Temporal weighting functions for interaural time and level differences. II. The effect of binaurally synchronous temporal jitter

Recent work has demonstrated that sensitivity to interaural time differences (ITD) carried by high-rate cochlear implant pulse trains or analogous acoustic signals can be enhanced by imposing random temporal variation on the stimulus rate [see Goupell et al. (2009). J. Acoust. Soc. Am. 126, 2511-2521]. The present study characterized the effect of such "temporal jitter" on normal-hearing listeners' weighting of ITD and interaural level differences (ILD) applied to brief trains of Gabor clicks (4 kHz center frequency) presented at nominal interclick intervals (ICI) of 1.25 and 2.5 ms. Lateral discrimination judgments were evaluated on the basis of the ITD or ILD carried by individual clicks in each train. Random perturbation of the ICI significantly reduced listeners' weighting of onset cues for both ITD and ILD discrimination compared to corresponding isochronous conditions, consistent with enhanced sensitivity to post-onset binaural cues in jittered stimuli, although the reduction of onset weighting was not statistically significant at 1.25 ms ICI. An additional analysis suggested greater weighting of ITD or ILD presented following lengthened versus shortened ICI, although weights for such "gaps" and "squeezes" were comparable to other post-onset weights. Results are discussed in terms of binaural information available in jittered versus isochronous stimuli.     

Temporal weighting of binaural cues revealed by detection of dynamic interaural differences in high-rate Gabor click trains

Listeners detected interaural differences of time (ITDs) or level (ILDs) carried by single 4000-Hz Gabor clicks (Gaussian-windowed tone bursts) and trains of 16 such clicks repeating at an interclick interval (ICI) of 2, 5, or 10 ms. In separate conditions, target interaural differences favored the right ear by a constant amount for all clicks (condition RR), attained their peak value at onset and diminished linearly to 0 at offset (condition R0), or grew linearly from 0 at onset to a peak value at offset (condition 0R). Threshold ITDs and ILDs were determined adaptively in separate experiments for each of these conditions and for single clicks. ITD thresholds were found to be lower for 16-click trains than for single clicks at 10-ms ICI, regardless of stimulus condition. At 2-ms ICI, thresholds in RR and R0 conditions were similar to single click thresholds at 2-ms ICI; thresholds in the 0R condition were significantly worse than for single clicks at 2-ms ICI, consistent with strong rate-dependent onset dominance in listeners' temporal weighting of ITD. ILD thresholds, in contrast, were predominantly unaffected by ICI, suggesting little or no onset dominance for ILD of high-rate stimuli.     

Detection of interaural differences of intensity in trains of high-frequency clicks as a function of interclick interval and number

Listeners were asked to detect interaural differences of intensity in trains of 4000-Hz clicks as the interclick interval (ICI) was varied from 10 to 1 ms and the number of clicks in a train (n) was varied from 1 to 32. As has previously been shown for differences of time [Hafter and Dye, J. Acoust. Soc. Am. 73, 644-651 (1983)], plots of log interaural threshold versus log n produced slopes that decrease with ICI. These results are explained in terms of a saturation model which argues that as the click rate increases, the evoked neural activity changes from what is essentially a tonic response toward one that is more phasic.     

Masking-level differences for trains of clicks

Masking-level differences (MLDs) were measured for trains of 2000-Hz bandpass clicks as a function of the interclick interval (ICI) and the number of clicks in the train. The magnitude of the MLD grew as the number of clicks in the train was increased from 1 to 32. While the MLDs tended to be larger at longer ICIs, the effect was mediated by changes in detectability in the homophasic conditions. For click trains consisting of 4-32 clicks, the improvement in detectability in the antiphasic conditions with increases in the number of clicks appears to be the result of integration of acoustic power, as is the case for the homophasic conditions. The absence of MLDs for short trains of high-frequency transients remains quite puzzling, since large MLDs are found with single, low-frequency transients.     

