Interclick Interval Research Articles

Echolocation variability of franciscana dolphins (Pontoporia blainvillei) between estuarine and open-sea habitats, with insights into foraging patterns.

Environmental and ecological factors can trigger changes in the acoustic repertoire of cetaceans. This study documents the first use of a well-established passive acoustic monitoring device (C-POD) to analyze echolocation sounds and behavior of franciscana dolphins in different habitats: estuary [Babitonga Bay (BB)] and open sea [Itapirubá Beach (IB)]. A total of 10 924 click trains were recorded in BB and 6 093 in IB. An inter-click interval < 10 ms (so called "feeding buzzes") was used as a proxy for foraging activity. The main difference in the acoustic parameters between the two habitats was related to the frequency spectrum, with higher maximum and lower modal and minimum click frequencies in BB, and a train frequency range of 17 kHz, against 10 kHz in IB. Also, the click emission rate (clicks/s) was almost 20% higher in BB. Both studied habitats showed a high proportion of feeding buzzes (BB = 68%; IB = 58%), but with a higher probability of occurrence in BB (p < 0.001) and at night (p < 0.001) in both habitats. The C-PODs showed great potential to monitor occurrence, bioacoustics parameters, and echolocation behavior of franciscana dolphins. Longer-term temporal and spatial monitoring are necessary for elucidating several issues raised in this study.

Description of the first acoustic recording of spinner dolphins (Stenella longirostris) from the northern straits of Malacca, Malaysia (L).

Relatively little is known about spinner dolphins in Malaysian waters and the wider Southeast Asian region. This note represents the first known acoustic recording of the species sighted opportunistically in the northern Straits of Malacca. Over a brief 20 min sighting, 46 whistles were recorded and four tonal types were detected, with 54.4% being upsweep whistles. The whistle duration ranged from 36 to 977 ms and the frequency ranged from 6.6 to 23.8 kHz. Fifty-seven click trains with a mean interclick interval of 41.5 ± 19.3 ms were detected. These findings provide a baseline for future regional acoustic research on this species.

Individual differences in the ability to access spatial information in lag-clicks

It may be difficult to determine whether a dichotic lag-click points to the left or right when preceded by a diotic lead-click. Previous research suggests that this loss of spatial information is most prominent at inter-click intervals (ICIs) <10 ms. However, a study by Nilsson, Tirado, and Szychowska [(2019). J. Acoust. Soc. Am. 145, 512-524] found support for loss of spatial information in lag-clicks at much longer ICIs using a stimulus setup differing from those in previous research. The present study used a setup similar to that of the Nilsson, Tirado, and Szychowska study [(2019). J. Acoust. Soc. Am. 145, 512-524] to measure 13 listeners' ability to lateralize (left versus right) and detect (present versus absent) the lag-click in lead-lag click pairs with ICIs of 6-48 ms. The main finding was distinct individual differences in performance. Some listeners could lateralize lag-clicks all the way down to their detection threshold, whereas others had lateralization thresholds substantially higher than their detection thresholds, i.e., they could not lateralize lag-clicks that they could easily detect. Two such listeners trained for 30 days and managed to improve their lateralization thresholds to reach their detection thresholds, but only at longer ICIs (>20 ms), suggesting different mechanisms underlying lag-click lateralization at short versus long ICIs.

The biosonar of the boto: evidence of differences among species of river dolphins (Inia spp.) from the Amazon.

Echolocation clicks can reflect the anatomy of the vocalizing animal, enabling the distinction of species. River dolphins from the family Iniidae are formally represented by one species and two subspecies (Inia geoffrensis geoffrensis and I. g. humboldtiana). Additionally, two other species have been proposed (I. boliviensis and I. araguaiaensis) regarding its level of restricted distribution and morph-genetics differences. For the Committee on Taxonomy of the Society for Marine Mammalogy, the specific status of the proposed species relies on further knowledge on morphology, ecology, and genetics. Given that species-specific status is required for conservation efforts, we described and compared the echolocation clicks of Inia spp., searching for specific differences on their vocalizations. The sounds were captured with a Cetacean Research ™ C54XRS (+3/−20 dB, −185 dB re: 1V/μPa) in Guaviare River (Orinoco basin), Madeira River (Madeira basin), Xingu River (Amazon Basin), and Araguaia River (Tocantins-Araguaia basin). We found significant differences in all analyzed parameters (peak frequency, 3 dB bandwidth, 10 dB bandwidth and inter-click interval) for all species and subspecies. Differences in acoustical parameters of clicks are mainly related to the animal’s internal morphology, thus this study may potentially support with information for the species-level classification mostly of I. araguaiaensis (the Araguaian boto). Classifying the Araguaian boto separately from I. geoffrensis has important implications for the species in terms of conservation status, since it is restricted to a highly impacted river system.

