Dolphin echo-delay resolution measured with a jittered-echo paradigm.

Biosonar echo delay resolution was investigated in four bottlenose dolphins (Tursiops truncatus) using a "jittered" echo paradigm, where dolphins discriminated between electronic echoes with fixed delay and those whose delay alternated (jittered) on successive presentations. The dolphins performed an echo-change detection task and produced a conditioned acoustic response when detecting a change from non-jittering echoes to jittering echoes. Jitter delay values ranged from 0 to 20 µs. A passive listening task was also conducted, where dolphins listened to simulated echoes and produced a conditioned acoustic response when signals changed from non-jittering to jittering. Results of the biosonar task showed a mean jitter delay threshold of 1.3 µs and secondary peaks in error functions suggestive of the click autocorrelation function. When echoes were jittered in polarity and delay, error functions shifted by approximately 5 µs and all dolphins discriminated echoes that jittered only in polarity. Results were qualitatively similar to those from big brown bats (Eptesicus fuscus) and indicate that the dolphin biosonar range estimator is sensitive to echo phase information. Results of the passive listening task suggested that the dolphins could not passively detect changes in timing and polarity of simulated echoes.

Detailed analysis of two detected overlaying transient components within the echolocation beam of a bottlenose dolphin (Tursiops truncatus).

Dolphin echolocation clicks measured far off-axis contain two time-separated components. Whether these components overlap and appear as a single signal on axis has received little attention. Here, the scaled reassigned spectrogram analysis was used to examine if bottlenose dolphin (Tursiops truncatus) clicks measured near- or on-axis of the echolocation beam contained overlapping components. Across click trains, the number of overlapping components spatially varied within the echolocation beam. Two overlapping components were found to predominantly occur in the upper portion of the beam, whereas the lower portion of the beam predominantly contained a single component. When components overlapped, the trailing component generally had a higher center frequency and arrived less than 5 µs after the leading component. The spatial relationship of components was consistent with previous findings of two vertically distinct beam lobes with separated frequency content. The two components in the upper portion of the beam possibly result from a single transient click propagating through a geometrically dispersive media; specifically, the slower sound speed of the dolphin melon's core slightly delays the more directional, high frequency energy of the click, whereas the less directional, lower frequency energy propagates through more peripheral but higher sound speed portions of the melon.

Jittered echo-delay resolution in bottlenose dolphins (Tursiops truncatus).

Psychophysical methods similar to those employed with bats were used to examine jittered echo-delay resolution in bottlenose dolphins (Tursiops truncatus). Two dolphins were trained to produce echolocation clicks and report a change from electronic echoes with a fixed delay of ~ 12.6 ms (~ 9.4 m simulated range) to echoes with delays that alternated (jittered) between successive emitted signals. Jitter delays varied from 0 to 50 µs. Jittered echo-delay thresholds were between 1 and 2 µs-the lowest achievable (non-zero) values with the hardware configuration. Error functions matched the click autocorrelation function near zero jitter delay, and were well within the envelope of the autocorrelation function; however, measured jitter delay thresholds were larger than predictions for a coherent or semicoherent receiver at comparable signal-to-noise ratios. When one of the two alternating jittered echoes was inverted in polarity, both dolphins reliably discriminated echoes at all jittered echo delays, including 0 µs (i.e., only jittering in polarity, not delay). Finally, both dolphins used unusual patterns of click emissions, where groups of echolocation clicks were interspersed with silent gaps. Further tests with sub-microsecond jitter values and various echo signal-to-noise ratios would be necessary for proper direct comparison with jitter detection values obtained for bats.

Dynamics of biosonar signals in free-swimming and stationary dolphins: The role of source levels on the characteristics of the signals.

The biosonar signals of two free-swimming Atlantic bottlenose dolphins performing a complex sonar search for a bottom target in San Diego Bay were compared with the biosonar signals of a dolphin performing a target discrimination task in a net pen in the same bay. A bite-plate device carried by the free-swimming dolphins supported a hydrophone that extended directly in front of the dolphin. A biosonar measuring tool attached to the bite plate measured the outgoing biosonar signals while the dolphins conducted sonar searches. Each of the free-swimming dolphins used different biosonar search strategy in solving the problem and the dolphins' biosonar signals reflect the difference in strategy. The dolphin in the pen stationed in a hoop while echolocating on a target 6 m away and reported if the indentation on a spherical target was directed toward it. The signals were parameterized by determining the peak-to-peak source levels, source energy flux density, peak frequency, center frequency, root-mean-square (rms) bandwidth, rms duration, and the Q of the signals. Some parameters were similar for the free-swimming and stationary dolphins while some were significantly different, suggesting biosonar signals used by free-swimming animals may be different than signals used by dolphins in a pen.

High-resolution measurement of a bottlenose dolphin's (Tursiops truncatus) biosonar transmission beam pattern in the horizontal plane.

