Modulation Rate Transfer Function Research Articles

GENETICS OF DEPRESSION

Auditory thalamus (medial geniculate body [MGB]) receives ascending inhibitory GABAergic inputs from inferior colliculus (IC) and descending GABAergic projections from the thalamic reticular nucleus (TRN) with both inputs postulated to play a role in shaping temporal responses. Previous studies suggested that enhanced processing of temporally rich stimuli occurs at the level of MGB, with our recent study demonstrating enhanced GABA sensitivity in MGB compared to IC. The present study used sinusoidal amplitude-modulated (SAM) stimuli to generate modulation transfer functions (MTFs), to examine the role of GABAergic inhibition in shaping the response properties of MGB single units in anesthetized rats. Rate MTFs (rMTFs) were parsed into “bandpass (BP)”, “mixed (Mixed)”, “highpass (HP)” or “atypical” response types, with most units showing the Mixed response type. GABAA receptor blockade with iontophoretic application of the GABAA receptor (GABAAR) antagonist gabazine (GBZ) selectively altered the response properties of most MGB neurons examined. Mixed and HP units showed significant GABAAR-mediated SAM-evoked rate response changes at higher modulation frequencies (fms), which were also altered by N-methyl-d-aspartic acid (NMDA) receptor blockade (2R)-amino-5-phosphonopentanoate (AP5). BP units, and the lower arm of Mixed units responded to GABAAR blockade with increased responses to SAM stimuli at or near the rate best modulation frequency (rBMF). The ability of GABA circuits to shape responses at higher modulation frequencies is an emergent property of MGB units, not observed at lower levels of the auditory pathway and may reflect activation of MGB NMDA receptors (Rabang and Bartlett, 2011; Rabang et al., 2012). Together, GABAARs exert selective rate control over selected fms, generally without changing the units’ response type. These results showed that coding of modulated stimuli at the level of auditory thalamus is at least, in part, strongly controlled by GABA neurotransmission, in delicate balance with glutamatergic neurotransmission.

Processing of fast amplitude modulations in bat auditory cortex matches communication call-specific sound features.

Bats use a large repertoire of calls for social communication. In the bat Phyllostomus discolor, social communication calls are often characterized by sinusoidal amplitude and frequency modulations with modulation frequencies in the range of 100-130 Hz. However, peaks in mammalian auditory cortical modulation transfer functions are typically limited to modulation frequencies below 100 Hz. We investigated the coding of sinusoidally amplitude modulated sounds in auditory cortical neurons in P. discolor by constructing rate and temporal modulation transfer functions. Neuronal responses to playbacks of various communication calls were additionally recorded and compared with the neurons' responses to sinusoidally amplitude-modulated sounds. Cortical neurons in the posterior dorsal field of the auditory cortex were tuned to unusually high modulation frequencies: rate modulation transfer functions often peaked around 130 Hz (median: 87 Hz), and the median of the highest modulation frequency that evoked significant phase-locking was also 130 Hz. Both values are much higher than reported from the auditory cortex of other mammals, with more than 51% of the units preferring modulation frequencies exceeding 100 Hz. Conspicuously, the fast modulations preferred by the neurons match the fast amplitude and frequency modulations of prosocial, and mostly of aggressive, communication calls in P. discolor. We suggest that the preference for fast amplitude modulations in the P. discolor dorsal auditory cortex serves to reliably encode the fast modulations seen in their communication calls. NEW & NOTEWORTHY Neural processing of temporal sound features is crucial for the analysis of communication calls. In bats, these calls are often characterized by fast temporal envelope modulations. Because auditory cortex neurons typically encode only low modulation frequencies, it is unclear how species-specific vocalizations are cortically processed. We show that auditory cortex neurons in the bat Phyllostomus discolor encode fast temporal envelope modulations. This property improves response specificity to communication calls and thus might support species-specific communication.

Modulation rate transfer functions from four species of stranded odontocete (Stenella longirostris, Feresa attenuata, Globicephala melas, and Mesoplodon densirostris).