The effects of intensity on the detection of interaural differences of time in high-frequency trains of clicks

Threshold values of interaural differences of time (delta IDTs ) were measured for trains of dichotic clicks whose levels were 20, 40, or 60 dB SPL. All clicks were bandpass filtered at 4 kHz, and the number of clicks in the train (n) was 1, 2, 4, 8, 16, or 32. The interclick interval (ICI) was 5, 2, or 1 ms. Performance was compared to that of an ideal integrator of information, which produces slopes of - 0.5 when log delta IDT versus log n is plotted. The results showed that increases in level had no effect on the slopes of the log-log functions regardless of the ICI but did decrease the intercepts. Shortening the ICI caused the slopes to go from nearly - 0.5 towards 0.0. The improvement with level could be explained by either a decrease in the temporal variability of neural discharges, or by an increase in the number of samples of IDT at higher intensities brought on by increased firing rates or the activation of more auditory units. A review of the physiological literature found the most parsimonious explanation to be that the decline in threshold IDT was mediated by an increase in the number of active units, each possessing the same degree of adaptation.     

Characteristics of biosonar signals from the northern bottlenose whale, Hyperoodon ampullatus

The biosonar pulses from free-ranging northern bottlenose whales (Hyperoodon ampullatus) were recorded with a linear hydrophone array. Signals fulfilling criteria for being recorded close to the acoustic axis of the animal (a total of 10 clicks) had a frequency upsweep from 20 to 55 kHz and durations of 207 to 377 μs (measured as the time interval containing 95% of the signal energy). The source level of these signals, denoted pulses, was 175-202 dB re 1 μPa rms at 1 m. The pulses had a directionality index of at least 18 dB. Interpulse intervals ranged from 73 to 949 ms (N?=?856). Signals of higher repetition rates had interclick intervals of 5.8-13.1 ms (two sequences, made up of 59 and 410 clicks, respectively). These signals, denoted clicks, had a shorter duration (43-200 μs) and did not have the frequency upsweep characterizing the pulses of low repetition rates. The data show that the northern bottlenose whale emits signals similar to three other species of beaked whale. These signals are distinct from the three other types of biosonar signals of toothed whales. It remains unclear why the signals show this grouping, and what consequences it has on echolocation performance.     

Echolocation range of captive and free-ranging baiji (Lipotes vexillifer), finless porpoise (Neophocaena phocaenoides), and bottlenose dolphin (Tursiops truncatus).

The interclick intervals of captive dolphins are known to be longer than the two-way transit time between the dolphin and a target. In the present study, the interclick intervals of free-ranging baiji, finless porpoises, and bottlenose dolphins in the wild and in captivity were compared. The click intervals in open waters ranged up to 100-200 ms, whereas the click intervals in captivity were in the order of 4-28 ms. Echolocation of free-ranging dolphins appears to adapt to various distance in navigation or ranging, sometimes up to 140 m. Additionally, the difference of waveform characteristics of clicks between species was recognized in the frequency of maximum energy and the click duration.     

The influence of the inter-click interval on moving sound source localization for navigation systems

In this paper an analysis of moving sound source localization via headphones is presented. Also the influence of the inter-click interval on this localization is studied. The experimental sound is a short delta sound of 5 ms, generated for the horizontal frontal plane, for distances from 0.5 m to 5 m and azimuth of 32° to both left and right sides with respect of the middle line of the listener head convolutioned with individual HRTFs. The results indicate that the best accurate localization was achieved for the ICI of 150 ms. Comparing the localization accuracy in distance and azimuth is deduced that the best results have been achieved for azimuth. The results show that the listeners are able to extract accurately the distance and direction of the moving sound for larger inter-click intervals.     