Echolocation click parameters and biosonar behaviour of the dwarf sperm whale (Kogia sima).

Dwarf sperm whales (Kogia sima) are small toothed whales that produce narrow-band high-frequency (NBHF) echolocation clicks. Such NBHF clicks, subject to high levels of acoustic absorption, are usually produced by small, shallow-diving odontocetes, such as porpoises, in keeping with their short-range echolocation and fast click rates. Here, we sought to address the problem of how the little-studied and deep-diving Kogia can hunt with NBHF clicks in the deep sea. Specifically, we tested the hypotheses that Kogia produce NBHF clicks with longer inter-click intervals (ICIs), higher directionality and higher source levels (SLs) compared with other NBHF species. We did this by deploying an autonomous deep-water vertical hydrophone array in the Bahamas, where no other NBHF species are present, and by taking opportunistic recordings of a close-range Kogia sima in a South African harbour. Parameters from on-axis clicks (n=46) in the deep revealed very narrow-band clicks (root mean squared bandwidth, BWRMS, of 3±1 kHz), with SLs of up to 197 dB re. 1 µPa peak-to-peak (μPapp) at 1 m, and a half-power beamwidth of 8.8deg. Their ICIs (mode of 245 ms) were much longer than those of porpoises (<100 ms), suggesting an inspection range that is longer than detection ranges of single prey, perhaps to facilitate auditory streaming of a complex echo scene. On-axis clicks in the shallow harbour (n=870) had ICIs and SLs in keeping with source parameters of other NBHF cetaceans. Thus, in the deep, dwarf sperm whales use a directional, but short-range echolocation system with moderate SLs, suggesting a reliable mesopelagic prey habitat.

Characterization and comparison of echolocation clicks of white-beaked dolphins (Lagenorhynchus albirostris) off the Northumberland coast, UK.

Odontocetes produce ultrasonic clicks for navigation and foraging. These are commonly categorized as regular or buzz clicks based on the inter-click interval. Buzz clicks are linked to foraging behaviors and may be subdivided into slow buzz clicks for prey chase, and regular buzz clicks for prey capture. This study recorded these three click types produced by white-beaked dolphins (Lagenorhynchus albirostris) off the Northumberland coast, UK. Acoustic parameters (including duration, centroid frequency, and root-mean-squared bandwidth) were calculated and compared across the three click types. The results showed that the regular clicks had shorter durations and higher frequencies than both the buzz click types. The regular buzz clicks had longer durations, lower frequencies, and narrower bandwidths than the slow buzz clicks. Additionally, regardless of click type, about 30% of the clicks had high-frequency (200-250 kHz) secondary peaks and >90% of the clicks displayed spectral peak and notch patterns between 20 and 80 kHz. These findings are useful for future quantitative assessment of the echolocation performance of white-beaked dolphins in the wild. The patterns of spectral peaks and notches identified may facilitate for acoustic identification of this species.

Characterisation and classification method of burst pulses produced by Guiana dolphins (Sotalia guianensis)

ABSTRACT Although burst pulses are the least studied sounds produced by odontocetes, their characterisation is essential to describe the acoustic repertoire of a species. This study aimed to characterise burst pulses produced by Guiana dolphins and propose a visual method to classify these sounds into structural categories. Data collection occurred during boat surveys in south-eastern Brazil. Five acoustic parameters were extracted: number of pulses, duration, interclick interval, click repetition rate, and minimum frequency. All sounds were visually classified into four categories and used in a discriminant function analysis. In addition, a K-means cluster was used to compare the visual classification with this unsupervised grouping method and confirm the representativeness of each category. According to the discriminant analysis, most sounds were correctly classified (77%) into the four categories. Both methods grouped burst pulses in similar patterns, confirming the use of the categories. Click repetition rate was the most important parameter to separate categories. The variation in this parameter could be related to different behaviours as already suggested for other species. Thus, each category could have a different function in Guiana dolphin communication. The visual classification method is efficient and could be used for other species that produce pulsed sounds.

