydrostatic pressure from compressing and interfering with axonal conductance. Altogether the whale skin has an exceptionally dense low-threshold mechanosensory system innervation most likely adapted for sensing hydrodynamic stimuli, as well as nerves that can likely withstand high pressure experienced during deep dives.

Auditory temporal resolution and evoked responses to pulsed sounds for the Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis).

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.

The influence of cochlear shape on low-frequency hearing.

The conventional theory about the snail shell shape of the mammalian cochlea is that it evolved essentially and perhaps solely to conserve space inside the skull. Recently, a theory proposed that the spiral's graded curvature enhances the cochlea's mechanical response to low frequencies. This article provides a multispecies analysis of cochlear shape to test this theory and demonstrates that the ratio of the radii of curvature from the outermost and innermost turns of the cochlear spiral is a significant cochlear feature that correlates strongly with low-frequency hearing limits. The ratio, which is a measure of curvature gradient, is a reflection of the ability of cochlear curvature to focus acoustic energy at the outer wall of the cochlear canal as the wave propagates toward the apex of the cochlea.

Possible limitations of dolphin echolocation: a simulation study based on a cross-modal matching experiment.

Dolphins use their biosonar to discriminate objects with different features through the returning echoes. Cross-modal matching experiments were conducted with a resident bottlenose dolphin (Tursiops aduncus). Four types of objects composed of different materials (water-filled PVC pipes, air-filled PVC pipes, foam ball arrays, and PVC pipes wrapped in closed-cell foam) were used in the experiments, respectively. The size and position of the objects remained the same in each case. The data collected in the experiment showed that the dolphin's matching accuracy was significantly different across the cases. To gain insight into the underlying mechanism in the experiments, we used finite element methods to construct two-dimensional target detection models of an echolocating dolphin in the vertical plane, based on computed tomography scan data. The acoustic processes of the click's interaction with the objects and the surrounding media in the four cases were simulated and compared. The simulation results provide some possible explanations for why the dolphin performed differently when discriminating the objects that only differed in material composition in the previous matching experiments.

Sound detection by the longfin squid (Loligo pealeii) studied with auditory evoked potentials: sensitivity to low-frequency particle motion and not pressure.

Although hearing has been described for many underwater species, there is much debate regarding if and how cephalopods detect sound. Here we quantify the acoustic sensitivity of the longfin squid (Loligo pealeii) using near-field acoustic and shaker-generated acceleration stimuli. Sound field pressure and particle motion components were measured from 30 to 10,000 Hz and acceleration stimuli were measured from 20 to 1000 Hz. Responses were determined using auditory evoked potentials (AEPs) with electrodes placed near the statocysts. Evoked potentials were generated by both stimuli and consisted of two wave types: (1) rapid stimulus-following waves, and (2) slower, high-amplitude waves, similar to some fish AEPs. Responses were obtained between 30 and 500 Hz with lowest thresholds between 100 and 200 Hz. At the best frequencies, AEP amplitudes were often >20 µV. Evoked potentials were extinguished at all frequencies if (1) water temperatures were less than 8°C, (2) statocysts were ablated, or (3) recording electrodes were placed in locations other than near the statocysts. Both the AEP response characteristics and the range of responses suggest that squid detect sound similarly to most fish, with the statocyst acting as an accelerometer through which squid detect the particle motion component of a sound field. The modality and frequency range indicate that squid probably detect acoustic particle motion stimuli from both predators and prey as well as low-frequency environmental sound signatures that may aid navigation.