Odontocete marine mammals explore the environment by rapidly producing echolocation signals and receiving the corresponding echoes, which likewise return at very rapid rates. Thus, it is important that the auditory system has a high temporal resolution to effectively process and extract relevant information from click echoes. This study used auditory evoked potential methods to investigate auditory temporal resolution of individuals from four different odontocete species, including a spinner dolphin (Stenella longirostris), pygmy killer whale (Feresa attenuata), long-finned pilot whale (Globicephala melas), and Blainville's beaked whale (Mesoplodon densirostris). Each individual had previously stranded and was undergoing rehabilitation. Auditory Brainstem Responses (ABRs) were elicited via acoustic stimuli consisting of a train of broadband tone pulses presented at rates between 300 and 2000Hz. Similar to other studied species, modulation rate transfer functions (MRTFs) of the studied individuals followed the shape of a low-pass filter, with the ability to process acoustic stimuli at presentation rates up to and exceeding 1250Hz. Auditory integration times estimated from the bandwidths of the MRTFs ranged between 250 and 333 µs. The results support the hypothesis that high temporal resolution is conserved throughout the diverse range of odontocete species.

GABAergic inhibition shapes SAM responses in rat auditory thalamus

Auditory thalamus (medial geniculate body [MGB]) receives ascending inhibitory GABAergic inputs from inferior colliculus (IC) and descending GABAergic projections from the thalamic reticular nucleus (TRN) with both inputs postulated to play a role in shaping temporal responses. Previous studies suggested that enhanced processing of temporally rich stimuli occurs at the level of MGB, with our recent study demonstrating enhanced GABA sensitivity in MGB compared to IC. The present study used sinusoidal amplitude-modulated (SAM) stimuli to generate modulation transfer functions (MTFs), to examine the role of GABAergic inhibition in shaping the response properties of MGB single units in anesthetized rats. Rate MTFs (rMTFs) were parsed into “bandpass (BP)”, “mixed (Mixed)”, “highpass (HP)” or “atypical” response types, with most units showing the Mixed response type. GABAA receptor blockade with iontophoretic application of the GABAA receptor (GABAAR) antagonist gabazine (GBZ) selectively altered the response properties of most MGB neurons examined. Mixed and HP units showed significant GABAAR-mediated SAM-evoked rate response changes at higher modulation frequencies (fms), which were also altered by N-methyl-d-aspartic acid (NMDA) receptor blockade (2R)-amino-5-phosphonopentanoate (AP5). BP units, and the lower arm of Mixed units responded to GABAAR blockade with increased responses to SAM stimuli at or near the rate best modulation frequency (rBMF). The ability of GABA circuits to shape responses at higher modulation frequencies is an emergent property of MGB units, not observed at lower levels of the auditory pathway and may reflect activation of MGB NMDA receptors (Rabang and Bartlett, 2011; Rabang et al., 2012). Together, GABAARs exert selective rate control over selected fms, generally without changing the units’ response type. These results showed that coding of modulated stimuli at the level of auditory thalamus is at least, in part, strongly controlled by GABA neurotransmission, in delicate balance with glutamatergic neurotransmission.

Evolutionary adaptations for the temporal processing of natural sounds by the anuran peripheral auditory system

Sensory systems function most efficiently when processing natural stimuli, such as vocalizations, and it is thought that this reflects evolutionary adaptation. Among the best-described examples of evolutionary adaptation in the auditory system are the frequent matches between spectral tuning in both the peripheral and central auditory systems of anurans (frogs and toads) and the frequency spectra of conspecific calls. Tuning to the temporal properties of conspecific calls is less well established, and in anurans has so far been documented only in the central auditory system. Using auditory-evoked potentials, we asked whether there are species-specific or sex-specific adaptations of the auditory systems of gray treefrogs (Hyla chrysoscelis) and green treefrogs (H. cinerea) to the temporal modulations present in conspecific calls. Modulation rate transfer functions (MRTFs) constructed from auditory steady-state responses revealed that each species was more sensitive than the other to the modulation rates typical of conspecific advertisement calls. In addition, auditory brainstem responses (ABRs) to paired clicks indicated relatively better temporal resolution in green treefrogs, which could represent an adaptation to the faster modulation rates present in the calls of this species. MRTFs and recovery of ABRs to paired clicks were generally similar between the sexes, and we found no evidence that males were more sensitive than females to the temporal modulation patterns characteristic of the aggressive calls used in male-male competition. Together, our results suggest that efficient processing of the temporal properties of behaviorally relevant sounds begins at potentially very early stages of the anuran auditory system that include the periphery.