Dual mechanisms in the perceptual processing of click train temporal regularity

Two experiments measured human sensitivity to temporal jitter in 25-click trains with inter-click intervals (ICIs) between 5 and 100 ms. In a naturalistic experiment using wideband clicks, jitter thresholds were a nonmonotonic function of ICI, peaking for ICIs near 40-60 ms. In a subsequent experiment, clicks were high-passed and presented against a low-frequency noise masker. Jitter threshold vs ICI functions lost the positive slope over short ICIs but retained the negative slope at long ICIs. The same behavior was seen in click rate discrimination tasks. Different processes mediate regularity analysis for click trains with ICIs above and below 40-60 ms.     

The monopulsed nature of sperm whale clicks

Traditionally, sperm whale clicks have been described as multipulsed, long duration, nondirectional signals of moderate intensity and with a spectrum peaking below 10 kHz. Such properties are counterindicative of a sonar function, and quite different from the properties of dolphin sonar clicks. Here, data are presented suggesting that the traditional view of sperm whale clicks is incomplete and derived from off-axis recordings of a highly directional source. A limited number of assumed on-axis clicks were recorded and found to be essentially monopulsed clicks, with durations of 100 micros, with a composite directionality index of 27 dB, with source levels up to 236 dB re: 1 microPa (rms), and with centroid frequencies of 15 kHz. Such clicks meet the requirements for long-range biosonar purposes. Data were obtained with a large-aperture, GPS-synchronized array in July 2000 in the Bleik Canyon off Vester?len, Norway (69 degrees 28' N, 15 degrees 40' E). A total of 14 h of sound recordings was collected from five to ten independent, simultaneously operating recording units. The sound levels measured make sperm whale clicks by far the loudest of sounds recorded from any biological source. On-axis click properties support previous work proposing the nose of sperm whales to operate as a generator of sound.     

Responses of neurons to click-pairs as simulated echoes: auditory nerve to auditory cortex

When two identical sounds are presented from different locations with a short interval between them, the perception is of a single sound source at the location of the leading sound. This "precedence effect" is an important behavioral phenomenon whose neural basis is being increasingly studied. For this report, neural responses were recorded to paired clicks with varying interstimulus intervals, from several structures of the ascending auditory system in unanesthetized animals. The structures tested were the auditory nerve, anteroventral cochlear nucleus, superior olivary complex, inferior colliculus, and primary auditory cortex. The main finding is a progressive increase in the duration of the suppressive effect of the leading sound (the conditioner) on the response to the lagging sound (the probe). The first major increase occurred between the lower brainstem and inferior colliculus, and the second between the inferior colliculus and auditory cortex. In neurons from the auditory nerve, cochlear nucleus, and superior olivary complex, 50% recovery of the response to the probe occurred, on average, for conditioner and probe intervals of approximately 2 ms. In the inferior colliculus, 50% recovery occurred at an average separation of approximately 7 ms, and in the auditory cortex at approximately 20 ms. Despite these increases in average recovery times, some neurons in every structure showed large responses to the probe within the time window for precedence (approximately 1-4 ms for clicks). This indicates that during the period of the precedence effect, some information about echoes is retained. At the other extreme, for some cortical neurons the conditioner suppressed the probe response for intervals of up to 300 ms. This is in accord with behavioral results that show dominance of the leading sound for an extended period beyond that of the precedence effect. Other transformations as information ascended included an increased variety in the shapes of the recovery functions in structures subsequent to the nerve, and neurons "tuned" to particular conditioner-probe intervals in the auditory cortex. These latter are reminiscent of neurons tuned to echo delay in bats, and may contribute to the perception of the size of the acoustic space.     

The combination of interaural time and intensity in the lateralization of high-frequency complex signals

In an effort to examine the rules by which information arising from interaural differences of time (IDT) and interaural differences of intensity (IDI) is combined, d"s were measured for trains of high-frequency clicks (4000 Hz, bandpass) possessing various combinations of IDT and IDI. The number of clicks was either 1 or 8, with the interclick interval either 2 or 10 ms. A 2-IFC task was employed in which the paired values of IDT and IDI favored one side during one interval and the other side during the other interval. Data obtained with the combined cues are compared to those obtained with IDTs or IDIs alone in order to determine the degree to which processing of the two cues is done independently. Results suggest that lateralization with such stimuli is based on the sum of the temporal and intensive differences and not on independent evaluations of their separate presences.     

