Click Level Research Articles

Acoustic localization of resident killer whales for source-level estimation of echolocation clicks

To study the efficacy of resident killer whale echolocation clicks for foraging and other activities requires knowledge of the acoustic characteristics of their clicks, in particular, source levels. To compute apparent source level (ASL) from recordings of killer whales in the wild requires localization of the clicking animals, ideally including rigorous uncertainty estimates, to account for transmission losses from source to receiver. This paper considers acoustic localization of killer whales based on time-of-arrival-differences for clicks recorded at a 2 × 2 m2 array of 23 hydrophones. To quantify uncertainties, a Bayesian localization approach is formulated and two methods of solution are considered, one based on a linearized approximation and the other a nonlinear three-dimensional (3D) grid search. Simulations indicate significant linearization errors in 3D uncertainty (probability) distributions, confirming the superiority of the nonlinear localization. Results of this localization approach are used to calculate ASLs (with uncertainties) for echolocation clicks from Southern and Northern Resident killer whales determined to be directed approximately at the array (i.e., on-axis). Comparison of ASL values across varying ranges indicates a roughly logarithmic range dependence, consistent with whales adjusting their click levels based on distance to the target, approximately accounting for two-way spherical-spreading loss.

Auditory evoked potentials: Differences by sex, race, and menstrual cycle and correlations with common psychoacoustical tasks.

Auditory brainstem responses (ABRs) and auditory middle-latency responses (AMLRs) to a click stimulus were measured in about 100 subjects. Of interest were the sex differences in those auditory evoked potentials (AEPs), the correlations between the various AEP measures, and the correlations between the AEP measures and measures of otoacoustic emissions (OAEs) and behavioral performance also measured on the same subjects. Also of interest was how the menstrual cycle affected the various AEP measures. Most ABR measures and several AMLR measures exhibited sex differences, and many of the former were substantial. The sex differences tended to be larger for latency than for amplitude of the waves, and they tended to be larger for a weak click stimulus than for a strong click. The largest sex difference was for Wave-V latency (effect size ~1.2). When subjects were dichotomized into Non-Whites and Whites, the race differences in AEPs were small within sex. However, sex and race interacted so that the sex differences often were larger for the White subjects than for the Non-White subjects, particularly for the latency measures. Contrary to the literature, no AEP measures differed markedly across the menstrual cycle. Correlations between various AEP measures, and between AEP and OAE measures, were small and showed no consistent patterns across sex or race categories. Performance on seven common psychoacoustical tasks was only weakly correlated with individual AEP measures (just as was true for the OAEs also measured on these subjects). AMLR Wave Pa unexpectedly did not show the decrease in latency and increase in amplitude typically observed for AEPs when click level was varied from 40 to 70 dB nHL (normal Hearing Level). For the majority of the measures, the variability of the distribution of scores was greater for the males than for the females.

Click evoked middle ear muscle reflex: Spectral and temporal aspects

This study describes a time series-based method of middle ear muscle reflex (MEMR) detection using bilateral clicks. Although many methods can detect changes in the otoacoustic emissions evoking stimulus to monitor the MEMR, they do not discriminate between true MEMR-mediated vs artifactual changes in the stimulus. We measured MEMR in 20 young clinically normal hearing individuals using 1-s-long click trains presented at six levels (65 to 95 dB peak-to-peak sound pressure level in 6 dB steps). Changes in the stimulus levels over the 1 s period were well-approximated by two-term exponential functions. The magnitude of ear canal pressure changes due to MEMR increased monotonically as a function of click level but non-monotonically with frequency when separated into 1/3 octave wide bands between 1 and 3.2 kHz. MEMR thresholds estimated using this method were lower than that obtained from a clinical tympanometer in ∼94% of the participants. A time series-based method, along with statistical tests, may provide additional confidence in detecting the MEMR. MEMR effects were smallest at 2 kHz, between 1 and 3.2 kHz, which may provide avenues for minimizing the MEMR influence while measuring other responses (e.g., the medial olivocochlear reflex).