Echolocation inter-click interval variation among specific behaviours in free-ranging bottlenose dolphins from the coast of Uruguay

To assess whether behaviour can be inferred from echolocation trains (inter-click intervals) this study examines acoustic recordings of free-ranging bottlenose dolphins using Passive Acoustic Monitoring (PAM). Inter-click intervals from 17 groups of free-ranging bottlenose dolphins were monitored over a 36 day period in an area within the Cerro Verde (Marine Protected Area), Uruguay. Simultaneous visual observations were made from shore nearby. Results show that inter-click intervals in the echolocation trains had significant differences for three specific behaviours: feeding, socialising and travelling. The natural environment was quiet, with no disturbances in the immediate vicinity from boats, drones or other man-made noises.

Study of the relationship between sound signals and behaviors of a sperm whale during the hunting process

In order to study the relationship between the vocal law and the hunting demand of a sperm whale in the process of approaching and capturing prey, time domain characteristics such as waveform amplitude and inter click interval (ICI) and frequency domain characteristics such as centroid frequency and rms bandwidth of the sperm whale pulses are calculated firstly. Secondly, the sonar waveform parameters such as range resolution, speed resolution and Doppler tolerance are calculated using the wideband ambiguity function (WAF). Finally, variation of the above seven parameters is analyzed. The results show that the seven parameters change continuously during the hunting process. When switching from the approaching stage to the capture stage, the values of waveform amplitude, inter click interval, centroid frequency and range resolution will decrease, the values of speed resolution and rms bandwidth will increase, and the values of Doppler tolerance remain relatively unchanged. By analyzing variation of the seven parameters of the pulses and the hunting demand of the sperm whale in different hunting stages, we can better understand the relationship between the vocal law of the sperm whale and its hunting demand. We prove that the click pulses of the sperm whale transmitted during the stage of approaching and capturing prey are Doppler tolerant signals. The sperm whale constantly changes the ICI and signal forms of the transmitted signals during the approach and capture stages to adapt to different hunting demands at different hunting stages. Unlike the sonar signals of bats, the sperm whale does not change the length of the signals when switching from approach stage to capture stage, and instead it will reduce the peak pressure of the clicks significantly. The click pulses of the sperm whale in the process of approaching and capturing prey are more likely specially designed to accurately distinguish the position rather than the speed between targets, which is similar to the results of the previous research on dolphins.

Covert underwater communication based on combined encoding of diverse time-frequency characteristics of sperm whale clicks

Bio-inspired camouflage covert underwater acoustic communication has been studied widely recently. In order to improve the camouflage ability and communication rate, a bio-inspired covert underwater acoustic communication method based on combined encoding of diverse time–frequency characteristics of sperm whale clicks is proposed. The characteristics of time–frequency contour shapes and frequency-band distributions are classified and counted, and the communication codes are constructed based on the two characteristics. According to the inter-click interval characteristic of sperm whale click pulse trains, the bionic communication sequences are constructed through differential time delay difference encoding between different communication codes. Based on the proposed combined encoding method of diverse time–frequency characteristics, the constructed communication sequence contains a large number of different modulated clicks. It conforms to the characteristic that there exists a large number of and a variety of clicks in the true click pulse train, which improves the camouflage ability. The true sperm whale click pulses are inserted into the communication sequences as camouflage pulses to further improve the camouflage ability. Furthermore, the communication rate is improved due to using multiple characteristics to represent different encoding information in the proposed method. Simulations are implemented to demonstrate the effectiveness, reliability and covertness of the proposed covert communication method.

