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Echolocation signals of the free-ranging Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientialis).

This paper describes the high-frequency echolocation signals from free-ranging Yangtze finless porpoise in the Tian-e-zhou Baiji National Natural Reserve in Hubei Province, China. Signal analysis showed that the Yangtze finless porpoise clicks are typical high-frequency narrow-band (relative width of the frequency spectrum Q = 6.6 +/- 1.56, N = 548) ultrasonic pulses. The peak frequencies of the typical clicks range from 87 to 145 kHz with an average of 125 +/- 6.92 kHz. The durations range from 30 to 122 micros with an average of 68 +/- 14.12, as. The characteristics of the signals are similar to those of other members of the Phocoenidae as well as the distantly related delphinids, Cephalorhynchus spp. Comparison of these signals to those of the baiji (Lipotes vexillifer), who occupies habitat similar to that of the Yangtze finless porpoise, showed that the peak frequencies of clicks produced by the Yangtze finless porpoise are remarkably higher than those produced by the baiji. Difference in peak frequency between the two species is probably linked to the different size of preferred prey fish. Clear double-pulse and multi-pulse reverberation structures of clicks are noticed, and there is no indication of any low-frequency (< 70 kHz) components during the recording period.

Animals↗

Echolocation clicks of free-ranging Cuvier's beaked whales (Ziphius cavirostris).

Strandings of beaked whales of the genera Ziphius and Mesoplodon have been reported to occur in conjunction with naval sonar use. Detection of the sounds from these elusive whales could reduce the risk of exposure, but descriptions of their vocalizations are at best incomplete. This paper reports quantitative characteristics of clicks from deep-diving Cuvier's beaked whales (Ziphius cavirostris) using a unique data set. Two whales in the Ligurian Sea were simultaneously tagged with sound and orientation recording tags, and the dive tracks were reconstructed allowing for derivation of the range and relative aspect between the clicking whales. At depth, the whales produced trains of regular echolocation clicks with mean interclick intervals of 0.43 s (+/- 0.09) and 0.40 s (+/- 0.07). The clicks are frequency modulated pulses with durations of approximately 200 micros and center frequencies around 42 kHz, -10 dB bandwidths of 22 kHz, and Q(3 dB) of 4. The sound beam is narrow with an estimated directionality index of more than 25 dB, source levels up to 214 dB(pp) re: 1 microPa at 1 m, and energy flux density of 164 dB re: 1 microPa2 s. As the spectral and temporal properties are different from those of nonziphiid odontocetes the potential for passive detection is enhanced.

Animals↗

Target representation of naturalistic echolocation sequences in single unit responses from the inferior colliculus of big brown bats.

Echolocating big brown bats (Eptesicus fuscus) emit trains of frequency-modulated (FM) biosonar signals whose duration, repetition rate, and sweep structure change systematically during interception of prey. When stimulated with a 2.5-s sequence of 54 FM pulse-echo pairs that mimic sounds received during search, approach, and terminal stages of pursuit, single neurons (N = 116) in the bat's inferior colliculus (IC) register the occurrence of a pulse or echo with an average of < 1 spike/sound. Individual IC neurons typically respond to only a segment of the search or approach stage of pursuit, with fewer neurons persisting to respond in the terminal stage. Composite peristimulus-time-histogram plots of responses assembled across the whole recorded population of IC neurons depict the delay of echoes and, hence, the existence and distance of the simulated biosonar target, entirely as on-response latencies distributed across time. Correlated changes in pulse duration, repetition rate, and pulse or echo amplitude do modulate the strength of responses (probability of the single spike actually occurring for each sound), but registration of the target itself remains confined exclusively to the latencies of single spikes across cells. Modeling of echo processing in FM biosonar should emphasize spike-time algorithms to explain the content of biosonar images.

Acoustic Stimulation↗

Source-to-sensation level ratio of transmitted biosonar pulses in an echolocating false killer whale.

