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The plainfin midshipman fish, Porichthys notatus, is a nocturnal marine teleost that uses social acoustic signals for communication during the breeding season. Nesting type I males produce multiharmonic advertisement calls by contracting their swim bladder sonic muscles to attract females for courtship and spawning while subsequently attracting cuckholding type II males. Here, we report intra- and intersexual dimorphisms of the swim bladder in a vocal teleost fish and detail the swim bladder dimorphisms in the three sexual phenotypes (females, type I and II males) of plainfin midshipman fish. Micro-computerized tomography revealed that females and type II males have prominent, horn-like rostral swim bladder extensions that project toward the inner ear end organs (saccule, lagena, and utricle). The rostral swim bladder extensions were longer, and the distance between these swim bladder extensions and each inner-ear end organ type was significantly shorter in both females and type II males compared to that in type I males. Our results revealed that the normalized swim bladder length of females and type II males was longer than that in type I males while there was no difference in normalized swim bladder width among the three sexual phenotypes. We predict that these intrasexual and intersexual differences in swim bladder morphology among midshipman sexual phenotypes will afford greater sound pressure sensitivity and higher frequency detection in females and type II males and facilitate the detection and localization of conspecifics in shallow water environments, like those in which midshipman breed and nest.

Previous work on auditory processing in Opsanus tau has focused on the descending octaval nucleus; however, the magnocellular octaval nucleus receives similar inputs from the otolithic endorgans. The purpose of this study was to assess whether cells in any of the three subdivisions of the magnocellular nucleus respond to auditory frequencies and encode sound source direction. Extracellular recording sites were chosen based on anatomical landmarks, and neurobiotin injections confirmed the location of auditory sites in subdivisions of the magnocellular nucleus. In general, the auditory cells in M2 and M3 responded best to frequencies at or below 100 Hz. Most auditory cells responded well to directional stimuli presented along axes in the horizontal plane. Cells in M3 (not M2) also responded to lateral line stimulation, consistent with otolithic endorgan and lateral line inputs to M3. The convergence of auditory and lateral line inputs in M3, the lack of Mauthner cells in this species, and previous evidence that the magnocellular nucleus does not contribute to ascending auditory pathways suggest to us that the large cells of M3 may play a role in rapid behavioral responses to particle motion stimuli in oyster toadfish.

Georg von Békésy was one of the first comparative auditory researchers. He not only studied basilar membrane (BM) movements in a range of mammals of widely different sizes, he also worked on the chicken basilar papilla and the frog middle ear. We show that, in mammals, at least, his data do not differ from those that could be collected using modern techniques but with the same, very loud sounds. There is in all cases a major difference to frequency maps collected using low-level sounds. In contrast, the same cannot be said of his chicken data, perhaps due to the different roles played by the BM in mammals and birds. In lizards, the BM is not tuned and it is perhaps good that Békésy did not begin with those species and get discouraged in his seminal comparative work.

Auditory evoked potentials (AEPs) have become popular for estimating hearing thresholds and audiograms. What is the utility of these measurements? How do AEP audiograms compare with behavioral audiograms? In general, AEP measurements for fishes and marine mammals often underestimate behavioral thresholds, but comparisons are especially complicated when the AEP and behavioral measures are obtained under different acoustic conditions. There is no single representative relationship between AEP and behavioral audiograms and these audiograms should not be considered equivalent. We suggest that the most valuable comparisons are those made by the same researcher using similar acoustic conditions for both measurements.

The lake sturgeon, Acipenser fulvescens, belongs to one of the few extant nonteleost ray-finned fishes and diverged from the main vertebrate lineage about 250 million years ago. The aim of this study was to use this species to explore the peripheral neural coding strategies for sound direction and compare these results to modern bony fishes (teleosts). Extracellular recordings were made from afferent neurons innervating the saccule and lagena of the inner ear while the fish was stimulated using a shaker system. Afferents were highly directional and strongly phase locked to the stimulus. Directional response profiles resembled cosine functions, and directional preferences occurred at a wide range of stimulus intensities (spanning at least 60 dB re 1 nm displacement). Seventy-six percent of afferents were directionally selective for stimuli in the vertical plane near 90° (up down) and did not respond to horizontal stimulation. Sixty-two percent of afferents responsive to horizontal stimulation had their best axis in azimuths near 0° (front back). These findings suggest that in the lake sturgeon, in contrast to teleosts, the saccule and lagena may convey more limited information about the direction of a sound source, raising the possibility that this species uses a different mechanism for localizing sound. For azimuth, a mechanism could involve the utricle or perhaps the computation of arrival time differences. For elevation, behavioral strategies such as directing the head to maximize input to the area of best sensitivity may be used. Alternatively, the lake sturgeon may have a more limited ability for sound source localization compared with teleosts.