A novel approach to investigate subcortical and cortical sensitivity to temporal structure simultaneously

Hearing loss is associated with changes at the peripheral, subcortical, and cortical auditory stages. Research often focuses on these stages in isolation, but peripheral damage has cascading effects on central processing, and different stages are interconnected through extensive feedforward and feedback projections. Accordingly, assessment of the entire auditory system is needed to understand auditory pathology. Using a novel stimulus paired with electroencephalography in young, normal-hearing adults, we assess neural function at multiple stages of the auditory pathway simultaneously. We employ click trains that repeatedly accelerate then decelerate (3.5 Hz click-rate-modulation) introducing varying inter-click-intervals (4 to 40 ms). We measured the amplitude of cortical potentials, and the latencies and amplitudes of Waves III and V of the auditory brainstem response (ABR), to clicks as a function of preceding inter-click-interval. This allowed us to assess cortical processing of click-rate-modulation, as well as adaptation and neural recovery time in subcortical structures (probably cochlear nuclei and inferior colliculi). Subcortical adaptation to inter-click intervals was reflected in longer latencies. Cortical responses to the 3.5 Hz modulation included phase-locking, probably originating from auditory cortex, and sustained activity likely originating from higher-level cortices. We did not observe any correlations between subcortical and cortical responses. By recording neural responses from different stages of the auditory system simultaneously, we can study functional relationships among levels of the auditory system, which may provide a new and helpful window on hearing and hearing impairment.

Passive acoustic monitoring of the distribution patterns of Irrawaddy dolphins (Orcaella brevirostris) in the middle reaches of the Ayeyarwady River, Myanmar

AbstractThe Irrawaddy dolphins (Orcaella brevirostris) are an endangered species. Thus, up‐to‐date information on the distribution pattern of dolphins is critical for its proper management and conservation. Using a towed passive acoustic monitoring device, the distribution pattern of the Irrawaddy dolphins in the middle reaches of the Ayeyarwady River, Myanmar, was investigated during a vessel‐based survey between Mingun and Katha. This region was successively divided into segments 1–4 from upstream to downstream. Sixteen echolocation encounters, with a series of click trains separated by &lt;8 min and 26 dolphin acoustic trajectories were recorded. The mean dolphin detection rate (animals/kilometer) across the four segments progressively increased from upstream to downstream. High relative abundance was observed in segment 4 (46%) and segment 1 (23%) which was consistent with findings from historical boat‐based visual surveys. The averaged interclick intervals of each click train in segment 2 and 4 was significantly shorter than that in segment 3, indicating that the dolphins in these segments frequently use shorter‐ranged biosonar. More frequent and consistent surveys with a systematic sampling track design that incorporates other factors and covering the whole distribution range along the Ayeyarwady River and at varied water levels are needed in the future.

Presentation of Binning-Based Inter-Click Interval Data from Passive Acoustic Monitoring of Free-Ranging Harbour Porpoises (Phocoena Phocoena)

C-PODs are used for Passive Acoustic Monitoring (PAM) of harbour porpoises (Phocoena phocoena) at an offshore open sea location in the German North Sea. Diel patterns of echolocation click trains are extracted from minimum inter-click interval (minICI) data by binning. The aim of this study is to reassess and refine minICI ranges of click train data with particular consideration to the binning widths. Emphasis is also placed on choosing an appropriate visualisation of these binned data. Key ecological results include presence of higher train rates during the day with intermediate minICI values defined by the range 6-28 ms and a higher train rate with short minICI values 1.25-2.00 ms at night. This indicates an increase in porpoise feeding behaviour, or change of style, at night. Click trains with long minICI values &gt; 35 ms occur at an equal rate throughout both diel phases, suggesting a more routine behaviour, such as navigation. Results could be revealed only by judicious choice of binning widths, e.g. previously overlooked patterns within historical echolocation data. The classification methodology can be used to analyse echolocation trains from a variety of species and can be applied to any PAM data with the relevant click parameters.

Modeling the acoustic repertoire of Cuvier's beaked whale clicks.

This paper investigates the evolution of spectral properties observed in Cuvier's beaked whale (Ziphius cavirostris) click trains recorded by fixed hydrophones in the Gulf of Mexico. In the context of deep water and high-frequency sounds and observed inter-click intervals, the authors assumed that the main effect responsible for the modification of the spectral content between adjacent clicks in the same click train is the source beam pattern. The spectral structure is studied by using the Wigner-Ville time-frequency distribution and is compared with the conventional Fourier spectrogram. The results show that the observed Cuvier's beaked whale clicks are a superposition of upsweep and downsweep chirps, unlike the currently accepted upsweep only structure of beaked whale clicks in bioacoustics literature. The spectral structure variations simulated by using a flat circular piston model as a beam pattern transmission model are consistent with the evolution of spectral click properties observed in experimental data. A better understanding of the properties of observed echolocation clicks of Cuvier's beaked whales will provide useful information for click annotations and, therefore, will contribute to improving accuracy of detecting, classifying, tracking, and estimating the density of Cuvier's beaked whales.