Effects of age on electrophysiological measures of cochlear synaptopathy in humans

Age-related cochlear synaptopathy (CS) has been shown to occur in rodents with minimal noise exposure, and has been hypothesized to play a crucial role in age-related hearing declines in humans. Because CS affects mainly low-spontaneous rate auditory nerve fibers, differential electrophysiological measures such as the ratio of the amplitude of wave I of the auditory brainstem response (ABR) at high to low click levels (WIH/WIL), and the difference between frequency following response (FFR) levels to shallow and deep amplitude modulated tones (FFRS-FFRD), have been proposed as CS markers. However, age-related audiometric threshold shifts, particularly prominent at high frequencies, may confound the interpretation of these measures in cross-sectional studies of age-related CS. To address this issue, we measured WIH/WIL and FFRS-FFRD using highpass masking (HP) noise to eliminate the contribution of high-frequency cochlear regions to the responses in a cross-sectional sample of 102 subjects (34 young, 34 middle-aged, 34 older). WIH/WIL in the presence of the HP noise did not decrease as a function of age. However, in the absence of HP noise, WIH/WIL showed credible age-related decreases even after partialing out the effects of audiometric threshold shifts. No credible age-related decreases of FFRS-FFRD were found. Overall, the results do not provide evidence of age-related CS in the low-frequency region where the responses were restricted by the HP noise, but are consistent with the presence of age-related CS in higher frequency regions.

Echolocation Clicks of Irrawaddy Dolphins (Orcaella brevirostris) During Foraging in the Bay of Brunei, Malaysia

The source parameters of Irrawaddy dolphins’ echolocation click in the Bay of Brunei were estimated. Analysis of eight parameters shows that the Irrawaddy dolphins produce broadband echolocation clicks with mean click duration of 21.1 ± 7.2 μs. The clicks had a mean peak-to-peak apparent source level (ASLpp) of 201.2 ± 7.3 dB re 1 μPa (N = 350), a mean peak frequency of 116.5 ± 15.2 kHz, a mean centroid frequency of 115.4 ± 13.9 kHz, − 3 dB bandwidth of 51.8 ± 17.7 kHz, and − 10 dB bandwidth of 100.9 ± 20.3 kHz. Foraging behaviour was characterized by the high movement of the animals in various directions with no obvious pattern and frequent deep dives. The source parameters of the Brunei Bay Irrawaddy dolphins’ clicks from the present study were compared to those of the populations in Bangladesh and Thailand. The apparent source level and frequency range of clicks for the population in Brunei Bay were wider than those of the population in Bangladesh and Thailand. The variations in the measured parameters might be due to environmental factors or behaviour related.

The Effect of Otoacoustic Emission Stimulus Level on the Strength and Detectability of the Medial Olivocochlear Reflex.

Previous work demonstrates the importance of a high signal to noise ratio (SNR) when using transient evoked otoacoustic emissions (TEOAEs) to assay the medial olivocochlear reflex (MOCR). Increasing stimulus level provides one means to increase TEOAE SNR. However, this may come at the expense of a smaller MOCR effect. It is not clear whether the gain in SNR associated with the use of higher stimulus levels outweighs the disadvantage of a potentially smaller MOCR effect. The present study investigated the strength and detectability of the MOCR when assayed using TEOAEs at different stimulus levels. The hypothesis was that although the strength of the MOCR decreases with increasing stimulus level, the occurrence of statistically significant MOCR effects increases due to an increase in TEOAE SNR. Twenty-five young adult females with normal hearing participated in the study. TEOAEs were measured in the right ear with and without broadband noise presented in the left ear. The strength of the MOCR was quantified as the percent difference in the TEOAE between the contralateral noise and quiet conditions. Statistical bootstrapping was used to detect significant MOCR effects in individual subjects across different frequency bands and stimulus levels. The relationship between a detectable MOCR (response variable) and frequency, stimulus level, TEOAE SNR, MOCR strength, and subject (predictor variables) was evaluated using generalized linear mixed-effect models. The number of statistically significant MOCR effects increased with stimulus level at all frequencies. Occurrence was highest for the 2-kHz TEOAE frequency band and lowest for the 4-kHz frequency band. The strength of the MOCR decreased with increasing click level. TEOAE SNR, MOCR strength, and stimulus level were significant predictors of a detectable MOCR: The likelihood of a detectable MOCR increased with TEOAE SNR, MOCR strength, and stimulus level. Despite a reduction in the strength of the MOCR with increasing stimulus level, the detectability of the MOCR increased. This is due, in part, to an increase in TEOAE SNR with stimulus level. For clinical implementation of TEOAE-based MOCR assays, achieving a high SNR is necessary to permit the detection of the MOCR in individual patients.