SUMMARY Acipenser fulvescens, the lake sturgeon, belongs to one of the few extant non-teleost ray-finned (bony) fishes. The sturgeons (family Acipenseridae) have a phylogenetic history that dates back about 250 million years. The study reported here is the first investigation of peripheral coding strategies for spectral analysis in the auditory system in a non-teleost bony fish. We used a shaker system to simulate the particle motion component of sound during electrophysiological recordings of isolated single units from the eighth nerve innervating the saccule and lagena. Background activity and response characteristics of saccular and lagenar afferents (such as thresholds, response–level functions and temporal firing) resembled the ones found in teleosts. The distribution of best frequencies also resembled data in teleosts (except for Carassius auratus, goldfish) tested with the same stimulation method. The saccule and lagena in A. fulvescens contain otoconia, in contrast to the solid otoliths found in teleosts, however, this difference in otolith structure did not appear to affect threshold, frequency tuning, intensity- or temporal responses of auditory afferents. In general, the physiological characteristics common to A. fulvescens, teleosts and land vertebrates reflect important functions of the auditory system that may have been conserved throughout the evolution of vertebrates.

Binaural computations involving the convergence of excitatory and inhibitory inputs have been proposed to explain directional sharpening and frequency tuning documented in the brainstem of a teleost fish, the oyster toadfish (Opsanus tau). To assess the presence of inhibitory neurons in the ascending auditory circuit, we used a monoclonal antibody to GABA to evaluate immunoreactivity at three levels of the circuit: the first order descending octaval nucleus (DON), the secondary octaval population (dorsal division), and the midbrain torus semicircularis. We observed a subset of immunoreactive (IR) cells and puncta distributed throughout the neuropil at all three locations. To assess whether contralateral inhibition is present, fluorescent dextran crystals were inserted into dorsal DON to fill contralateral, commissural inputs retrogradely prior to GABA immunohistochemistry. GABA-IR somata and puncta co-occurred with retrogradely filled, GABA-negative auditory projection cells. GABA-IR projection cells were more common in the dorsolateral DON than in the dorsomedial DON, but GABA-IR puncta were common in both dorsolateral and dorsomedial divisions. Our findings demonstrate that GABA is present in the ascending auditory circuit in the brainstem of the toadfish, indicating that GABA-mediated inhibition participates in shaping auditory response characteristics in a teleost fish as in other vertebrates.

SUMMARYComparisons of left and right auditory input are required for sound source localization in most terrestrial vertebrates. Previous physiological and neuroanatomical studies have indicated that binaural convergence is present in the ascending auditory system of the toadfish. In this study, we introduce a new technique, otolith tipping, to reversibly alter directional auditory input to the central nervous system of a fish. The normal directional response pattern (DRP) was recorded extracellularly for auditory cells in the first-order descending octaval nucleus (DON) or the midbrain torus semicircularis (TS) using particle motion stimuli in the horizontal and mid-sagittal planes. The same stimuli were used during tipping of the saccular otolith to evaluate changes in the DRPs. Post-tipping DRPs were generated and compared with the pre-tipping DRPs to ensure that the data had been collected consistently from the same unit. In the DON, ipsilateral or contralateral tipping most often eliminated spike activity, but changes in spike rate(±) and DRP shape were also documented. In the TS, tipping most often caused a change in spike rate (±) and altered the shape or best axis of the DRP. The data indicate that there are complex interactions of excitatory and inhibitory inputs in the DON and TS resulting from the convergence of binaural inputs. As in anurans, but unlike other terrestrial vertebrates,binaural processing associated with encoding the direction of a sound source begins in the first-order auditory nucleus of this teleost.

