The continued importance of comparative auditory research to modern scientific discovery.

A rich history of comparative research in the auditory field has afforded a synthetic view of sound information processing by ears and brains. Some organisms have proven to be powerful models for human hearing due to fundamental similarities (e.g.

Ontogeny of the middle ear and auditory sensitivity in the Natterjack toad (Epidalea calamita).

In most anuran amphibians, acoustic communication is of prime importance for mate localization and selection. The tympanic middle ear increases auditory sensitivity and directionality and is therefore expected to be favoured by natural selection. However, especially within the family of true toads (Bufonidae) there is a tendency for species to lose parts of the middle ear apparatus and consequently have a reduced sensitivity to high-frequency sounds (above 1 kHz).

Neuroethology of sound localization in anurans.

Albert Feng pioneered the study of neuroethology of sound localization in anurans by combining behavioral experiments on phonotaxis with detailed investigations of neural processing of sound direction from the periphery to the central nervous system. The main advantage of these studies is that many species of female frogs readily perform phonotaxis towards loudspeakers emitting the species-specific advertisement call. Behavioral studies using synthetic calls can identify which parameters are important for phonotaxis and also quantify localization accuracy.

Hearing without a tympanic ear.

The ability to sense and localize sound is so advantageous for survival that it is difficult to understand the almost 100 million year gap separating the appearance of early tetrapods and the emergence of an impedance-matching tympanic middle ear - which we normally regard as a prerequisite for sensitive hearing on land - in their descendants. Recent studies of hearing in extant atympanate vertebrates have provided significant insights into the ancestral state(s) and the early evolution of the terrestrial tetrapod auditory system. These reveal a mechanism for sound pressure detection and directional hearing in 'earless' atympanate vertebrates that may be generalizable to all tetrapods, including the earliest terrestrial species.

Peripheral Auditory System Divergence Does Not Explain Species Differences in Call Preference.

Receiver sensory systems have long been cited as an important source of variation in mate preferences that could lead to signal diversification and behavioral isolation between lineages, with a general assumption that animals prefer the most conspicuous signals. The matched filter hypothesis posits that tuning of the frog peripheral auditory system matches dominant frequencies in advertisement calls used to attract mates. However, little work has characterized species with frequency modulation in their calls.

Lung mediated auditory contrast enhancement improves the Signal-to-noise ratio for communication in frogs.

Environmental noise is a major source of selection on animal sensory and communication systems. The acoustic signals of other animals represent particularly potent sources of noise for chorusing insects, frogs, and birds, which contend with a multi-species analog of the human "cocktail party problem" (i.e.

Strongly directional responses to tones and conspecific calls in the auditory nerve of the Tokay gecko, .

The configuration of lizard ears, where sound can reach both surfaces of the eardrums, produces a strongly directional ear, but the subsequent processing of sound direction by the auditory pathway is unknown. We report here on directional responses from the first stage, the auditory nerve. We used laser vibrometry to measure eardrum responses in Tokay geckos and in the same animals recorded 117 auditory nerve single fiber responses to free-field sound from radially distributed speakers.

Three auditory brainstem response (ABR) methods tested and compared in two anuran species.

Hearing sensitivity has been extensively investigated, often by measuring the auditory brainstem response (ABR). ABR measurements are relatively non-invasive, easy to reproduce, and allow the assessment of sensitivity when psychophysical data are difficult to obtain. However, the experimental methods differ greatly in respect to stimulation, which may result in different audiograms.

Lung-to-ear sound transmission does not improve directional hearing in green treefrogs ().

Amphibians are unique among extant vertebrates in having middle ear cavities that are internally coupled to each other and to the lungs. In frogs, the lung-to-ear sound transmission pathway can influence the tympanum's inherent directionality, but what role such effects might play in directional hearing remains unclear. In this study of the American green treefrog (), we tested the hypothesis that the lung-to-ear sound transmission pathway functions to improve directional hearing, particularly in the context of intraspecific sexual communication.

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

The perception of airborne infrasound (sounds below 20 Hz, inaudible to humans except at very high levels) has been documented in a handful of mammals and birds. While animals that produce vocalizations with infrasonic components (e.g.

Amphibious hearing in a diving bird, the great cormorant ().

Diving birds can spend several minutes underwater during pursuit-dive foraging. To find and capture prey, such as fish and squid, they probably need several senses in addition to vision. Cormorants, very efficient predators of fish, have unexpectedly low visual acuity underwater.

Sound localization by the internally coupled ears of lizards: From biophysics to biorobotics.

As they are generally small and only hear low frequencies, lizards have few cues for localizing sound. However, their ears show extreme directionality (up to 30 dB direction-dependent difference in eardrum vibrations) created by strong acoustical coupling of the eardrums, with almost perfect internal transmission from the contralateral ear over a broad frequency range. The activity of auditory nerve fibers reflects the eardrum directionality, so all auditory neurons are directional by default.

Modeling underwater hearing and sound localization in the frog .

Animals that are small compared to sound wavelengths face the challenge of localizing a sound source since the main cues to sound direction-interaural time differences (ITD) and interaural level differences (ILD)-both depend on size. Remarkably, the majority of terrestrial vertebrates possess internally coupled ears (ICE) with an air-filled cavity connecting the two eardrums and producing an inherently directional middle-ear system. Underwater, longer wavelengths and faster sound-speed reduce both ITD and ILD cues.