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena)

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I-VI), and 1.4 ms (II-IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal's size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.

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 100kHz 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 1,250Hz, after which response amplitudes declined until extinguished beyond 2,500Hz. 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.

Live CT imaging of sound reception anatomy and hearing measurements in the pygmy killer whale, Feresa attenuata

In June 2008, two pygmy killer whales (Feresa attenuata) were stranded alive near Boca Grande, FL, USA, and were taken into rehabilitation. We used this opportunity to learn about the peripheral anatomy of the auditory system and hearing sensitivity of these rare toothed whales. Three-dimensional (3-D) reconstructions of head structures from X-ray computed tomography (CT) images revealed mandibles that were hollow, lacked a bony lamina medial to the pan bone and contained mandibular fat bodies that extended caudally and abutted the tympanoperiotic complex. Using auditory evoked potential (AEP) procedures, the modulation rate transfer function was determined. Maximum evoked potential responses occurred at modulation frequencies of 500 and 1000 Hz. The AEP-derived audiograms were U-shaped. The lowest hearing thresholds occurred between 20 and 60 kHz, with the best hearing sensitivity at 40 kHz. The auditory brainstem response (ABR) was composed of seven waves and resembled the ABR of the bottlenose and common dolphins. By changing electrode locations, creating 3-D reconstructions of the brain from CT images and measuring the amplitude of the ABR waves, we provided evidence that the neuroanatomical sources of ABR waves I, IV and VI were the auditory nerve, inferior colliculus and the medial geniculate body, respectively. The combination of AEP testing and CT imaging provided a new synthesis of methods for studying the auditory system of cetaceans.

Encoding of amplitude modulations by auditory neurons of the locust: influence of modulation frequency, rise time, and modulation depth

Using modulation transfer functions (MTF), we investigated how sound patterns are processed within the auditory pathway of grasshoppers. Spike rates of auditory receptors and primary-like local neurons did not depend on modulation frequencies while other local and ascending neurons had lowpass, bandpass or bandstop properties. Local neurons exhibited broader dynamic ranges of their rate MTF that extended to higher modulation frequencies than those of most ascending neurons. We found no indication that a filter bank for modulation frequencies may exist in grasshoppers as has been proposed for the auditory system of mammals. The filter properties of half of the neurons changed to an allpass type with a 50% reduction of modulation depths. Contrasting to reports for mammals, the sensitivity to small modulation depths was not enhanced at higher processing stages. In ascending neurons, a focus on the range of low modulation frequencies was visible in the temporal MTFs, which describe the temporal locking of spikes to the signal envelope. To investigate the influence of stimulus rise time, we used rectangularly modulated stimuli instead of sinusoidally modulated ones. Unexpectedly, steep stimulus onsets had only small influence on the shape of MTF curves of 70% of neurons in our sample.Electronic supplementary materialThe online version of this article (doi:10.1007/s00359-010-0587-4) contains supplementary material, which is available to authorized users.

Auditory evoked potentials in a stranded Gervais’ beaked whale (Mesoplodon europaeus)

Efforts to identify the specific causal mechanisms responsible for beaked whale strandings coincident with naval exercises have been hampered by lack of data concerning the hearing abilities of beaked whales and their physiological and behavioral responses to sound. In this study, auditory capabilities of a stranded Gervais' beaked whale (Mesoplodon europaeus) were investigated by measuring auditory evoked potentials. Click-evoked potentials, auditory thresholds as a function of frequency, and the modulation rate transfer function were determined. The evoked potentials and modulation rate transfer function were similar to those measured in other echolocating odontocetes; the upper limit of functional hearing was 80-90 kHz.