SUMMARY Zebrafish (Danio rerio) were placed in small wells that could be driven vertically with a series of calibrated sinusoids. Video images of the fish were obtained and analyzed to determine the levels and frequencies at which the fish responded to the stimulus tones. It was found that fish 4 days post fertilization (dpf) did not respond to the stimulus tones, whereas fish 5 dpf to adult did respond. It was further found that the stimulus thresholds and frequency bandwidth to which the fish responded did not change from 5 dpf to adult; indicating that the otolithic organ adaptations for high-frequency hearing are already present in larval fish. Deflating the swimbladders in adult fish eliminated their response, which is consistent with sensing sound pressure. Deflating the swimbladder in larval fish did not affect their thresholds, which is consistent with sensing the particle motion of the fluid directly. Because adult fish with Weberian ossicles have a greater input to the inner ear for a given sound pressure level (SPL), the finding that the adult and larval fish respond at the same SPL with intact swimbladders suggests that the acoustic startle response threshold is adjusted as the fish develop in order to maintain appropriate reactions to relevant stimuli.

Bimodal cells in the torus semicircularis of the toadfish respond to both directional acoustic stimuli and hydrodynamic stimuli. Our previous physiological work indicated that bimodal cells may be distributed throughout the torus semicircularis. In this study, neurobiotin was used to compare the distribution of auditory-only and bimodal sites and to assess the inputs to those sites. A brief neurobiotin injection with short survival time was used to document the recording location. In other fish, a longer injection and survival time was used at an auditory-only or a bimodal site to fill the axons of the medullary inputs. Auditory-only sites were located in the most dorsal and medial sites in nucleus centralis. Bimodal sites were identified within both nucleus centralis and nucleus ventrolateralis. The greatest number of retrogradely filled cell bodies was found in the descending octaval nucleus following injection at auditory-only recording sites in nucleus centralis. In contrast, retrogradely filled cell bodies were found in both the descending octaval nucleus and the lateral line nucleus medialis following injection at bimodal sites in nucleus centralis or nucleus ventrolateralis. Retrogradely filled cell bodies were located in the dorsal and ventral divisions of the secondary octaval population from injections at either bimodal or auditory-only sites. The secondary octaval population has been implicated in auditory processing based on previous studies of both auditory specialist and auditory generalist fishes; however, this study is the first to reveal the potential role of the secondary octaval population in directional hearing in a fish. Relatively large numbers of retrogradely filled cells around the lateral lemniscus at consistent locations in the medulla indicate that a perilemniscal cell group also might be a component of the directional hearing circuit.

Single-unit recordings were made from areas in the midbrain (torus semicircularis) of the oyster toadfish. We evaluated frequency tuning and directional responses using whole-body oscillation to simulate auditory stimulation by particle motion along axes in the horizontal and mid-sagittal planes. We also tested for bimodality in responses to auditory and hydrodynamic stimuli. One recording location in each animal was marked by a neurobiotin injection to confirm the recording site. Recordings were made in nucleus centralis, nucleus ventrolateralis, and the deep cell layer. Most units were frequency-selective with best frequencies between 50 and 141 Hz. Suppression of activity was apparent in 10% of the cells. Bimodality was common, including inhibition and suppression of background activity by auditory or hydrodynamic stimulation. The majority of the cells were directionally selective with directional response patterns that were sharpened compared with those of primary saccular afferents. The best directional axes were arrayed widely in spherical space, covering most azimuths and elevations. This representation is adequate for the computation of the motional axis of an auditory stimulus for sound source localization.

While particle motion is thought to directly stimulate the inner ear of most fish species, it is difficult to measure and might not be predictable from pressure measurements in a small tank. It is therefore important to replicate experiments conducted relative to pressure measurements using stimuli of known particle motion, to ensure that unmeasured components of the stimulus field do not produce misleading frequency response profiles. The frequency sensitivity of the inner ear of the plainfin midshipman fish, Porichthys notatus, in response to isopressure stimuli has been described. This study now examines the frequency and directional response properties of midshipman saccular afferents in response to whole-body displacements simulating acoustic particle motion. Best frequencies were distributed bimodally, with peaks at 50 Hz and 100 Hz. Most units had cosinusoidally shaped directional response profiles in the horizontal and vertical planes, though some units showed slight deviations from this pattern. A few units (probably saccular efferents) had omnidirectional directional response profiles and did not phase lock to the stimulus waveform. These results are consistent with responses of the midshipman saccular nerve to isopressure stimuli, and strengthen the hypothesis that the frequency sensitivity of the midshipman ear matches the frequency content of behaviorally relevant vocalizations.