Restarting the adapted binaural system

Previous experiments using trains of high-frequency filtered clicks have shown that for lateralization based on interaural difference of time or level, there is a decline in the usefulness of interaural information after the signal's onset when the clicks are presented at a high rate. This process has been referred to as "binaural adaptation." Of interest here are the conditions that produce a recovery from adaptation and allow for a resampling of the interaural information. A train of clicks with short interclick intervals is used to produce adaptation. Then, during its course, a treatment such as the insertion of a temporal gap or the addition of another "triggering" sound is tested for its ability to restart the binaural process. All of the brief triggers tested are shown to be capable of promoting recovery from adaptation. This suggests that, while the binaural system deals with the demands of high-frequency stimulation with rapid adaptation, it quickly cancels the adaptation in response to stimulus change.     

The influence of later-arriving sounds on the ability of listeners to judge the lateral position of a source

This study examined the deleterious effects of a later-arriving sound on the processing of interaural differences of time (IDTs) from a preceding sound. A correlational analysis assessed the relative weight given to IDTs of source and echo clicks for echo delays of 1-64 ms when the echo click was attenuated relative to the source click (0-36 dB). Also measured were proportion correct and the proportion of responses predicted from the weights. The IDTs of source and echo clicks were selected independently from Gaussian distributions (mu=0 s, sigma = 100/s). Listeners were instructed to indicate the laterality of the source click. Equal weight was given to the source and echo clicks for echo delays of 64 ms with no echo attenuation. For echo delays of 16-64 ms, attenuating the echo had no substantial effect on source weight or proportion correct until the echo was attenuated by 18-30 dB. At echo delays < or =4 ms, source weights and proportions correct remained high regardless of echo attenuation. The proportions of responses predicted from the weights were lower at echo delays > or =16 ms. Results were discussed in terms of backward recognition masking and binaural sluggishness and compared to measurements of echo disturbance.     

Male sperm whale acoustic behavior observed from multipaths at a single hydrophone

Sperm whales generate transient sounds (clicks) when foraging. These clicks have been described as echolocation sounds, a result of having measured the source level and the directionality of these signals and having extrapolated results from biosonar tests made on some small odontocetes. The authors propose a passive acoustic technique requiring only one hydrophone to investigate the acoustic behavior of free-ranging sperm whales. They estimate whale pitch angles from the multipath distribution of click energy. They emphasize the close bond between the sperm whale's physical and acoustic activity, leading to the hypothesis that sperm whales might, like some small odontocetes, control click level and rhythm. An echolocation model estimating the range of the sperm whale's targets from the interclick interval is computed and tested during different stages of the whale's dive. Such a hypothesis on the echolocation process would indicate that sperm whales echolocate their prey layer when initiating their dives and follow a methodic technique when foraging.     

Sperm whale clicks: directionality and source level revisited

In sperm whales (Physeter catodon L. 1758) the nose is vastly hypertrophied, accounting for about one-third of the length or weight of an adult male. Norris and Harvey [in Animal Orientation and Navigation, NASA SP-262 (1972), pp. 397-417] ascribed a sound-generating function to this organ complex. A sound generator weighing upward of 10 tons and with a cross-section of 1 m is expected to generate high-intensity, directional sounds. This prediction from the Norris and Harvey theory is not supported by published data for sperm whale clicks (source levels of 180 dB re 1 microPa and little, if any, directionality). Either the theory is not borne out or the data is not representative for the capabilities of the sound-generating mechanism. To increase the amount of relevant data, a five-hydrophone array, suspended from three platforms separated by 1 km and linked by radio, was deployed at the slope of the continental shelf off Andenes, Norway, in the summers of 1997 and 1998. With this system, source levels up to 223 dB re 1 microPa peRMS were recorded. Also, source level differences of 35 dB for the same click at different directions were seen, which are interpreted as evidence for high directionality. This implicates sonar as a possible function of the clicks. Thus, previously published properties of sperm whale clicks underestimate the capabilities of the sound generator and therefore cannot falsify the Norris and Harvey theory.     