Click-Evoked Auditory Efferent Activity: Rate and Level Effects.

There currently are no standardized protocols to evaluate auditory efferent function in humans. Typical tests use broadband noise to activate the efferents, but only test the contralateral efferent pathway, risk activating the middle ear muscle reflex (MEMR), and are laborious for clinical use. In an attempt to develop a clinical test of bilateral auditory efferent function, we have designed a method that uses clicks to evoke efferent activity, obtain click-evoked otoacoustic emissions (CEOAEs), and monitor MEMR. This allows for near-simultaneous estimation of cochlear and efferent function. In the present study, we manipulated click level (60, 70, and 80dB peak-equivalent sound pressure level [peSPL]) and rate (40, 50, and 62.5Hz) to identify an optimal rate-level combination that evokes measurable efferent modulation of CEOAEs. Our findings (n = 58) demonstrate that almost all click levels and rates used caused significant inhibition of CEOAEs, with a significant interaction between level and rate effects. Predictably, bilateral activation produced greater inhibition compared to stimulating the efferents only in the ipsilateral or contralateral ear. In examining the click rate-level effects during bilateral activation in greater detail, we observed a 1-dB inhibition of CEOAE level for each 10-dB increase in click level, with rate held constant at 62.5Hz. Similarly, a 10-Hz increase in rate produced a 0.74-dB reduction in CEOAE level, with click level held constant at 80dB peSPL. The effect size (Cohen's d) was small for either monaural condition and medium for bilateral, faster-rate, and higher-level conditions. We were also able to reliably extract CEOAEs from efferent eliciting clicks. We conclude that clicks can indeed be profitably employed to simultaneously evaluate cochlear health using CEOAEs as well as their efferent modulation. Furthermore, using bilateral clicks allows the evaluation of both the crossed and uncrossed elements of the auditory efferent nervous system, while yielding larger, more discernible, inhibition of the CEOAEs relative to either ipsilateral or contralateral condition.

The effects of click and masker spectrum on the auditory brainstem response of bottlenose dolphins (Tursiops truncatus).

Two experiments were performed that investigated the effects of (1) click level and (2) continuous broadband noise on the binaural auditory brainstem response (ABR) of normal-hearing and hearing-impaired bottlenose dolphins (Tursiops truncatus). In addition to spectrally uncompensated clicks and noise, stimuli were digitally compensated to achieve "white" spectra (flat spectral density level) or "pink" spectra (spectral density level rolling off at -3 dB/octave). For experiment 1, in all spectral conditions, ABR peak latencies increased and peak amplitudes decreased with decreasing click level, but interwave intervals changed little. Latency-intensity function (LIF) slopes ranged from -3 to -11 μs/dB. The LIF slopes of ABR peaks evoked by uncompensated clicks were steeper in dolphins with hearing loss. Click level was held constant during experiment 2, and the effect of bilaterally delivered broadband masking noise on the ABR was investigated. Clicks and noise were filtered to create a pink click/noise condition and a white click/noise condition. With increasing levels of masking noise, peak latencies increased (although only P1-P4 white reached significance), peak amplitudes decreased, and interpeak intervals increased (although not significantly). These effects are compared to results reported for terrestrial mammals, and implications for auditory health assessment and biosonar function are discussed.

The echolocation beam of bottlenose dolphins (Tursiops truncatus): High-resolution measurements of horizontal beam patterns and nearfield/farfield transitions

The work of Whitlow Au and colleagues has demonstrated that dolphin biosonar forms a highly directional, forward-facing beam. In our recent studies, we have expanded upon previous work by making biosonar beam measurements using high-resolution hydrophone arrays with up to 48 hydrophones. Bottlenose dolphins were trained to echolocate on both physical targets and phantom echo generators, with clicks simultaneously recorded on all hydrophones at a sampling rate of 2 MHz. Target ranges (and simulated target ranges for phantom echoes) were varied in order to examine the resulting effects on the spatial characteristics of the acoustic field. The directivity index of the echolocation beam increased with increasing click level and center frequency, and recordings from extreme off-axis azimuths displayed a two-pulse pattern best explained by internal reflections off of the premaxillary bones. High-density hydrophone arrays placed near echolocating dolphins’ heads demonstrated that a transition from the geometric ...