This article provides an overview to articles on fish bioacoustics – the study of hearing, sound production, communication, and the lateral line of fishes. Since bioacoustics includes the ear, this section also includes articles on the anatomy of the ear as well as the other important role of the ear beyond hearing, the vestibular sense.

The neurocranium of the toadfish (Opsanus tau) exhibits a distinct translucent region in the otic capsule (OC) that may have functional significance for the auditory pathway. This study used ultrahigh resolution computerized tomography (100 µm voxels) to compare the relative density of three sites along the OC (dorsolateral, midlateral, and ventromedial) and two reference sites (dorsal: supraoccipital crest; ventral: parasphenoid bone) in the neurocranium. Higher attenuation occurs where structural density is greater; thus, we compared the X-ray attenuations measured, which provided a measure of relative density. The maximum attenuation value was recorded for each of the five sites (x and y) on consecutive sections throughout the OC and for each of the three calcareous otoliths associated with the sensory maculae (lagena, saccule, and utricle) in the OC. All three otoliths had higher attenuations than any sites in the neurocranium. Both dorsal and ventral reference sites (supraoccipital crest and parasphenoid bone, respectively) had attenuation levels consistent with calcified bone and had relatively small, irregular variations along the length of the OC in all individuals. The lowest relative attenuations (lowest densities) occurred consistently at the three sites along the OC. In addition, the lowest attenuations measured along the OC occurred at the ventromedial site around the saccular otolith for all seven fish. The decrease in bone density along the OC is consistent with the hypothesis that there is a low-density channel in the skull to facilitate transmission of acoustic stimuli to the auditory endorgans of the ear.

Chapters cannot be read stand-alone. Please see complete SpringerBrief at: http://link.springer.com/book/10.1007/978-3-319-06659-2.

Chapters cannot be read stand-alone. Please see complete SpringerBrief at: http://link.springer.com/book/10.1007/978-3-319-06659-2.

This chapter chronicles the evolution of experiments from 1966 through 2011 that led to determining what fish listen to. By 2009, it became clear that goldfish are listening to the soundscape, like most other animals. This conclusion is based on three conceptions: (1) the fish brain is similar in morphotype to all vertebrate brains; (2) the goldfish, at least, has the processing power to analyze complex soundscapes; and (3) the processing of fish vocalizations couldn’t be a determining factor in the evolution of fish hearing. Goldfish possess the sense of hearing required to process the various sources likely making up a fish’s soundscape. They seem to be able to process the acoustic spectrum. They can process reverberations and reflections and various temporal patterns, and some species can locate the sources of sound using binaural hearing. Suggested here, then, is the notion of environmental soundscapes as the most probably important sources of information to organisms. Environmental information exists to be exploited for appropriate behavior with respect to audible sound sources and events, and fish have the capacity to exploit it for general orientation. This is what fish and other species listen to.

Chapters cannot be read stand-alone. Please see complete SpringerBrief at: http://link.springer.com/book/10.1007/978-3-319-06659-2.

We investigated the roles of the swim bladder and the lateral line system in sound localization behavior by the plainfin midshipman fish (Porichthys notatus). Reproductive female midshipman underwent either surgical deflation of the swim bladder or cryoablation of the lateral line and were then tested in a monopolar sound source localization task. Fish with nominally "deflated" swim bladders performed similar to sham-deflated controls; however, post-experiment evaluation of swim bladder deflation revealed that a majority of "deflated" fish (88%, 7 of the 8 fish) that exhibited positive phonotaxis had partially inflated swim bladders. In total, 95% (21/22) of fish that localized the source had at least partially inflated swim-bladders, indicating that pressure reception is likely required for sound source localization. In lateral line experiments, no difference was observed in the proportion of females exhibiting positive phonotaxis with ablated- (37%) versus sham-ablated (47%) lateral line systems. These data suggest that the lateral line system is likely not required for sound source localization, although this system may be important for fine-tuning the approach to the sound source. We found that midshipman can solve the 180° ambiguity of source direction in the shallow water of our test tank, which is similar to their nesting environment. We also found that the potential directional cues (phase relationship between pressure and particle motion) in shallow water differs from a theoretical free-field. Therefore, the general question of how fish use acoustic pressure cues to solve the 180° ambiguity of source direction from the particle motion vector remains unresolved.