Transmitted biosonar pulses, and the brain auditory evoked potentials (AEPs) associated with those pulses, were synchronously recorded in a false killer whale Pseudorca crassidens trained to accept suction-cup EEG electrodes and to detect targets by echolocation. AEP amplitude was investigated as a function of the transmitted biosonar pulse source level. For that, a few thousand of the individual AEP records were sorted according to the spontaneously varied amplitude of synchronously recorded biosonar pulses. In each of the sorting bins (in 5-dB steps) AEP records were averaged to extract AEP from noise; AEP amplitude was plotted as a function of the biosonar pulse source level. For comparison, AEPs were recorded to external (in free field) sound pulses of a waveform and spectrum similar to those of the biosonar pulses; amplitude of these AEPs was plotted as a function of sound pressure level. A comparison of these two functions has shown that, depending on the presence or absence of a target, the sensitivity of the whale's hearing to its own transmitted biosonar pulses was 30 to 45 dB lower than might be expected in a free acoustic field.

Acoustics↗

Sonar gain control in echolocating finless porpoises (Neophocaena phocaenoides) in an open water.

Source levels of echolocating free-ranging Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis) were calculated using a range estimated by measuring the time delays of the signals via the surface and bottom reflection paths to the hydrophone, relative to the direct signal. Peak-to-peak source levels for finless porpoise were from 163.7 to 185.6 dB re: 1 microPa. The source levels are highly range, dependent and varied approximately as a function of the one-way transmission loss for signals traveling from the animals to the hydrophone.

Acoustics↗

Echolocation transmitting beam of the Atlantic bottlenose dolphin.

The transmitting beam patterns of echolocation signals emitted by an Atlantic bottlenose dolphin Tursiops truncatus were measured in the vertical and horizontal planes with an array of seven hydrophones. Particular emphasis was placed on accurately verifying the animal's position on a bite-plate/tail-rest stationing device using underwater video monitoring equipment. The major axis of the vertical beam was directed at an angle of 5 degrees above the plane defined by the animal's lips. This angle was 15 degrees lower than previously measured. The vertical beam measurements indicate that the major axis of the transmitting beam is aligned with the major axis of the receiving beam. The horizontal beam was directed forward. The directivity index of 26.5 dB calculated from the beam pattern measured in both planes agreed well with previous calculation of 25.4 dB.

Acoustics↗

Propagation of beluga echolocation signals.

The propagation characteristics of high-frequency echolocation signals (peak energies above 100 kHz) of the beluga (Delphinapterus leucas) were measured while the animal performed a target detection task. The whale was trained to station on a bite plate so that its transmission beam could be measured in the vertical and horizontal planes using hydrophone arrays. The transitional region between the acoustic near- and farfields was also located using an array of hydrophones that extended directly in front of the animal in the horizontal plane. Three distinct modes of signals were observed. Mode 1 signals had click intervals greater than the time required for the signals to travel to the target and back (two-way transit time). Mode 2 signals had click intervals shorter than the two-way transit time, and mode 3 signals had high repetition rates with an average click interval of 1.7 ms, approximately 2% of the two-way transit time. The average click intervals for the modes 1 and 2 signals were 193 and 44 ms, respectively. The vertical and horizontal beam patterns of the mode 1 signals had similar 3-dB beamwidths of approximately 6.5 degrees. The major axis of the vertical beam was directed approximately 5 degrees above the plane defined by the animal's teeth. The near- to farfield transition region was approximately 0.64-0.75 m from the tip of the animal's mouth.

Animals↗

The sound emission pattern and the acoustical role of the noseleaf in the echolocating bat, Carollia perspicillata.

Carollia perspicillata (Phyllostomidae) is a frugivorous bat that emits low-intensity, broadband, frequency-modulated echolocation pulses through nostrils surrounded by a noseleaf. The emission pattern of this bat is of interest because the ratio between the nostril spacing and the emitted wavelength varies during the pulse, causing complex interference patterns in the horizontal dimension. Sound pressures around the bat were measured using a movable microphone and were referenced to those at a stationary microphone positioned directly in front of the animal. Interference between the nostrils was confirmed by blocking one nostril, which eliminated sidelobes and minima in the emission pattern, and by comparison of real emission patterns with simple computer models. The positions of minima in the patterns indicate effective nostril spacings of over a half-wavelength. Displacement of the dorsal lancet of the noseleaf demonstrated that this structure directs sound in the vertical dimension.