Modeling the cochlear nucleus: a site for monaural echo suppression?

Echo suppression plays an important role in identifying and localizing auditory objects. One can distinguish between binaural and monaural echo suppression, although the former is the one commonly referred to. Based on biological findings we introduce and analyze a mathematical model for a neural implementation of monaural echo suppression in the cochlear nucleus. The model's behavior has been verified by analytical calculations as well as by numerical simulations for several types of input signal. It shows that in the perception of a pair of clicks the leading click suppresses the lagging one and that suppression is maximal for an interclick interval of 2-3 ms. Similarly, ongoing stimuli will be affected by the suppression mechanism primarily a couple of milliseconds after onset, resulting in a reduced perception of a sound shortly after its start. Both effects match experimental data.     

Modeling of the dolphin’s clicking sound source: The influence of the critical parameters

A physical and a mathematical models of the dolphin’s source of echolocation clicks have been recently proposed. The physical model includes a bottle of pressurized air connected to the atmosphere with an underwater rubber tube. A compressing rubber ring is placed on the underwater portion of the tube. The ring blocks the air jet passing through the tube from the bottle. This ring can be brought into self-oscillation by the air jet. In the simplest case, the ring displacement follows a repeated triangular waveform. Because the acoustic pressure gradient is proportional to the second time derivative of the displacement, clicks arise at the bends of the displacement waveform. The mathematical model describes the dipole oscillations of a sphere “frozen” in the ring and calculates the waveform and the sound pressure of the generated clicks. The critical parameters of the mathematical model are the radius of the sphere and the peak value and duration of the triangular displacement curve. This model allows one to solve both the forward (deriving the properties of acoustic clicks from the known source parameters) and the inverse (calculating the source parameters from the acoustic data) problems. Data from click records of Odontocetes were used to derive both the displacement waveforms and the size of the “frozen sphere” or a structure functionally similar to it. The mathematical model predicts a maximum source level of up to 235 dB re 1 μPa at 1-m range when using a 5-cm radius of the “frozen” sphere and a 4-mm maximal displacement. The predicted sound pressure level is similar to that of the clicks produced by Odontocetest.     

Psychophysical evidence against the autocorrelation theory of auditory temporal processing.

Nowadays, it is widely believed that the temporal structure of the auditory nerve fibers' response to sound stimuli plays an important role in auditory perception. An influential hypothesis is that information is extracted from this temporal structure by neural operations akin to an autocorrelation algorithm. The goal of the present work was to test this hypothesis. The stimuli consisted of sequences of unipolar clicks that were high-pass filtered and mixed with low-pass noise so as to exclude spectral cues. In experiment 1, "interfering" clicks were inserted in an otherwise periodic (isochronous) click sequence. Each click belonging to the periodic sequence was followed, after a random portion of the period, by one interfering click. This disrupted the detection of temporal regularity, even when the interfering clicks were 5 dB less intense than the periodic clicks. Experiments 2-4 used click sequences that showed a single peak in their autocorrelation functions. For some sequences, this peak originated from "first-order" temporal regularities, that is from the temporal relations between consecutive clicks. For other sequences, the peak originated instead from "second-order" regularities, relative to nonconsecutive clicks. The detection of second-order regularities appeared to be much more difficult than the detection of comparable first-order regularities. Overall, these results do not tally with the current autocorrelation models of temporal processing. They suggest that the extraction of temporal information from a group of closely spaced spectral components makes no use of time intervals between nonconsecutive peaks of the amplitude envelope.