The Gems of the Past: A Brief History of Lateral Line Research in the Context of the Hearing Sciences.- Morphological Diversity, Development, and Evolution of the Mechanosensory Lateral Line System.- The Hydrodynamic of Flow Stimuli.- The Biophysics of the Fish Lateral Line.- Sensory Ecology and Neuroethology of the Lateral Line.- Information Encoding and Processing by the Peripheral Lateral Line System.- The Central Nervous Organization of the Lateral Line System.- Central Processing of Lateral Line Information.- Functional Overlap and Nonoverlap Between Lateral Line and Auditory Systems.- The Hearing Loss, Protection, and Regeneration in the Larval Zebrafish Lateral Line.

This paper considers aspects of the evolution of the vertebrate auditory system from an 'ichthyocentric' perspective. It is argued that all vertebrate auditory systems are required to do certain basic tasks including acoustic feature discrimination, sound source localization, frequency analysis, and auditory scene analysis, among others. These sorts of capabilities arose very early in the evolution of the vertebrates and have been modified by selection in different species. In some cases the same structures have been involved in detection and analysis throughout the vertebrates, while in other cases the mechanism by which the same type of analysis takes place may have changed.

This paper reviews and discusses several investigations of the peripheral neural code for the directional axis of acoustical particle motion in the saccule of two fishes: goldfish ( Carassius auratus ) and toadfish ( Opsanus tau ). Most saccular afferents are directional in the manner of hair cells, having a cosine–shaped directional response pattern. The saccular sensory epithelia are orientated almost vertically in a parasagittal plane. In the horizontal plane, these epithelia are orientated obliquely with respect to the midline. Hair–cell stereocilia project perpendicularly. Thus, directional response patterns of saccular afferents tend to be orientated in azimuth parallel to the orientation of the epithelia in the head. The oblique angle of the toadfish saccule is greater than that of the goldfish, and the range of best directions in the horizontal plane for each species reflects those differing orientations. The azimuth of acoustical particle motion could be computed by comparing the relative activation of the two saccules, as is the case for the ears of most terrestrial vertebrates. The spatial patterns of saccular hair–cell orientation of most fishes thus appear to have little function in azimuthal source location, but for toadfish are probably most important for determining the elevation of monopole sources.

Neurobiotin was injected iontophoretically into saccular afferents of toadfish (Opsanus tau) after intracellular recording to examine dendritic arbors and central projections with respect to the physiological and directional response properties of the cells. Dendritic arbors of 36 afferents were examined in detail. Maximum diameter of the arbor and the number of terminal points were positively correlated with each other, but neither was predictive of spontaneous activity or sensitivity. Best azimuths were centered around 30 degrees -40 degrees, which corresponds to the angle of the saccule with respect to the fish's midline. In general, best elevations for afferents corresponded to hair cell orientations in the region innervated; unexpectedly low elevations obtained from afferents innervating the middle saccule may reflect curvature of the sensory epithelium against the otolith. Three efferent cells were filled partially. The location and large size of the efferent projections indicate that activity along the saccule could be modulated by a single efferent. All afferents projected to the dorsal zone of the descending octaval nucleus (dDON); many afferents bifurcated to terminate in the anterior octaval nucleus, and a few of those also had terminal fields in the medial zone of DON. All afferent projections into the dDON consisted of multiple axon collaterals projecting to numerous sites along the rostral-caudal extent of the nucleus. Variation in terminal field sites also was noted in the medial to lateral axis of the dDON; however, there were no consistent correlations between terminal field locations, physiology, and best directions of the saccular afferents.

Introduction and overview.- Neuronal response properties and voltage-gated ion channels.- The hair cell synapse.- The endbulbs of Held.- The calyces of Held.- Synaptic coincidence detection.- Synaptic inhibition in the auditory system.- Modulatory mechanisms for controlling auditory processing.- Long-term plasticity in the auditory system.

SUMMARY Sound-source localization behavior was studied in the plainfin midshipman fish (Porichthys notatus) by making use of the naturally occurring phonotaxis response of gravid females to playback of the male's advertisement call. The observations took place outdoors in a circular concrete tank. A dipole sound projector was placed at the center of the tank and an 80–90 Hz tone (the approximate fundamental frequency to the male's advertisement call) was broadcast to gravid females that were released from alternative sites approximately 100 cm from the source. The phonotaxic responses of females to the source were recorded, analyzed and compared with the sound field. One release site was approximately along the vibratory axis of the dipole source, and the other was approximately orthogonal to the vibratory axis. The sound field in the tank was fully characterized through measurements of the sound pressure field using hydrophones and acoustic particle motion using an accelerometer. These measurements confirmed that the sound field was a nearly ideal dipole. When released along the dipole vibratory axis, the responding female fish took essentially straight paths to the source. However, when released approximately 90 deg to the source's vibratory axis, the responding females took highly curved paths to the source that were approximately in line with the local particle motion axes. These results indicate that the acoustic cues used by fish during sound-source localization include the axes of particle motion of the local sound field.