The Masked ABR (mABR): a New Measurement Method for the Auditory Brainstem Response.

The auditory brainstem response (ABR) is relatively non-invasive, and in many species, the only practical way to assess auditory sensitivity. The two main methods for measuring ABR are using either transients or tone bursts as a stimulus. The transient stimulus produces strong neural responses that contain no frequency information.

Sensitive high-frequency hearing in earless and partially eared harlequin frogs ().

Harlequin frogs, genus , communicate at high frequencies despite most species lacking a complete tympanic middle ear that facilitates high-frequency hearing in most anurans and other tetrapods. Here, we tested whether are better at sensing high-frequency acoustic sound compared with other eared and earless species in the Bufonidae family, determined whether middle ear variation within affects hearing sensitivity and tested potential hearing mechanisms in We determined that at high frequencies (2000-4000 Hz), are 10-34 dB more sensitive than other earless bufonids but are relatively insensitive to mid-range frequencies (900-1500 Hz) compared with eared bufonids. Hearing among species is fairly consistent, evidence that the partial middle ears present in a subset of species do not convey a substantial hearing advantage.

Adaptive Processes in Hearing.

Our auditory environment is constantly changing and evolving over time, requiring us to rapidly adapt to a complex dynamic sensory input. This adaptive ability of our auditory system can be observed at different levels, from individual cell responses to complex neural mechanisms and behavior, and is essential to achieve successful speech communication, correct orientation in our full environment, and eventually survival. These adaptive processes may differ in individuals with hearing loss, whose auditory system may cope via "readapting" itself over a longer time scale to the changes in sensory input induced by hearing impairment and the compensation provided by hearing devices.

Evolution of Sound Source Localization Circuits in the Nonmammalian Vertebrate Brainstem.

The earliest vertebrate ears likely subserved a gravistatic function for orientation in the aquatic environment. However, in addition to detecting acceleration created by the animal's own movements, the otolithic end organs that detect linear acceleration would have responded to particle movement created by external sources. The potential to identify and localize these external sources may have been a major selection force in the evolution of the early vertebrate ear and in the processing of sound in the central nervous system.

Earless toads sense low frequencies but miss the high notes.

Sensory losses or reductions are frequently attributed to relaxed selection. However, anuran species have lost tympanic middle ears many times, despite anurans' use of acoustic communication and the benefit of middle ears for hearing airborne sound. Here we determine whether pre-existing alternative sensory pathways enable anurans lacking tympanic middle ears (termed earless anurans) to hear airborne sound as well as eared species or to better sense vibrations in the environment.

Evidence of auditory insensitivity to vocalization frequencies in two frogs.

The emergence and maintenance of animal communication systems requires the co-evolution of signal and receiver. Frogs and toads rely heavily on acoustic communication for coordinating reproduction and typically have ears tuned to the dominant frequency of their vocalizations, allowing discrimination from background noise and heterospecific calls. However, we present here evidence that two anurans, Brachycephalus ephippium and B.

Implementation and evaluation of a Danish test battery for auditory processing disorder in children.

Objective: This study presents a Danish test battery for auditory processing disorder (APD). The tests were evaluated as to normative cut-off values (pass-fail criteria) and their test-retest reliability.

Design: The battery consists of four behavioural tests: the filtered words (FW) test, the dichotic digits (DD) test, the gap detection (GD) test and the binaural masking level difference (BMLD) test.

Animals and ICE: meaning, origin, and diversity.

ICE stands for internally coupled ears. More than half of the terrestrial vertebrates, such as frogs, lizards, and birds, as well as many insects, are equipped with ICE that utilize an air-filled cavity connecting the two eardrums. Its effect is pronounced and twofold.

Better late than never: effective air-borne hearing of toads delayed by late maturation of the tympanic middle ear structures.

Most vertebrates have evolved a tympanic middle ear that enables effective hearing of airborne sound on land. Although inner ears develop during the tadpole stages of toads, tympanic middle ear structures are not complete until months after metamorphosis, potentially limiting the sensitivity of post-metamorphic juveniles to sounds in their environment. We tested the hearing of five species of toads to determine how delayed ear development impairs airborne auditory sensitivity.

Coupled ears in lizards and crocodilians.

Lizard ears are coupled across the pharynx, and are very directional. In consequence all auditory responses should be directional, without a requirement for computation of sound source location. Crocodilian ears are connected through sinuses, and thus less tightly coupled.

Sound source localization and segregation with internally coupled ears: the treefrog model.

Acoustic signaling plays key roles in mediating many of the reproductive and social behaviors of anurans (frogs and toads). Moreover, acoustic signaling often occurs at night, in structurally complex habitats, such as densely vegetated ponds, and in dense breeding choruses characterized by high levels of background noise and acoustic clutter. Fundamental to anuran behavior is the ability of the auditory system to determine accurately the location from where sounds originate in space (sound source localization) and to assign specific sounds in the complex acoustic milieu of a chorus to their correct sources (sound source segregation).

From "ear" to there: a review of biorobotic models of auditory processing in lizards.

The peripheral auditory system of lizards has been extensively studied, because of its remarkable directionality. In this paper, we review the research that has been performed on this system using a biorobotic approach. The various robotic implementations developed to date, both wheeled and legged, of the auditory model exhibit strong phonotactic performance for two types of steering mechanisms-a simple threshold decision model and Braitenberg sensorimotor cross-couplings.