Auditory Streaming of Amplitude-Modulated Sounds in the Songbird Forebrain

Streaming in auditory scene analysis refers to the perceptual grouping of multiple interleaved sounds having similar characteristics while sounds with different characteristics are segregated. In human perception, auditory streaming occurs on the basis of temporal features of sounds such as the rate of amplitude modulation. We present results from multiunit recordings in the auditory forebrain of awake European starlings (Sturnus vulgaris) on the representation of sinusoidally amplitude modulated (SAM) tones to investigate the effect of temporal envelope structure on neural stream segregation. Different types of rate modulation transfer functions in response to SAM tones were observed. The strongest responses were found for modulation frequencies (fmod) <160 Hz. The streaming stimulus consisted of sequences of alternating SAM tones with the same carrier frequency but differing in fmod (ABA-ABA-ABA-...). A signals had a modulation frequency evoking a large excitation, whereas the fmod of B signals was <or=4 octaves higher. Synchrony of B signal responses to the modulation decreased as fmod increased. Spike rate in response to B signals dropped as fmod increased. Faster signal repetition resulted in fewer spikes, suggesting the contribution of forward suppression to the response that may be due to both signals having similar spectral energy and that is not related to the temporal pattern of modulation. These two effects are additive and may provide the basis for a more separated representation of A and B signals by two populations of neurons that can be viewed as a neuronal correlate of segregated streams.

Auditory temporal resolution of a wild white-beaked dolphin (Lagenorhynchus albirostris)

Adequate temporal resolution is required across taxa to properly utilize amplitude modulated acoustic signals. Among mammals, odontocete marine mammals are considered to have relatively high temporal resolution, which is a selective advantage when processing fast traveling underwater sound. However, multiple methods used to estimate auditory temporal resolution have left comparisons among odontocetes and other mammals somewhat vague. Here we present the estimated auditory temporal resolution of an adult male white-beaked dolphin, (Lagenorhynchus albirostris), using auditory evoked potentials and click stimuli. Ours is the first of such studies performed on a wild dolphin in a capture-and-release scenario. The white-beaked dolphin followed rhythmic clicks up to a rate of approximately 1,125-1,250 Hz, after which the modulation rate transfer function (MRTF) cut-off steeply. However, 10% of the maximum response was still found at 1,450 Hz indicating high temporal resolution. The MRTF was similar in shape and bandwidth to that of other odontocetes. The estimated maximal temporal resolution of white-beaked dolphins and other odontocetes was approximately twice that of pinnipeds and manatees, and more than ten-times faster than humans and gerbils. The exceptionally high temporal resolution abilities of odontocetes are likely due primarily to echolocation capabilities that require rapid processing of acoustic cues.

Intracellular Responses of Neurons in the Mouse Inferior Colliculus to Sinusoidal Amplitude-Modulated Tones

Changes in the temporal envelope are important defining features of natural acoustic signals. Many cells in the inferior colliculus (IC) respond preferentially to certain modulation frequencies, but how they accomplish this is not yet clear. We therefore made whole cell patch-clamp recordings in the IC of anesthetized mice while presenting sinusoidal amplitude-modulated (SAM) tones. The relation between the number of evoked spikes and modulation frequency was used to construct rate modulation transfer functions (rMTFs). We observed different types of rate tuning, including band-pass (16%), band-reject (13%), high-pass (6%), and low-pass (6%) tuning. In the high-pass rMTF neurons and some of the low-pass rMTF neurons, the tuning characteristics appeared to be already present in the inputs. In both band-pass and band-reject rMTF neurons, the nonlinear relation between membrane potential and spike probability ensured preferential spiking during only a small part of the modulation period. Band-pass rMTF neurons had rapidly rising excitatory postsynaptic potentials, allowing good phase-locking to brief tones and intermediate modulation frequencies. At low modulation frequencies, adaptation of their spike threshold contributed to the onset response. In contrast, band-reject rMTF neurons responded with small excitatory or inhibitory postsynaptic potentials to brief tones. In these cells, a power law could describe the supralinear relation between average membrane potential and spike rate. Differences in timing of synaptic input and presence or absence of spike adaptation therefore define band-pass and band-reject rate tuning to SAM tones in the mouse IC.