Underwater soundscapes have probably played an important role in the adaptation of ears and auditory systems of fishes throughout evolutionary time, and for all species. These sounds probably contain important information about the environment and about most objects and events that confront the receiving fish so that appropriate behavior is possible. For example, the sounds from reefs appear to be used by at least some fishes for their orientation and migration. These sorts of environmental sounds should be considered much like "acoustic daylight," that continuously bathes all environments and contain information that all organisms can potentially use to form a sort of image of the environment. At present, however, we are generally ignorant of the nature of ambient sound fields impinging on fishes, and the adaptive value of processing these fields to resolve the multiple sources of sound. Our field has focused almost exclusively on the adaptive value of processing species-specific communication sounds, and has not considered the informational value of ambient "noise." Since all fishes can detect and process acoustic particle motion, including the directional characteristics of this motion, underwater sound fields are potentially more complex and information-rich than terrestrial acoustic environments. The capacities of one fish species (goldfish) to receive and make use of such sound source information have been demonstrated (sound source segregation and auditory scene analysis), and it is suggested that all vertebrate species have this capacity. A call is made to better understand underwater soundscapes, and the associated behaviors they determine in fishes.

The behavioral capacity for directional hearing and the discrimination of sound sources at different locations has been demonstrated in a number of fish species. Evidence suggests that all fishes can detect the acoustic particle motion component of sound using their ears. Few experiments, however, provide conclusive evidence that fishes can find the location of sound sources, and debate remains about whether fishes know where sound sources are in the same way as human listeners or if they have other strategies for finding sound sources. It is still uncertain how sound source localization by fishes is accomplished.

A conditioned feeding response of the mottled sculpin was used to measure level discrimination by the lateral line system as a function of source level, frequency, and distance. Level discrimination limens (LDLs) were determined with a 6‐mm‐diam, sinusoidally vibrating sphere placed in front of the trunk lateral line system at varying distances from the trunk of the fish. LDLs were relatively independent of frequency over most of the detection bandwidth, with mean values ranging from 5–6 dB from 10 to 50 Hz at a fish‐to‐source distance of 15 mm. The mean LDL at 100 Hz, near the upper end of the frequency range of the lateral line system was 8–9 dB at the same source distance. Mean LDLs at 10 and 20 dB above detection threshold were not significantly different, nor were mean LDLs at distances ranging from 7.5 to 30 mm for a 50‐Hz, 10‐dB SL source. The level‐encoding features of posterior lateral line fibers innervating the trunk were also determined from data previously collected under stimulus conditions nearly identical to those used for behavioral studies. Afferent fibers responded to level increments with increases in both spike rate and phase‐locking over most of the conditions used in behavioral experiments. Although it was impossible to eliminate spike rate as a level‐encoding mechanism at any stimulus condition, phase‐locking appeared to be a strong candidate at low source levels and vibration frequencies. These results show that level discrimination abilities by the sculpin lateral line system are within the range of capabilities of vertebrate auditory systems and that phase‐locking in peripheral fibers may be important for encoding stimulus level by the lateral line.

The aim of this study was to use plainfin midshipman fish (Porichthys notatus) as a general model to explore how fishes localize an underwater sound source in the relatively simple geometry of a monopole sound field. The robust phonotaxic responses displayed by gravid females toward a monopole sound projector (J-9) broadcasting a low-frequency (90 Hz) tone similar to the fundamental frequency of the male's advertisement call were examined. The projector's sound field was mapped at 5 cm resolution azimuth using an eight-hydrophone array. Acoustic pressure was measured with the array and acoustic particle motion was calculated from pressure gradients between hydrophones. The response pathways of the fish were analyzed from video recordings and compared to the sound field. Gravid females at initial release were directed toward the sound source, and the majority (73%) swam to the playback projector with straight to slightly curved tracks in the direction of the source and in line with local particle motion vectors. In contrast, the initial direction of the control (sound-off) group did not differ from random. This paper reports on a comparison of fish localization behavior with directional cues available in the form of local particle motion vectors.

