Thesis (M.S.)--University of North Carolina Wilmington, 2007. / Includes bibliographical references (leaves: 64-66)

Bottlenose dolphins (Tursiops truncatus) kept in zoos and dolphinarias rarely get an outlet for their echolocation abilities as their pool environment is often quite barren. Not much research has been carried out on enrichments promoting echolocation for dolphins in human care. In the present study a setup with live fish was compared to a setup with air-filled floats (providing strong sonar targets, similar to the swim bladders of large fish) and a control setup. A PCL (porpoise click logger) was used to record the echolocation click trains produced by the dolphins and aimed at the three setups. Behavioural data was also collected from video footage. Both the PCL data and all the behavioural observations indicated that the fish setup was more interesting than the float and the control setup, for the dolphins to echolocate towards. However, there were some contradictions with some parameters, where the floats and control seemed to be more interesting. This was probably due to the location of the PCL hydrophone in relation to the floats and fish, and not because the dolphins had a real bigger interest in these setups. To increase the possibility for dolphins to perform more echolocation in human care and increase their welfare, live fish can be recommended as echolocation enrichment.

To avoid masking incoming echoes with outgoing calls, bats use a low duty cycle (LDC) or high duty cycle (HDC) echolocation strategy. LDC echolocation is the most common and involves short pulses of broadband sound followed by relatively long periods of silence. In contrast, HDC echolocators emit long, narrowband sounds with short periods of silence and use Doppler shifts to detect the relative speed of prey. HDC echolocators are almost exclusively found in the families Rhinolophidae and Hipposidaridae. However, there are two known exceptions that have evolved echolocation strategies independent of the families of which they reside: Pternotus parnelli is from an LDC family but uses HDC echolocation, and conversely, Coelops frithii is from an HDC family but uses LDC echolocation. In our previous work, we used engineering software to model sound transmission from the larynx to the auditory bulla via the stylohyal-tympanic bone articulation and found that sound transmitted through the bony chain during echolocation call emission is likely loud enough for bats to hear. We also noticed differences in the morphology of the stylohyal and its placement on the tympanic bulla that might correlate with echolocation strategy. Pteronotus parnelli and C. frithii, having evolved echolocation strategies that differ from the other species in their respective families, provide the opportunity to test whether these morphologies are simply “evolutionary baggage” or driven by a selective pressure associated with function. To test this, we used µCT image data to compare the morphology of the stylohyal bones from P. parnelli, C. frithii, and other LDC and HDC echolocators. Preliminary results show that HDC echolocators have a stylohyal that sits on the medial side of the bulla, while the stylohyal in LDC echolocators sits on the lateral side of the bulla. The stylohyal in P. parnelli and C. frithi appear to have characteristics of both HDC and LDC echolocators. Further analysis will include a Multivariant Functional Shape Analysis (MFSA) to determine if these morphological differences are statistically significant and identify what variant(s) are likely driving these differences. Lastly, digital 3D models from each bat species have been built from µCT data to test for functional differences via acoustic engineering simulations. Since bone conducted sound can reach the cochlea through direct stimulation or stimulation of the eardrum, we predict that an LDC echolocator, with its stylohyal sitting on the lateral side of the bulla, will transmit sound to the cochlea more effectively through the eardrum. And given the stylohyal in HDC echolocators sits on the medial side of the bulla, we expect to see better direct stimulation of the cochlea than that of a LDC echolocator. This research uses an integrative approach to address a long-held assumption concerning the function of the stylohyal-tympanic bone articulation in laryngeally echolocating bats.

Bats are important components of biodiversity within forested ecosystems. This research addressed habitat characteristics that influence species occupancy and stable isotopes and wing morphology to assess community structure within the Sam D. Hamilton Noxubee National Wildlife Refuge. To meet these objectives, I deployed echolocation recorders, mist-nets and conducted roost checks to capture bat acoustics; fur samples were also collected to measure ratios of carbon (C13/12) and nitrogen (N15/14). Relationships between occupancy, habitat class and features were not apparent for most species. However, Lasiurus and Mytois spp. showed positive relationships with proximity to roads, Lasiurus, positive with stem density and Perimyotis subflavus, negative with basal area. Stable isotope analysis revealed some distinction of trophic niches while wing morphometrics indicated bats of similar wing shape and size show greater trophic overlap. Collectively, these results suggest that habitat management, as current within the study area, will have limited influence on local bat distributions.