Previous measurements of toothed whale echolocation transmission beam patterns have utilized few hydrophones and have therefore been limited to fine angular resolution only near the principal axis or poor resolution over larger azimuthal ranges. In this study, a circular, horizontal planar array of 35 hydrophones was used to measure a dolphin's transmission beam pattern with 5° to 10° resolution at azimuths from -150° to +150°. Beam patterns and directivity indices were calculated from both the peak-peak sound pressure and the energy flux density. The emitted pulse became smaller in amplitude and progressively distorted as it was recorded farther off the principal axis. Beyond ±30° to 40°, the off-axis signal consisted of two distinct pulses whose difference in time of arrival increased with the absolute value of the azimuthal angle. A simple model suggests that the second pulse is best explained as a reflection from internal structures in the dolphin's head, and does not implicate the use of a second sound source. Click energy was also more directional at the higher source levels utilized at longer ranges, where the center frequency was elevated compared to that of the lower amplitude clicks used at shorter range.

Dolphin biosonar signals measured at extreme off-axis angles: insights to sound propagation in the head.

Biosonar signals radiated along the beam axis of an Atlantic bottlenose dolphin resemble short transient oscillations. As the azimuth of the measuring hydrophones in the horizontal plane progressively increases with respect to the beam axis the signals become progressively distorted. At approximately ±45°, the signals begin to divide into two components with the time difference between the components increasing with increasing angles. At ±90° or normal to the longitudinal axis of the animal, the time difference between the two pulses measured by the hydrophone on the right side of the dolphin's head is, on average, ∼11.9 µs larger than the time differences observed by the hydrophone on the left side of the dolphin's head. The center frequency of the first pulse is generally lower, by 33-47 kHz, than the center frequency of the second pulse. When considering the relative locations of the two phonic lips, the data suggest that the signals are being produced by one of the phonic lips and the second pulse resulting from a reflection within the head of the animal. The generation of biosonar signals is a complex process and the propagation pathways through the dolphin's head are not well understood.

Directional properties of bottlenose dolphin (Tursiops truncatus) clicks, burst-pulse, and whistle sounds.

The directional properties of bottlenose dolphin clicks, burst-pulse, and whistle signals were measured using a five element array, at horizontal angles of 0°, 45°, 90°, 135°, and 180° relative to a dolphin stationed on an underwater biteplate. Clicks and burst-pulse signals were highly directional with directivity indices of ~11 dB for both signal types. Higher frequencies and higher amplitudes dominated the forward, on-axis sound field. A similar result was found with whistles, where higher frequency harmonics had greater directivity indices than lower frequency harmonics. The results suggest the directional properties of these signals not only provide enhanced information to the sound producer (as in echolocation) but can provide valuable information to conspecific listeners during group coordination and socialization.

The biosonar field around an Atlantic bottlenose dolphin (Tursiops truncatus).

The use of remote autonomous passive acoustic recorders (PAR) to determine the distribution of dolphins at a given locations has become very popular. Some investigators are using echolocation clicks to gather information on the presence of dolphins and to identify species. However, in all of these cases, the PAR probably recorded mainly off-axis clicks, even some from behind the animals. Yet there is a very poor understanding of the beam pattern and the click waveform and spectrum from different azimuths around the animal's body. The beam pattern completely around an echo locating dolphin was measured at 16 different but equally spaced angles in the horizontal plane using an 8-hydrophone array in sequence. Eight channels of data were digitized simultaneously at a sampling rate of 500 kHz. The resulting beam patterns in both planes showed a continuous drop off in sound pressure with azimuth around the animal and reached levels below -50 dB relative to the signal recorded on the beam axis. The signals began to break up into two components at angles greater than ± 45° in the horizontal plane. The center frequency dropped off from its maximum at 0° in a non-uniform matter.

Frequency-dependent variation in the two-dimensional beam pattern of an echolocating dolphin.

Recent recordings of dolphin echolocation using a dense array of hydrophones suggest that the echolocation beam is dynamic and can at times consist of a single dominant peak, while at other times it consists of forward projected primary and secondary peaks with similar energy, partially overlapping in space and frequency bandwidth. The spatial separation of the peaks provides an area in front of the dolphin, where the spectral magnitude slopes drop off quickly for certain frequency bands. This region is potentially used to optimize prey localization by directing the maximum pressure slope of the echolocation beam at the target, rather than the maximum pressure peak. The dolphin was able to steer the beam horizontally to a greater extent than previously described. The complex and dynamic sound field generated by the echolocating dolphin may be due to the use of two sets of phonic lips as sound sources, or an unknown complexity in the sound propagation paths or acoustic properties of the forehead tissues of the dolphin.

The acoustic field on the forehead of echolocating Atlantic bottlenose dolphins (Tursiops truncatus).