This study is a continuation of a long-term investigation of the auditory circuit in the oyster toadfish, Opsanus tau. Input from the auditory periphery projects to the ipsilateral descending octaval nucleus (DON). Ipsilateral and contralateral DONs project to the auditory midbrain, where a previous study indicated that both frequency tuning and directional sharpening are present. To better understand the transformation of auditory information along the auditory pathway, we have examined over 400 units in the DON to characterize frequency and directional information encoded in the dorsolateral division of the nucleus. Background activity was primarily low (

This chapter, written from the perspective of four authors who have been studying fish bioacoustics for over 120 years (cumulative!), examines the major issues of the field. Each topic is put in some historical perspective, but the chapter emphasizes current thinking about acoustic communication, hearing (including bandwidth, sensitivity, detection of signals in noise, discrimination, and sound source localization), the functions of the ear (both auditory and vestibular, and including the role(s) of the otoliths and sensory hair cells) and their relationships to peripheral structures such as the swim bladder, and the interactions between the ear and the lateral line. Hearing in fishes is not only for acoustic communication and detection of sound-emitting predators and prey but can also play a major role in telling fishes about the acoustic scene at distances well beyond the range of vision. The chapter concludes with the personal views of the authors as to the major challenges and questions for future study. There are still many gaps in our knowledge of fish bioacoustics, including questions on ear function and the significance of interspecific differences in otolith size and shape and hair cell orientation, the role of the lateral line vis-a-vis the ear, the mechanisms of central processing of acoustic (and lateral line) signals, the mechanisms of sound source localization and whether fishes can determine source distance as well as direction, the evolution and functional significance of hearing specializations in taxonomically diverse fish species, and the origins of fish (and vertebrate) hearing and hearing organs.

Otoacoustic Emissions: Concepts and Origins.- Traveling Waves, Second Filters, and Physiological Vulnerability: A Short History of the Discovery of Active Processes in Hearing.- Critical Oscillators as Active Elements in Hearing.- Active Hair-Bundle Motility of the Hair Cells of Vestibular and Auditory Organs.- The Morphological Specializations and Electromotility of the Mammalian Outer Hair Cell.- Active Processes in Insect Hearing.- Otoacoustic Emissions in Amphibians, Lepidosaurs, and Archosaurs.- Otoacoustic Emissions: Basic Studies in Mammalian Models.- Mechanisms of Mammalian Otoacoustic Emission.- Cellular and Molecular Mechanisms in the Efferent Control of Cochlear Nonlinearities.- Cochlear Models Incorporating Active Processes.- Relationships Between Otoacoustic and Psychophysical Measures of Cochlear Function.- Otoacoustic Emissions as a Diagnostic Tool in a Clinical Context.- Future Directions in the Study of Active Processes and Otoacoustic Emissions.

Overview: Regeneration and Repair.- Morphological Correlates of Regeneration and Repair in the Inner Ear.- Recovery of Function in the Avian Auditory System After Ototrauma.- Functional Recovery After Hair Cell Regeneration in Birds.- Hair Cell Regeneration: Mechanisms Guiding Cellular Proliferation and Differentiation.- Protection and Repair of Inner Ear Sensory Cells.- Gene Arrays, Cell Lines, Stem Cells, and Sensory Regeneration in Mammalian Ears.

Vertebrate Hair Cells: Modern and Historic Perspectives.- The Development of Hair Cells in the Inner Ear.- The Structure and Composition of the Stereociliary Bundle of Vertebrate Hair Cells.- Mechanoelectrical Transduction in Auditory Hair Cells.- Contribution of Ionic Currents to Tuning in Auditory Hair Cells.- The Synaptic Physiology of Hair Cells.- The Piezoelectric Outer Hair Cell.- Mammalian Vestibular Hair Cells.