Acoustics↗

Clutter interference along the target range axis in the echolocating bat, Eptesicus fuscus.

The sensitivity of the echolocating bat, Eptesicus fuscus, for detection of a sonar target is impaired by the presence of additional targets located at similar distances. At a range of 54 cm, sensitivity to one target declines if the range separation to other targets is smaller than 8-9 cm. This loss of sensitivity is an example of clutter interference along the range axis, created by simultaneous masking of one set of echoes by another. Echoes that fall within an experimentally defined critical range band may sum together to contribute collectively to detection in that band. Echoes falling into separate bands may be detected independently. Acoustic glints within a band appear to be grouped together to be perceived as a single range-extended target of complex structure. Range bands may thus define what a "target" is by specifying within-target and between-target differences in range. The width of critical range bands appears to depend upon target range, with wider bands at greater ranges.

Animals↗

Clutter interference and the integration time of echoes in the echolocating bat, Eptesicus fuscus.

The ability of the echolocating bat, Eptesicus fuscus, to detect a sonar target is affected by the presence of other targets along the same axis at slightly different ranges. If echoes from one target arrive at about the same delay as echoes from another target, clutter interference occurs and one set of echoes masks the other. Although the bat's sonar emissions and the echoes themselves are 2 to 5 ms long, echoes (of approximately equal sensation levels--around 15 dB SL) only interfere with each other if they arrive within 200 to 400 microseconds of the same arrival time. This figure is an estimate of the integration time of the bat's sonar receiver for echoes. The fine structure of the clutter-interference data reflects the reinforcement and cancellation of echoes according to their time separation. When clutter interference first occurs, the waveforms of test and cluttering echoes already overlap for much of their duration. The masking effect underlying clutter interference appears specifically due to overlap, not between raw echo waveforms, but between the patterns of mechanical excitation created when echoes pass through bandpass filters equivalent to auditory-nerve tuning curves. While the time scale of clutter interference is substantially shorter than the duration of echo waveforms, it still is much longer than the eventual width of a target's range-axis image expressed in terms of echo delay.

Animals↗

The acoustic basis for target discrimination by FM echolocating bats.

Past experiments show that echolocating bats of the species Myotis lucifugus and Eptesicus fuscus can discriminate among airborne sonar targets presented in the context of pursuit maneuvers for the interception of prey. These bats distinguish between edible mealworms and inedible spheres of various sizes. Myotis can distinguish between disks and mealworms similar enough in size that the bat's performance requires the ability to perceive the acoustic equivalent of target shape. Previously observed small differences in the spectrum of echoes from mealworms and disks appear insufficient to distinguish these targets at the performance levels achieved by bats. We measured the acoustic properties of the targets by broadcasting ultrasonic impulses at mealworms, spheres, and disks and recording their echoes, displaying the results in terms of impulse echo waveforms and the frequency response of targets derived from the target transfer function. The echoes from disks and mealworms at various orientations convey the range-axis profile of the target (number and spacing of reflecting points or glints distributed at different ranges) in terms of the impulse structure of their waveforms and in terms of the locations and spacing of notches or nulls in their spectra. For targets that bats can discriminate and that reflect echoes which do not clearly differ in overall amplitude, the targets appear distinguishable from the acoustic representation of their range profile, which is a feature of targets that bats can perceive with great acuity.

Animals↗

Intrinsic echolocation capability of Hector's dolphin, Cephalorhynchus hectori.

A sonar system's echolocation capabilities can be inferred from the ambiguity distribution (defined here in terms of the conventional signal response function) of each of its transmitted signals. Several records of sounds emitted by Hector's dolphin are analyzed. The computed ambiguity distributions indicate that the sonar clicks of Hector's dolphins should be capable of resolving the ranges of targets as close together as 2 cm apart, but that target velocities cannot be resolved to any useful degree from a single echo.