Variability in Click-Evoked Potentials in Killer Whales (Orcinus orca) and Determination of a Hearing Impairment in a Rehabilitated Killer Whale

An immature female killer whale (Orcinus orca) stranded in the Wadden Sea in 2010 and was later transferred to Loro Parque, Tenerife, Spain, for rehabilitation. The killer whale, named “Morgan,” was suspected to have a hearing impairment. To test whether Morgan has a hearing deficit, auditory brainstem responses to short-duration, broadband click stimuli were recorded. The same procedure was conducted with five other killer whales at Loro Parque for comparative purposes. Stereotypical click-evoked responses were recorded in all of the killer whales except Morgan, even at the highest click level that could be projected. Reductions in the amplitude of the click-evoked response paralleled reductions in the stimulus amplitude of the clicks presented to all of the other whales. The lack of a click-evoked response in Morgan indicates that she suffers from a hearing deficit. The magnitude and frequency range over which the hearing deficit occurs cannot be specified with the techniques used here. Nevertheless, it can be concluded that Morgan’s hearing sensitivity to broadband signals is at least 20 to 30 dB worse than the hearing sensitivity of the other killer whales tested. Morgan potentially suffers from a profound hearing deficit or even a complete loss of hearing, but this cannot be determined through the electrophysiological tests used in this experiment.

Medial olivocochlear efferent reflex inhibition of human cochlear nerve responses

Inhibition of cochlear amplifier gain by the medial olivocochlear (MOC) efferent system has several putative roles: aiding listening in noise, protection against damage from acoustic overexposure, and slowing age-induced hearing loss. The human MOC reflex has been studied almost exclusively by measuring changes in otoacoustic emissions. However, to help understand how the MOC system influences what we hear, it is important to have measurements of the MOC effect on the total output of the organ of Corti, i.e., on cochlear nerve responses that couple sounds to the brain. In this work we measured the inhibition produced by the MOC reflex on the amplitude of cochlear nerve compound action potentials (CAPs) in response to moderate level (52–60 dB peSPL) clicks from five, young, normal hearing, awake, alert, human adults. MOC activity was elicited by 65 dB SPL, contralateral broadband noise (CAS). Using tympanic membrane electrodes, approximately 10 h of data collection were needed from each subject to yield reliable measurements of the MOC reflex inhibition on CAP amplitudes from one click level. The CAS produced a 16% reduction of CAP amplitude, equivalent to a 1.98 dB effective attenuation (averaged over five subjects). Based on previous reports of efferent effects as functions of level and frequency, it is possible that much larger effective attenuations would be observed at lower sound levels or with clicks of higher frequency content. For a preliminary comparison, we also measured MOC reflex inhibition of DPOAEs evoked from the same ears with f2's near 4 kHz. The resulting effective attenuations on DPOAEs were, on average, less than half the effective attenuations on CAPs.

The Source Parameters of Echolocation Clicks from Captive and Free-Ranging Yangtze Finless Porpoises (Neophocaena asiaeorientalis asiaeorientalis).

The clicks of Yangtze finless porpoises (Neophocaena asiaeorientalis asiaeorientalis) from 7 individuals in the tank of Baiji aquarium, 2 individuals in a netted pen at Shishou Tian-e-zhou Reserve and 4 free-ranging individuals at Tianxingzhou were recorded using a broadband digital recording system with four element hydrophones. The peak-to-peak apparent source level (ASL_pp) of clicks from individuals at the Baiji aquarium was 167 dB re 1 μPa with mean center frequency of 133 kHz, -3dB bandwidth of 18 kHz and -10 dB duration of 58 μs. The ASL_pp of clicks from individuals at the Shishou Tian-e-zhou Reserve was 180 dB re 1 μPa with mean center frequency of 128 kHz, -3dB bandwidth of 20 kHz and -10 dB duration of 39 μs. The ASL_pp of clicks from individuals at Tianxingzhou was 176 dB re 1 μPa with mean center frequency of 129 kHz, -3dB bandwidth of 15 kHz and -10 dB duration of 48 μs. Differences between the source parameters of clicks among the three groups of finless porpoises suggest these animals adapt to their echolocation signals depending on their surroundings.