The hyoid apparatus in laryngeally echolocating bats is unique as it forms a mechanical connection between the larynx and auditory bullae which has been hypothesized to transfer the outgoing echolocation call to the middle ear during call emission. Previous finite element modeling (FEM) found that hyoid-borne sound can reach the bulla at an amplitude likely heard by echolocating bats; however, that study did not model how or if the signal could reach the inner ear (or cochlea). One route that sound could take is via stimulation of the eardrum – similarly to that of air-conducted sound. We used µCT data to build models of the hyoid apparatus and middle ear from six species of bats with variable morphology. Using FEM, we ran harmonic response analyses to measure the vibroacoustic response of the tympanic membrane to hyoid-borne sound generated during echolocation and found that hyoid-borne sound in all six species stimulated the eardrum within a range likely heard by bats. Although there was variation in the efficiency between models at higher frequencies, there are no obvious morphological patterns to account for it. This suggests that hyoid morphology in laryngeal echolocators is likely driven by other associated functions and warrants further inquiry. Note: This work was published open access in the Journal of Integrative Organismal Biology (https://doi.org/10.1093/iob/obad004)

<p>Developing bat pups produce distinct vocalizations called isolation calls (I‐calls) that serve to attract the bat’s mother. How individual pups shift their vocalizations from I‐calls to downward frequency modulated (FM) sweeps during development remains unclear. By recording individual bat pups from the day of birth to twenty‐five days postnatal we observed behavioural and bioacoustic (temporal and spectral) changes in pup calls. Temporal characteristics examined were call duration and call rate; spectral characteristics were minimum frequency, maximum frequency, peak spectral frequency, total signal bandwidth, maximum frequency of the fundamental acoustic element and bandwidth of the fundamental. I‐calls were produced only until a certain point in development, after which pups change from emitting long‐duration, tonal I‐calls to downward FM signals and eventually short‐duration biosonar vocalizations. We discovered additional spectral changes in the harmonic structure of pup calls, with the number of harmonic elements decreasing with age. We also recorded pup vocalizations during prolonged separation from their mothers to determine if extended isolation alters the type, number or acoustic structure of emitted vocalizations. Rate of calling was influenced by prolonged separation; younger pups had higher calling rates and called longer than older pups. We also compared temporal and spectral characteristics of spontaneous and provoked calls. We found that provoked calls were more similar to vocalizations produced by younger pups. By documenting the vocal behaviour and acoustic structure of pups calling in different situations, this research provides groundwork for further studies on the ontogeny and development of FM vocalizations in bats and other mammals.</p> / Master of Science (MSc)

<p>Neurons throughout the auditory pathway respond selectively to the frequency and amplitude of sound. In the auditory midbrain there exists a class of neurons that are also selective to the duration of sound. These duration-tuned neurons (DTNs) provide a potential neural mechanism underlying temporal processing in the central nervous system. Temporal processing is necessary for human speech, discriminating species-specific acoustic signals as well as echolocation. This dissertation aims to explore the role and underlying mechanisms of DTNs through single-unit in vivo electrophysiological recordings in the auditory midbrain of the big brown bat. The durations that DTNs are selective to in echolocating and non-echolocating species are first compared to the durations of each species vocalizations. This comparison reveals that the durations DTNs respond best to correlates to the durations of echolocation calls in echolocating species and to species-specific communication calls in non-echolocating species. The ability of DTNs in the bat to respond to stimulus parameters thought to be important for echolocation processing, such as pairs of pulses and binaural sound localization cues, are subsequently tested. The responses of DTNs to a paired tone spike suppressing paradigm presented monaurally and binaurally are also compared to characterize the role each ear plays in recruiting inhibition known to be involved in duration tuning. The results show that DTNs are able to respond to pairs of pulses at a timescale relevant to bat echolocation, and a majority also responded selectively to binaural sound localizing cues. Nearly half (48%) of DTNs did not show spike suppression to an ipsilaterally presented suppressing tone. When ipsilaterally evoked spike suppression occurred, the effect was significantly smaller than the suppression evoked by a contralateral suppressing tone. These findings provide evidence that DTNs may play a role in echolocation in bats as DTNs are able to respond to the outgoing pulse and returning echoes and localize the echo source and that the neural mechanism underlying duration tuning is monaural in nature.</p> / Doctor of Philosophy (PhD)