Acoustics↗

Sonar gain control and echo detection thresholds in the echolocating bat, Eptesicus fuscus.

The echolocating bat, Eptesicus fuscus, detects sonar echoes with a sensitivity that changes according to the time elapsed between broadcasting of each sonar signal and reception of echoes. When tested in an electronic target simulator on a two-choice echo-detection task, the bat's threshold improved by 11.5 dB as echo delay changed from 2.3 to 4.6 ms (target ranges of 40 and 80 cm). Earlier experiments measured the change in detection threshold for delays from 1 to 6.4 ms (target ranges from about 17 to 110 cm) and obtained about 11 dB of improvement per doubling of delay. The new experiments used electronic delay lines to simulate echo delay, thus avoiding movement of loudspeakers to different distances and the possible creation of delay-dependent backward masking between stimulus echoes and cluttering echoes from the loudspeaker surfaces. The slope of the threshold shift defines an echo gain control that keeps echoes from point targets at a fixed sensation level--reducing sensitivity by 11 to 12 dB as echo amplitude increases by 12 dB per halving of range during the bat's approach to the target. A recent experiment using loudness discrimination of echoes at 70 to 80 dB SPL (roughly 50 dB above threshold) found a slope of about 6 dB per halving of range, so the gain-control effect may be level dependent. The observed effect is operationally equivalent to forward masking of echoes by the transmission, but any events correlated with vocalization which impair hearing sensitivity for a short interval following transmissions could cause a decline in sensitivity to echoes. Contractions of the bat's middle-ear muscles synchronized to transmissions may account for the observed threshold shift, at least for a span of echo delays associated with the most critical portion of the approach stage of pursuit. Forward masking by the sonar transmissions may contribute to the threshold shift, too, but middle-ear muscle contractions do occur and must be a significant part of the cause.

Animals↗

Perception of structured phantom targets in the echolocating bat, Megaderma lyra.

Bats can discriminate among objects with different texture using echolocation. The mechanisms involved in texture discrimination are unknown. In real echoes reflected from three-dimensional objects, the spatial structure of the target creates a specific temporal and spectral pattern. In the experiments described here, simply structured phantom targets mimicking echoes reflected from an object with two parallel planes were generated by adding two differentially delayed copies of the bat's vocalizations. The discrimination performance of Megaderma lyra was studied in a two-alternative, forced-choice experiment using phantom targets with different internal delays and relative amplitudes of the two copies. When the reference target to which the bats were trained was presented, targets differing in internal delay by about 1 microseconds were discriminated. However, discrimination performance was not a globally monotonic function of the internal delay of the unrewarded target. When both targets presented in a trial differed from the reference, the bats still preferred one of them. Changes in overall level of the phantom target echo had little effect on performance; however, performance dropped considerably if only one of the two copies in an echo was attenuated. A model framework is introduced to account for these results. Models based on frequency processing yield better approximations than does a model based on time differences. The model based on spectral correlation gives a unified description of all the data. Although time domain models for texture discrimination cannot be generally refuted, the results presented here show that the discrimination of target surface structure can be explained as a discrimination of echo spectra.

Animals↗

Passive acoustic localization of the Atlantic bottlenose dolphin using whistles and echolocation clicks.

A method for localization and tracking of calling marine mammals was tested under realistic field conditions that include noise, multipath, and arbitrarily located sensors. Experiments were performed in two locations using four and six hydrophones with captive Atlantic bottlenose dolphins (Tursiops truncatus). Acoustic signals from the animals were collected in the field using a digital acoustic data acquisition system. The data were then processed off-line to determine relative hydrophone positions and the animal locations. Accurate hydrophone position estimates are achieved by pinging sequentially from each hydrophone to all the others. A two-step least-squares algorithm is then used to determine sensor locations from the calibration data. Animal locations are determined by estimating the time differences of arrival of the dolphin signals at the different sensors. The peak of a matched filter output or the first cycle of the observed waveform is used to determine arrival time of an echolocation click. Cross correlation between hydrophones is used to determine inter-sensor time delays of whistles. Calculation of source location using the time difference of arrival measurements is done using a least-squares solution to minimize error. These preliminary experimental results based on a small set of data show that realistic trajectories for moving animals may be generated from consecutive location estimates.