Influence of the stimulus presentation rate on medial olivocochlear system assays.

Click evoked otoacoustic emissions (CEOAEs) are commonly used both in research and clinics to assay the medial olivocochlear system (MOC). Clicks presented at rates >50 Hz in the contralateral ear have previously been reported to evoke contralateral MOC activity. However, in typical MOC assays, clicks are presented in the ipsilateral ear in conjunction with MOC elicitor (noise) in the contralateral ear. The effect of click rates in such an arrangement is currently unknown. A forward masking paradigm was used to emulate typical MOC assays to elucidate the influence of ipsilateral click presentation rates on MOC inhibition of CEOAEs in 28 normal hearing adults. Influence of five click rates (20.83, 25, 31.25, 41.67, and 62.5 Hz) presented at 55 dB peSPL was tested. Results indicate that click rates as low as 31.25 Hz significantly enhance contralateral MOC inhibition, possibly through the activation of ipsilateral and binaural MOC neurons with potential contributions from the middle ear muscle reflex. Therefore, click rates ≤25 Hz are recommended for use in MOC assays, at least for 55 dB peSPL click level.

Mechanisms of distance-invariant recognition of target strength in the biosonar of odontocetes

Invariant target recognition by sonar requires assessment of the target strength invariably of distance despite wide variation of the echo level. Information on the echo delay and the level of the emitted pulse allows computing of the target strength. The question is: which particular mechanisms perform this computation in the biosonar of odontocetes? Investigations of the auditory evoked potentials (AEPs) during echolocation in several odontocetes have shown that comparison of the emitted pulse level, echo delay, and echo level is based on the forward masking of the echo-response by the preceding self-heard emitted click. Prolongation of the echo delay results in releasing of the echo-related AEP from masking. This release from masking compensated for the echo attenuation with distance. As a result, the echo-related AEP amplitude depended on the target strength and little depended on the target distance. Moreover, the forward-masking duration depended on the level of the emitted pulse. As a result, the echo-related AEP featured amplitude little depended on the level of the emitted pulse. The constancy of the echo-related AEP amplitude indicates the dependence of sensation level of the echo only on the target strength and the independence of both emitted click level and target distance.

Google’s Search Data and its Application in Finance

This paper examines the relationship of asset price determination via Google data. To capture this relation, I create a model and estimate several time series’ regressions. I use weekly data from 2004 to 2010 from 30 international banks. To my knowledge this is the first study which differentiates between Google’s search volume and Google’s search clicks. I show that asset prices are positively related to the rate of change in Google’s search volume, trading volume and the level of Google search clicks. Secondly, I demonstrate that the absolute level of Google’s search volume and Google’s search clicks behave differently regarding the asset price dynamics. Google’s search volume, which measures long-run searches, is negatively related while Google’s search clicks have a positive relationship to asset prices. Hence, Google’s data offer new insights on both measuring attention and pricing financial assets.

A whale better adjusts the biosonar to ordered rather than to random changes in the echo parameters

A false killer whale's (Pseudorca crassidens) sonar clicks and auditory evoked potentials (AEPs) were recorded during echolocation with simulated echoes in two series of experiments. In the first, both the echo delay and transfer factor (which is the dB-ratio of the echo sound-pressure level to emitted pulse source level) were varied randomly from trial to trial until enough data were collected (random presentation). In the second, a combination of the echo delay and transfer factor was kept constant until enough data were collected (ordered presentation). The mean click level decreased with shortening the delay and increasing the transfer factor, more at the ordered presentation rather than at the random presentation. AEPs to the self-heard emitted clicks decreased with shortening the delay and increasing the echo level equally in both series. AEPs to echoes increased with increasing the echo level, little dependent on the echo delay at random presentations but much more dependent on delay with ordered presentations. So some adjustment of the whale's biosonar was possible without prior information about the echo parameters; however, the availability of prior information about echoes provided additional whale capabilities to adjust both the transmitting and receiving parts of the biosonar.