Research investigating the behavioral and physiological responses of bats to echoes typically includes analysis of acoustic signals from microphones and/or microphone arrays, using time difference of arrival (TDOA) between array elements or the microphones to locate flying bats (azimuth and elevation). This has provided insight into transmission adaptations with respect to target distance, clutter, and interference. Microphones recording transmitted signals and echoes near a stationary bat provide sound pressure as a function of time but no directional information. This dissertation introduces spatial audio techniques to bat biosonar studies as a complementary method to the current TDOA based acoustical study methods. This work proposes a couple of feasible methods based on spatial audio techniques, that both track bats in flight and pinpoint the directions of echoes received by a bat. A spatial audio/soundfield microphone array is introduced to measure sounds in the sonar frequency range (20-80 kHz) of the big brown bat (Eptesicus fuscus). The custom-built ultrasonic tetrahedral soundfield microphone consists of four capacitive microphones that were calibrated to match magnitude and phase responses using a transfer function approach. Ambisonics, a signal processing technique used in three-dimensional (3D) audio applications, is used for the basic processing and reproduction of the signals measured by the soundfield microphone. Ambisonics provides syntheses and decompositions of a signal containing its directional properties, using the relationship between the spherical harmonics and the directional properties. As the first proposed method, a spatial audio decoding technique called HARPEx (High Angular Resolution Planewave Expansion) was used to build a system providing angle and elevation estimates. HARPEx can estimate the direction of arrivals (DOA) for up to two simultaneous sources since it decomposes a signal into two dominant planewaves. Experiments proved that the estimation system based on HARPEx provides accurate DOA estimates of static or moving sources. It also reconstructed a smooth flight-path of a bat by accurately estimating its direction at each snapshot of pulse measurements in time. The performance of the system was also assessed using statistical analyses of simulations. A signal model was built to generate microphone capsule responses to a virtual source emitting an LFM signal (3 ms, two harmonics: 40-22 kHz and 80-44 kHz) at an angle of 30° in the simulations. Medians and RMSEs (root-mean-square error) of 10,000 simulations for each case represent the accuracy and precision of the estimations, respectively. Results show lower d (distance between a capsule and the soundfield microphone center) or/and higher SNR (signal-to-noise ratio) are required to achieve higher estimator performance. The Cramer-Rao lower bounds (CRLB) of the estimator are also computed with various d and SNR conditions. The CRLB which is for TDOA based methods does not cover the effects of different incident angles to the capsules and signal delays between the capsules due to a non-zero d, on the estimation system. This shows the CRLB is not a proper tool to assess the estimator performance. For the second proposed method, the matched-filter technique is used instead of HARPEx to build another estimation system. The signal processing algorithm based on Ambisonics and the matched-filter approach reproduces a measured signal in various directions, and computes matched-filter responses of the reproduced signals in time-series. The matched-filter result points a target(s) by the highest filter response. This is a sonar-like estimation system that provides information of the target (range, direction, and velocity) using sonar fundamentals. Experiments using a loudspeaker (emitter) and an artificial or natural target (either stationary or moving) show the system provides accurate estimates of the target's direction and range. Simulations of imitating a situation where a bat emits a pulse and receives an echo from a target (30°) were also performed. The echo sound level is determined using the sonar equation. The system processed the virtual bat pulse and echo, and accurately estimated the direction, range, and velocity of the target. The simulation results also appear to recommend an echo level over -3 dB for accurate and precise estimations (below 15% RMSE for all parameters). This work proposes two methods to track bats in flight or/and pinpoint the directions of targets using spatial audio techniques. The suggested methods provide accurate estimates of the direction, range, or/and velocity of a bat based on its pulses or of a target based on echoes. This demonstrates these methods can be used as key tools to reconstruct bat biosonar. They would be also an independent tool or a complementary option to TDOA based methods, for bat echolocation studies. The developed methods are believed to be also useful in improving man-made sonar technology. / Doctor of Philosophy / While bats are one of the most intriguing creatures to the general population, they are also a popular subject of study in various disciplines. Their extraordinary ability to navigate and forage irrespective of clutter using echolocation has gotten attention from many scientists and engineers. Research investigating bats typically includes analysis of acoustic signals from microphones and/or microphone arrays. Using time difference of arrival (TDOA) between the array elements or the microphones is probably the most popular method to locate flying bats (azimuth and elevation). Microphone responses to transmitted signals and echoes near a bat provide sound pressure but no directional information. This dissertation proposes a complementary way to the current TDOA methods, that delivers directional information by introducing spatial audio techniques. This work shows a couple of feasible methods based on spatial audio techniques, that can both track bats in flight and pinpoint the directions of echoes received by a bat. An ultrasonic tetrahedral soundfield microphone is introduced as a measurement tool for sounds in the sonar frequency range (20-80 kHz) of the big brown bat (Eptesicus fuscus). Ambisonics, a signal processing technique used in three-dimensional (3D) audio applications, is used for the basic processing of the signals measured by the soundfield microphone. Ambisonics also reproduces a measured signal containing its directional properties. As the first method, a spatial audio decoding technique called HARPEx (High Angular Resolution Planewave Expansion) was used to build a system providing angle and elevation estimates. HARPEx can estimate the direction of arrivals (DOA) for up to two simultaneous sound sources. Experiments proved that the estimation system based on HARPEx provides accurate DOA estimates of static or moving sources. The performance of the system was also assessed using statistical analyses of simulations. Medians and RMSEs (root-mean-square error) of 10,000 simulations for each simulation case represent the accuracy and precision of the estimations, respectively. Results show shorter distance between a capsule and the soundfield microphone center, or/and higher SNR (signal-to-noise ratio) are required to achieve higher performance. For the second method, the matched-filter technique is used to build another estimation system. This is a sonar-like estimation system that provides information of the target (range, direction, and velocity) using matched-filter responses and sonar fundamentals. Experiments using a loudspeaker (emitter) and an artificial or natural target (either stationary or moving) show the system provides accurate estimates of the target's direction and range. Simulations imitating a situation where a bat emits a pulse and receives an echo from a target (30°) were also performed. The system processed the virtual bat pulse and echo, and accurately estimated the direction, range, and velocity of the target. The suggested methods provide accurate estimates of the direction, range, or/and velocity of a bat based on its pulses or of a target based on echoes. This demonstrates these methods can be used as key tools to reconstruct bat biosonar. They would be also an independent tool or a complementary option to TDOA based methods, for bat echolocation studies. The developed methods are also believed to be useful in improving sonar technology.