Animals↗

Sensorimotor model of bat echolocation and prey capture.

A model of the bat sensorimotor system is developed using acoustics, signal processing, and control theory to illustrate the fundamental issues in accomplishing prey capture with echolocation. This model indicates that successful nonpredictive tracking of an ideal prey can be accomplished with a very simple system. Circular apertures approximate the mouth and ears for deriving acoustic beam patterns, using the big brown bat Eptesicus fuscus as a model. Fundamental and overtone frequency components in the emissions allow two simultaneous acoustic beams to be defined. A pair of nonlinear, time-variable, sampled-data controllers alter the bat's heading by applying yaw and pitch heading corrections. The yaw correction attempts to position the prey in the midsagittal plane by nulling the interaural intensity difference of the fundamental component. The pitch correction compares the intensities of the overtone and fundamental components and acts to null their difference. By initiating pitch correction when the overtone intensity first exceeds that of the fundamental, the ambiguity problem is solved and the prey is directed to the capture region. Simulations of passive prey capture indicate that the capture probability decreases as the prey speed increases. Both quick and sluggish prey are considered, with sluggish prey found to be caught with slightly better efficiency. The magnitude of the prey's lateral motion just prior to capture is observed to be an important factor determining capture. The presence of a blind stage is considered, during which the interference of the emission with the echo is assumed to disrupt any sonar information. The presence of such a blind stage is found to have negligible effect on capture efficiency.

Acoustics↗

Echolocation signals and transmission beam pattern of a false killer whale (Pseudorca crassidens).

The echolocation transmission beam pattern of a false killer whale (Pseudorca crassidens) was measured in the vertical and horizontal planes. A vertical array of seven broadband miniature hydrophones was used to measure the beam pattern in the vertical plane and a horizontal array of the same hydrophones was used in the horizontal plane. The measurements were performed in the open waters of Kaneohe Bay, Oahu, Hawaii, while the whale performed a target discrimination task. Four types of signals, characterized by their frequency spectra, were measured. Type-1 signals had a single low-frequency peak at 40 +/- 9 kHz and a low-amplitude shoulder at high frequencies. Type-2 signals had a bimodal frequency characteristic with a primary peak at 46 +/- 7 kHz and a secondary peak at 88 +/- 13 kHz. Type-3 signals were also bimodal but with a primary peak at 100 +/- 7 kHz and a secondary peak at 49 +/- 9 kHz. Type-4 signals had a single high-frequency peak at 104 +/- 7 kHz. The center frequency of the signals were found to be linearly correlated to the peak-to-peak source level, increasing with increasing source level. The major axis of the vertical beam was directed slightly downward between 0 and -5 degrees, in contrast to the +5 to 10 degrees for Tursiops and Delphinapterus. The beam in the horizontal plane was directed forward between 0 degrees and -5 degrees.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Acoustic basis for recognition of aspect-dependent three-dimensional targets by an echolocating bottlenose dolphin.

The relationships between acoustic features of target echoes and the cognitive representations of the target formed by an echolocating dolphin will influence the ease with which the dolphin can recognize a target. A blindfolded Atlantic bottlenose dolphin (Tursiops truncatus) learned to match aspect-dependent three-dimensional targets (such as a cube) at haphazard orientations, although with some difficulty. This task may have been difficult because aspect-dependent targets produce different echoes at different orientations, which required the dolphin to have some capability for object constancy across changes in echo characteristics. Significant target-related differences in echo amplitude, rms bandwidth, and distributions of interhighlight intervals were observed among echoes collected when the dolphin was performing the task. Targets could be classified using a combination of energy flux density and rms bandwidth by a linear discriminant analysis and a nearest centroid classifier. Neither statistical model could classify targets without amplitude information, but the highest accuracy required spectral information as well. This suggests that the dolphin recognized the targets using a multidimensional representation containing amplitude and spectral information and that dolphins can form stable representations of targets regardless of orientation based on varying sensory properties.

Acoustics↗