Asset Prices and Google's Search Data

This paper investigates the novel relationship of asset price determination via Google data and trading volume. To capture this relation, we construct a model and secondly estimate panel and time series regressions. We use weekly data from 2004 to 2010 for 30 international banks. Our study is the first which differentiate between Google’s search volume and Google’s search clicks. We find that asset prices are positively related to the growth rate of Google’s search, trading volume and the level of Google search clicks. Moreover, this finding is in line with our theoretical model. Secondly, we find that the absolute level of Google’s search volume and Google’s search clicks behave differently regarding asset price dynamics. Google’s search volume, which measures long-run searches, is negatively related to asset prices and Google’s search click is positively related. Thus, Google’s search data offer a new insight into two different aspects: On the one hand it is a natural way of measuring attention to assets, and on the other hand it provides a timely and unbiased measurement of asset price dynamics especially during turbulent times. We conclude that Google’s data contain important information for the identification of asset bubbles.

Isolating mechanisms that influence measures of the precedence effect: Theoretical predictions and behavioral tests

This study tests how peripheral auditory processing and spectral dominance impact lateralization of precedence effect (PE) stimuli consisting of a pair of leading and lagging clicks. Predictions from a model whose parameters were set from established physiological results were tested with specific behavioral experiments. To generate predictions, an auditory nerve model drove a binaural, cross correlation computation whose outputs were summed across frequency using weightings derived from past physiological studies. The model predicted that lateralization (1) depends on stimulus center frequency and the inter-stimulus delay (ISD) between leading and lagging clicks for narrowband clicks and (2) changes differently with lead click level for different ISDs. Behaviorally, subjects lateralized narrowband and wideband click pairs whose stimulus parameters were chosen based on modeling results to test how peripheral processing and frequency dominance contribute to lateralization of PE stimuli. Behavioral results (including unique measures with the lead attenuated relative to the lag) suggest that peripheral interactions between leading and lagging clicks on the basilar membrane and strong weighting of cues around 750 Hz influence lateralization of paired clicks with short ISDs. When combined with auditory nerve adaptation, which emphasizes onset information, lateralization of PE click pairs with a short ISD can be well predicted.

Dolphin hearing during echolocation: evoked potential responses in an Atlantic bottlenose dolphin (Tursiops truncatus)

Auditory evoked potential (AEP) responses were recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to accept suction-cup EEG electrodes and detect targets by echolocation. AEP recording was triggered by the echolocation clicks of the animal. Three targets with target strengths of -34, -28 and -22 dB were used at a target distance of 2 to 6.5 m for each target. The results demonstrated that the AEP appeared to both outgoing echolocation clicks and echoes during echolocation, with AEP complexes consisting of alternative positive and negative waves. The echo-related AEP amplitudes were obviously lower than the outgoing click-related AEP amplitudes for all the targets at the investigated target distances. However, for targets with target strengths of -22 and -28 dB, the peak-to-peak amplitudes of the echo-related AEPs were dependent on the target distances. The echo-related AEP response amplitudes increased at further target distances, demonstrating an overcompensation of echo attenuation with target distance in the echo-perception system of the dolphin biosonar. Measurement and analysis of outgoing click intensities showed that the click levels increased with target distance (R) by a factor of approximately 10 to 17.5 logR depending on target strength. The results demonstrated that a dual-component biosonar control system formed by intensity compensation behavior in both the transmission and receiving phases of a biosonar cycle exists synchronously in the dolphin biosonar system.

Precision rodent whisker stimulator with integrated servo-locked control and displacement measurement

We developed a high speed voice coil based whisker stimulator that delivers precise deflections of a single whisker or group of whiskers in a repeatable manner. The device is miniature, quiet, and inexpensive to build. Multiple stimulators fit together for independent stimulation of four or more whiskers. The system can be used with animals under anesthesia as well as awake animals with head-restraint, and does not require trimming the whiskers. The system can deliver 1–2 mm deflections in 2 ms resulting in velocities up to 900 mm/s to attain a wide range of evoked responses. Since auditory artifacts can influence behavioral studies using whisker stimulation, we tested potential effects of auditory noise by recording somatosensory evoked potentials (SEP) with varying auditory click levels, and with/without 80 dBa background white noise. We found that auditory clicks as low as 40 dBa significantly influence the SEP. With background white noise, auditory clicks as low as 50 dBa were still detected in components of the SEP. For behavioral studies where animals must learn to respond to whisker stimulation, these sounds must be minimized. Together, the stimulator and data system can be used for psychometric vigilance tasks, mapping of the barrel cortex and other electrophysiological paradigms.