Asymmetry and Microstructure of Temporal-Suppression Patterns in Basilar-Membrane Responses to Clicks: Relation to Tonal Suppression and Traveling-Wave Dispersion.

The cochlea's wave-based signal processing allows it to efficiently decompose a complex acoustic waveform into frequency components. Because cochlear responses are nonlinear, the waves arising from one frequency component of a complex sound can be altered by the presence of others that overlap with it in time and space (e.g., two-tone suppression). Here, we investigate the suppression of basilar-membrane (BM) velocity responses to a transient signal (a test click) by another click or tone. We show that the BM response to the click can be reduced when the stimulus is shortly preceded or followed by another (suppressor) click. More surprisingly, the data reveal two curious dependencies on the interclick interval, Δt. First, the temporal suppression curve (amount of suppression vs. Δt) manifests a pronounced and nearly periodic microstructure. Second, temporal suppression is generally strongest not when the two clicks are presented simultaneously (Δt =0), but when the suppressor click precedes the test click by a time interval corresponding to one to two periods of the best frequency (BF) at the measurement location. By systematically varying the phase of the suppressor click, we demonstrate that the suppression microstructure arises from alternating constructive and destructive interference between the BM responses to the two clicks. And by comparing temporal and tonal suppression in the same animals, we test the hypothesis that the asymmetry of the temporal-suppression curve around Δt =0 stems from cochlear dispersion and the well-known asymmetry of tonal suppression around the BF. Just as for two-tone suppression, BM responses to clicks are most suppressed by tones at frequencies just above the BF of the measurement location. On average, the frequency place of maximal suppressibility of the click response predicted from temporal-suppression data agrees with the frequency at which tonal suppression peaks, consistent with our hypothesis.

Coda repertoire and vocal clans of sperm whales in the western Atlantic Ocean

Sperm whales live in culture-based multilevel societies in which the fundamental social level is the nearly permanent social unit of females and immature individuals, and the largest level is the clan constituted by social units that share a common coda repertoire. This study describes the coda repertoire and vocal clans of sperm whales in Brazilian waters recorded between 2011 and 2016. Inter-click intervals of codas were used to quantify similarity between repertoires. The analysis showed two evident clans, in the north and the south, distinct in their repertoires. The clan of the north, termed the “5R” clan, presented a predominant production of codas containing five regularly spaced clicks, not found in the clan of the south, termed “D” clan, which repertoire was dominated by long codas with 10–13 clicks with descending inter-click intervals. Codas with five regularly spaced clicks with the same rhythm and tempo as those found in the “5R” clan of Brazil are also predominant in sperm whales recorded in the Island of Dominica, at a distance of approximately 2700 km from the north of Brazil, which corresponds to the extension of the areas of sympatric clans. The sharing of 5R type by the both distinct clans is likely the result of cultural transmission in which conformism through social learning homogenizes coda repertoire. The results of this study corroborated the hypothesis that social boundaries, here the clan level, are maintained by cultural identities.

Abnormal beta and gamma frequency neural oscillations mediate auditory sensory gating deficit in schizophrenia