Our previous studies have shown that the peripheral auditory system of the toadfish encodes the direction of a sound source. Here, we compare directional responses of peripheral saccular afferents, cells in the descending octaval nucleus (DON) of the medulla, and the torus semicircularis (TS) of the midbrain. Recording locations in the brain were labeled with neurobiotin to confirm the site. To compare directional responses among cells, we calculated an index [sharpening ratio (SR)] that weights the relative strength of responses to the best direction for that cell and to the adjacent stimulus angles tested. Unsharpened saccular afferents tend to have a cosinusoidal directional response pattern (DRP) with an expected SR of 0.87. In DON, more than 60% of the cells exhibited directional sharpening (defined as SR

The octavolateralis system consists of several submodalities, including the inertial-sensitive inner ear, the pressure-sensitive ear/air cavity complex (when present), and acceleration- and velocity-sensitive components of the lateral line system (canal and superficial neuromasts, respectively). All four of these channels are responsive to many of the same stimulus sources, particularly moving or vibrating objects within a short distance from the receiver. We therefore argue that the octavolateralis system is an excellent model for the study of multisensory interactions. We focus on the possible ways in which these channels may contribute to source localization mechanisms and to the multisensory guidance of behaviors with strong directional components (e.g., predator avoidance, prey capture and mate attraction). Finally, we define four ways in which information from multiple senses might interact. These include fractionation, synergy, accessory stimulation, and complementation. Although evidence for all types of octavolateralis interactions can be found, the primary modes of interaction appear to be complementation and fractionation. For example, the inertial and pressure-sensitive submodalities of the auditory system provide complementary pieces of information about the direction (e.g., left/right) and polarity (advancing or receding) of a moving source. In contrast, the lateral line canal system subserves short-range localization tasks, whereas the auditory system may subserve longer-range detection and localization tasks.

Experiments examined differential coding of acoustic particle motion axis in the auditory midbrain of goldfish. Animals were exposed to vibratory stimuli varying in axis orientation as action potentials were recorded from single units in the central neuropil of nucleus centralis in the torus semicircularis. Response magnitudes as a function of stimulation axis were visualized in three dimensional plots called directional response profiles. These are generally comparable to directional responses observed among primary saccular afferents in having substantially vertical orientations. Distortions in shape from the peripheral patterns indicate neural information processing. A three-dimensional model was used to evaluate the hypothesis that responses in the auditory midbrain reflect the convergence of excitatory and inhibitory primary afferent-like responses. Model afferent inputs were generated and combined arithmetically. This analysis gives insight into the mechanisms of information processing that appear to occur in brainstem nuclei. The lack of diversity in best axis directions suggests that this mechanism alone cannot account for directional hearing abilities in this species. The roles that this directional representation and processing may play in directional hearing and sound source localization are not yet clear. Implications of these data on current models of fish directional hearing are discussed.

Introduction by Donata Oertel * From the cochlea to the cortex and back by Philip H. Smith and George Spirou * Cellular mechanisms for information coding in auditory brain stem nuclei by Laurence Trussell * Detection of interaural time and intensity differences for localization in the binaural pathways through the brainstem by Tom Yin * Circuitry and function of the dorsal cochlear nucleus by Eric D. Young and Kevin A. Davis * Ascending pathways through the ventral nuclei of the lateral lemniscus nuclei and their possible role in pattern recognition of natural sounds by Donata Oertel and Robert E. Wickesberg * The inferior colliculus: a hub for the central auditory system by John H. Casseday, Thane Fremouw, and Ellen Covey * Localizing sound by cortical neurons by John C. Middlebrooks, Li Xu, Shigeto Furukawa, and Brian J. Mickey * Feature detection by the auditory cortex by Israel Nelken

Introduction and overview: Genetics in auditory science and clinical audiology by Bronya JB Keats and Charles I. Berlin * Genes and mutations in hearing impairment by Karen B. Avraham and Tama Hasson * Cytogenetics and cochlear expressed sequence tags (ESTs) for identification of genes involved in hearing and deafness by Ann B. Skvorak Giersch and Cynthia C. Morton * Mapping and cloning of genes for inherited hearing impairment by Robert F. Mueller, Guy Van Camp, and Nick J. Lench * Genetic epidemiology of deafness by Walter E. Nance and Arti Pandya * Autosomal and X-linked auditory disorders by Andrew J. Griffith and Thomas B. Friedman * Hearing loss and mitochondrial DNA mutations: Clinical implications and biological lessons by Nathan Fischel-Ghodsian * Mice as models for human hereditary deafness by Karen P. Steel, Alexandra Erven, and Amy E. Kierman * Genetic couseling for deafness by Kathy S. Arnos and M. Katherine Oerlrich