Coevolution of auditory sensitivity and temporal resolution with acoustic signal space in three songbirds

Coevolution between senders and receivers is expected to produce a close match between signal design and sensory biology. We evaluated this hypothesis in songbirds by comparing aspects of acoustic signal space with the frequency range of auditory sensitivity and temporal resolution in tufted titmice, Baeolophus bicolor ; house sparrows, Passer domesticus ; and white-breasted nuthatches, Sitta carolinensis . Auditory measurements were made electrophysiologically from the scalp using two classes of auditory-evoked potentials: the auditory brain-stem response (ABR) and the envelope-following response (EFR). ABRs to tone-burst stimuli indicated maximum sensitivity from 2.2 to 3.2 kHz in all species, but 12–14 dB greater sensitivity in titmice than in sparrows and nuthatches at 6.4 kHz (the highest frequency tested). Modulation rate transfer functions based on EFRs to amplitude-modulated tones suggested greater temporal resolution in titmice and sparrows than in nuthatches. Conservation of the frequency range of maximum sensitivity across species resulted in a mismatch with the dominant frequency of song in sparrows. The mismatch may reflect auditory constraints coupled with selection for high-frequency song and relaxed selection for a close match between sender and receiver due to small territory size. Consistent with coevolution between senders and receivers, high-frequency sensitivity varied with the maximum frequency of species-specific vocalizations, whereas temporal resolution varied with the maximum rate of envelope periodicity. Enhanced high-frequency sensitivity of the titmouse may reflect a specialization for processing high-frequency communication signals such as alarm calls.

Temporal resolution in the hearing system and auditory evoked potentials

A popular type of investigation with auditory evoked potentials (AEP) consists of mapping the dependency of the envelope following response to the AM frequency. This results in what is called the modulation rate transfer function (MRTF). The physiological interpretation of the MRTF is not straight forward, but is often used as a measure of the ability of the auditory system to encode temporal changes. It is, however, shown here that the MRTF must depend on the waveform of the click-evoked AEP (ceAEP), which does not relate directly to temporal resolution. The theoretical relationship between the spectrum of the ceAEP and MRTF is such that the MRTF should be identical to the ceAEP if a 1/f weighting is applied to the ceAEP. Deviations from this relationship indeed reflect temporal resolution capabilities. We measured this in a harbour porpoise. Our devised stimulus was a sequence of 0.5 ms Hann weighted 130 kHz tone pips presented at an increasing rate (chirped) over a time span of 32 ms. The results reveal that the system's responsiveness declines roughly exponentially as a function of click rate with a rate constant of about -0.7 kHz and appears more rate limited than implied by traditional MRTF.

Dolphin modulation rate transfer functions resulting from exposure to frequency modulated signals

Frequency modulated (FM) signals are useful in audiometric studies where the test environment contributes to multipath interference and produces spatial variations in the sound pressure field. Rhythmic FM stimuli are useful in evoked potential audiometry because of the auditory steady state response (ASSR) that is created within the auditory system. Although some research on dolphin ASSRs resulting from rhythmic FM signals has been performed, relationships between modulation depth, modulation rate, and ASSR amplitude have not been fully explored. To address this, two dolphins were exposed to rhythmic FM signals (120 dB SPL and center frequency=40 kHz) and the resulting ASSR recorded. The modulation depth (peak‐to‐peak) and modulation rate were varied from 0.1‐10% and 200‐2500 Hz, respectively. A total of 512 epochs were recorded for each stimulus presentation and the averaged waveform was analyzed in the frequency domain. The amplitude peak corresponding to the modulation rate was used to create the modulation rate transfer function (MRTF). Both dolphins showed a MRTF peak at a 1 kHz modulation rate, while one dolphin showed a second peak at 500 Hz. The use of rhythmic FM signals in ASSR studies may be a useful for testing in less than ideal acoustic environments.

Aged-related loss of temporal processing: Altered responses to amplitude modulated tones in rat dorsal cochlear nucleus