L’interface air-eau est un milieu où la tension de surface intervient en permanence pour minimiser l’énergie de la surface. Cette force est à l’origine des phénomènes capillaires que les organismes semi-aquatiques exploitent. L’objectif de cette thèse est d’étudier l’importance de ces phénomènes dans l’écologie des organismes semi-aquatiques en prenant les gyrins comme modèle. Les résultats montrent que le ménisque bipolaire produit par les gyrins est responsable d’auto-assemblables statiques entre individus immobiles. Les gyrins utilisent également différents types de nage pour minimiser les forces de résistance qu’ils rencontrent, notamment la force liée à la production d’onde. L’étude expérimentale de la perception d’objets immobiles suggère la perception des ménisques par les gyrins. L’écholocation grâce aux ondes de surface, supposée dans la littérature, ne peut cependant être exclue. / The surface tension at the air-water interface is responsible for capillary phenomena minimizing surface energies. Semi-aquatic organisms have evolved to exploit this capillarity. The purpose of this thesis is to study the importance of capillary phenomena in the ecology of semi-aquatic organisms using whirligig beetles as a model. Results show that the bipolar meniscus produced around whirligig beetles is responsible for static self-assembly between individuals. These insects use different types of swimming to minimize resistance forces, especially the wave drag. The experimental study of the detection of immobile objects suggests that whirligig beetles perceive the menisci. Echolocation using surface waves, assumed in literature, cannot however be excluded.

コウモリは1送信2受信器のミニマルな超音波センシングデザインからは想像できない，高度な3次元飛行を実現させている．本論文では，①繰り返し同じ障害物環境下を飛行するコウモリの未知と既知の空間に対する音響センシング行動の違いを比較した．さらに，②未知環境飛行時に見られる特徴的な空間スキャニングの行動パターンをモデル化し，自律走行車を用いてコウモリの行動の有用性を実環境センシングのふるまいから定量的に評価した． / Bats possess a highly developed biosonar system that can be regarded as the minimum sensor requirement for three-dimensional spatial sensing. The present study 1) experimentally investigated changes in the pulse direction, pulse emission timing and flight path of CF-FM bats during an obstacle avoidance flight as the bats became familiar with the space around them and 2) expressed behavioral principles observed in the bats during flight, especially in an unfamiliar space, using an algorithm and then embedded the principles into an autonomous vehicle equipped with simple ultrasound sensors. The findings of this world-leading biomimetic research offer new possibilities for artificial-intelligence navigation systems. / 博士(工学) / Doctor of Philosophy in Engineering / 同志社大学 / Doshisha University