BackgroundSensory gating is a process in which the brain's response to irrelevant and repetitive stimuli is inhibited. The sensory gating deficit in schizophrenia (SZ) is typically measured by the ratio or difference score of the P50 event-related potential (ERP) amplitudes in response to a paired click paradigm. While the P50 gating effect has usually been measured in relation to the peak amplitude of the S1 and S2 P50 ERPs, there is increasing evidence that inhibitory processes may be reflected by evoked or induced oscillatory activity during the inter-click interval in the beta (20–30 Hz) and gamma (30–50 Hz) frequency bands. We therefore examined the relationship between frequency specific activity in the inter-click interval with gating effects in the time and frequency domains. MethodPaired-auditory stimuli were presented to 131 participants with schizophrenia and 196 healthy controls (HC). P50 ERP amplitudes to S1 and S2as well as averaged- and single-trial beta (20–30 Hz) and gamma (30–50 Hz) frequency power during the inter-click interval were measured from the CZ electrode site. ResultsIn the time domain, P50 gating deficits were apparent in both ratio and difference scores. This effect was mainly due to smaller S1 amplitudes in the patient group. SZ patients exhibited less evoked beta and gamma power, particularly at the 0–100 ms time point, in response to S1. Early (0–100 ms) evoked beta and gamma responses were critical in determining the S1 amplitude and extent of P50 gating across the delay interval for both HC and SZ. ConclusionOur findings support a disruption in initial sensory registration in those with SZ, and do not support an active mechanism throughout the delay interval. The degree of response to S1 and early beta and gamma frequency oscillations in the delay interval provides information about the mechanisms supporting auditory sensory gating, and may provide a framework for studying the mechanisms that support sensory inhibition.

The Binaural Interaction Component of the Auditory Brainstem Response Under Precedence Effect Conditions.

The purpose of this study was to measure the binaural interaction component (BIC) derived from click-evoked auditory brainstem responses (ABRs) using stimuli configured to elicit the Precedence Effect. The hypothesis was that the contribution of binaural processing to echo suppression can be evidenced by a diminished or absent BIC associated with the echo. Ten normal-hearing young adults provided ABRs generated by sequences of click pairs. Results showed that BICs elicited by diotic clicks in isolation were obliterated when those diotic clicks were preceded by a click pair having an interaural time difference of 400 µs and where the interclick interval was 8.4 ms. The presence of the leading click pair increased the latency of the ABR generated by the lagging diotic click pair but did not decrease its amplitude. The results were interpreted as indicating a contribution of binaural processing at the level of the brainstem to echo suppression, at least for the conditions tested here.

Sounds associated with foraging and prey capture in individual fish-eating killer whales, Orcinus orca.

Foraging behavior in odontocetes is fundamentally tied to the use of sound. Resident-type killer whales use echolocation to locate and capture elusive salmonid prey. In this investigation, acoustic recording tags were suction cup-attached to endangered Southern Resident killer whales to describe their acoustic behavior during different phases of foraging that, along with detections of prey handling sounds (e.g., crunches) and observed predation events, allow confirmation of prey capture. Echolocation click trains were categorized based on the inter-click interval (ICI) according to hypothesized foraging function. Whales produced slow click trains (ICI >100 ms) at shallowest depths but over the largest change of depth, fast click trains (10 ms < ICI ≤100 ms) at intermediate depths, and buzz trains (ICI ≤10 ms) at deepest depths over the smallest depth change. These results align with hypotheses regarding biosonar use to search, pursue and capture prey. Males exhibited a higher probability of producing slow click trains, buzzes and prey handling sounds, indicating higher levels of prey searching and capture to support the energy requirement of their larger body size. These findings identify relevant acoustic indicators of subsurface foraging behaviors of killer whales, enabling investigations of human impacts on sound use and foraging.

Developing a click type “library” for Hawaiian odontocetes using machine learning methods

Though odontocete echolocation clicks are highly useful in passive acoustic monitoring, their acoustic features vary depending on behavior, orientation, and location relative to the receiver. Using unsupervised machine learning, regional sets of echolocation click types can be identified that allow for automated labeling of bioacoustic data while accounting for within-type signal variability. This methodology derives labels for clusters of click types based on parameters including spectral shape, inter-click interval, and click duration. Here we present classification results of echolocation clicks produced by Hawaiian odontocetes. Unsupervised methods were used to establish a set of click types for 10 years of recordings (200 kHz) collected from a bottom-mounted hydrophone off the coast of Kona, HI. These automated labels were compared with manual labels for species with established and recognizable clicks, which allowed for ground-truthing of some click types identified by the unsupervised machine learning method. Towed array data (500 kHz) was used to assign click types to species with visual verification, expanding the set of known species-specific click types for the region. The click types and method established here will be useful in future analyses of Hawaiian odontocete density estimation, abundance, and distribution patterns using acoustic data without requiring visual confirmation.