Loss of temporal processing is characteristic of age-related loss of speech understanding observed in the elderly. Inhibitory glycinergic circuits provide input onto dorsal cochlear nucleus (DCN) projection neurons which likely serve to modulate excitatory responses to time-varying complex acoustic signals. The present study sought to test the hypothesis that age-related loss of inhibition would compromise the ability of output neurons to encode sinusoidally amplitude modulated (SAM) tones. Extracellular recordings were obtained from young and aged FBN rat DCN putative fusiform cells. Stimuli were SAM tones at three modulation depths (100, 50, and 20%) at 30 dB hearing level with the carrier frequency set to the unit's characteristic frequency. Discharge rate and synchrony were calculated to describe SAM responses. There were significant age-related changes in the shape and peak vector strength [best modulation frequency (BMF)] of temporal modulation transfer functions (tMTFs), with no significant age-related changes in rate modulation transfer functions (rMTFs) at BMF. Young neurons exhibited band-pass tMTFs for most SAM conditions while aged fusiform cells exhibited significantly more low-pass or double-peaked tMTFs. There were significant differences in tMTFs between buildup, pauser-buildup, and wide-chopper temporal response types. Young and aged wide-choppers displayed significantly lower vector strength values than the other two temporal DCN response types. Age-related decreases in the number of pauser-buildup response types and increases in wide-chopper types reported previously, could account, in part, for the observed loss of temporal coding of the aged fusiform cell. Age-related changes in SAM coding were similar to changes observed with receptor blockade of glycinergic inhibition onto fusiform cells and consistent with previously observed age-related loss of endogenous glycine levels and changes in normal adult glycine receptor function. DCN changes in SAM coding could, in part, underpin temporal processing deficits observed in the elderly.

Modulation rate transfer functions in bottlenose dolphins (Tursiops truncatus) with normal hearing and high-frequency hearing loss

Envelope following responses were measured in two bottlenose dolphins in response to sinusoidal amplitude modulated tones with carrier frequencies from 20 to 60 kHz and modulation rates from 100 to 5,000 Hz. One subject had elevated hearing thresholds at higher frequencies, with threshold differences between subjects varying from +/-4 dB at 20 and 30 kHz to +40 dB at 50 and 60 kHz. At each carrier frequency, evoked response amplitudes and phase angles were plotted with respect to modulation frequency to construct modulation rate transfer functions. Results showed that both subjects could follow the stimulus envelope components up to at least 2,000 Hz, regardless of carrier frequency. There were no substantial differences in modulation rate transfer functions for the two subjects suggesting that reductions in hearing sensitivity did not result in reduced temporal processing ability. In contrast to earlier studies, phase data showed group delays of approximately 3.5 ms across the tested frequency range, implying generation site(s) within the brainstem rather than the periphery at modulation rates from 100 to 1,600 Hz. This discrepancy is believed to be the result of undersampling of the modulation rate during previous phase measurements.

Auditory evoked potential hearing measures of a group of bottlenosed dolphins (Tursiops truncatus)

The hearing of the bottlenosed dolphins at Kolmrdens Djurpark, Sweden, was examined using a portable auditory evoked potential (AEP) system in order to examine the hearing capabilities of dolphins housed in quiet environments with an emphasis on age-related hearing variation. Audiograms were measured on two dolphins and modulation rate transfer functions (MRTF) were also measured on two animals. A complete audiogram (4–200 kHz) was obtained for the oldest dolphin, a 34 year old female named Vicky. Her audiogram showed no evidence of high-frequency hearing loss. Notwithstanding this finding, her click evoked auditory brainstem response (ABR) revealed a marked increase in interpeak latency that could be indicative of an underlying hearing defect. The audiogram obtained from for the male dolphin, a 24 year old named Pichi (measured from 4–107 kHz), demonstrated profound hearing loss at all frequencies. The measured MRTF results indicated typical odontocete following responses to rates exceeding 1000 Hz.

Hearing and Auditory Evoked Potential Methods Applied to Odontocete Cetaceans

Auditory evoked potential (AEP) procedures have been increasingly used to measure hearing processes in aquatic mammals. They have been demonstrated to be useful in measuring the audiograms of stranded animals like infant sperm whales (Physeter macrocephalus) and Risso’s dolphins (Grampus griseus). Modulation rate transfer functions (MRTF) demonstrating appropriate stimulus presentation rates are usually measured prior to recording audiograms with odontocetes. Measures comparing behavioral and AEP audiograms with the same animals have generally shown good correspondence between data gathered using the two procedures. AEPs and acoustic brainstem responses (ABRs) also have been used to measure hearing while an animal is actively echolocating. This technique of measuring the animal’s ability to hear its own outgoing signals, as well as the returning echoes, allows experimenters to develop a new understanding of the processes underlying echolocation.