Arrays of up to six broadband suction cup hydrophones were placed on the forehead of two bottlenose dolphins to determine the location where the beam axis emerges and to examine how signals in the acoustic near-field relate to signals in the far-field. Four different array geometries were used; a linear one with hydrophones arranged along the midline of the forehead, and two around the front of the melon at 1.4 and 4.2 cm above the rostrum insertion, and one across the melon in certain locations not measured by other configurations. The beam axis was found to be close to the midline of the melon, approximately 5.4 cm above the rostrum insert for both animals. The signal path coincided with the low-density, low-velocity core of the melon; however, the data suggest that the signals are focused mainly by the air sacs. Slight asymmetry in the signals were found with higher amplitudes on the right side of the forehead. Although the signal waveform measured on the melon appeared distorted, when they are mathematically summed in the far-field, taking into account the relative time of arrival of the signals, the resultant waveform matched that measured by the hydrophone located at 1 m.

Relationship of blood flow and metabolism to acoustic processing centers of the dolphin brain.

Odontocete brain tissues associated with auditory processing are hypertrophied and modified relative to their terrestrial counterparts. The relationship between the functional demand on these tissues and metabolic substrate requirements is unknown. Using positron emission tomography (PET), relative cerebral blood flow was measured in a bottlenose dolphin. Approximately 60 mCi (13)NH(3) was administered to the dolphin via a catheter inserted into the hepatic vein and threaded proximate to the vena cava. Radiolabel initially appeared as distributed focal points in the cerebellum. Increasing scan time resulted in an increase in the number of focal regions and in the diffusivity of label activity throughout the brain. The time course and spatial distribution of radiolabel was consistent with a cerebral blood supply dominated by the spinal meningeal arteries. Blood flow was predominantly observed in the cerebellum and neocortex, particularly the auditory and visual cortex. Differential brain glucose uptake, previously measured in a separate dolphin, showed good agreement with the differential supply of blood to brain tissues. Rates of blood supply and glucose uptake in the auditory cortex, inferior colliculus, and cerebellum are consistent with a high metabolic demand of tissues which are important to the integration of auditory and other sensory inputs.

A method to enable a bottlenose dolphin (Tursiops truncatus) to echolocate while out of water.

The study of site-specific brain activity associated with dolphin echolocation has been hampered by the difficulties inherent in administering radiolabels and performing medical imaging while a dolphin echolocates in an aquatic environment. To overcome these limitations, a system has been developed to allow a bottlenose dolphin to echolocate while out of the water. The system relies on a "phantom echo generator" (PEG) consisting of a Texas Instruments C6713 digital signal processor with an analog input/output daughtercard. Echolocation clicks produced by the dolphin are detected with a hydrophone embedded in a suction cup on the melon, then digitized within the PEG. Clicks exceeding a user-defined threshold are convolved with a target impulse response, delayed, and scaled before being converted to analog and transmitted through a sound projector embedded in a suction cup attached to the dolphin's lower jaw. Dolphin in-air echolocation behavior, inter-click intervals, and overall performance were analogous to those observed during comparable underwater testing with physical targets, demonstrating that the dolphin was indeed performing an echolocation task while out of water.

Beamwidth control and angular target detection in an echolocating bottlenose dolphin (Tursiops truncatus).

Bottlenose dolphin (Tursiops truncatus) echolocation beams are typically characterized as symmetrical -3 dB beamwidths; however, the functional width of the beam during target detection has not been explored. Angular target detection thresholds of an echolocating dolphin were examined to more fully describe the functional characteristics of the echolocation beam. The dolphin performed an echolocation detection task with its head held in a fixed orientation. Targets were placed 9 m in front of the dolphin [0 degrees position (P(0))] and systematically moved right or left until target detection reached chance probability. A 24-element hydrophone array placed 1 m in front of the dolphin was used to measure vertical and horizontal echolocation beamwidths. Detection thresholds were 26 degrees left and 21 degrees right of P(0) for a sphere target and 19 degrees left and 13 degrees right of P(0) for a cylinder target. Estimates of maximum horizontal and vertical beamwidths ranged up to 40 degrees and 29 degrees , respectively, and exhibited large variability. The dolphin nominally steered the maximum response axis of the echolocation beam up to 18 degrees in the horizontal, 12 degrees in the upward vertical, and 4 degrees in the downward vertical. These results suggest that the dolphin can steer and modify the width of the echolocation beam.

Bio-inspired wideband sonar signals based on observations of the bottlenose dolphin (Tursiops truncatus).

This paper uses advanced time-frequency signal analysis techniques to generate new models for bio-inspired sonar signals. The inspiration comes from the analysis of bottlenose dolphin clicks. These pulses are very short duration, between 50 and 80 micros, but for certain examples we can delineate a double down-chirp structure using fractional Fourier methods. The majority of clicks have energy distributed between two main frequency bands with the higher frequencies delayed in time by 5-20 micros. Signal syntheses using a multiple chirp model based on these observations are able to reproduce much of the spectral variation seen in earlier studies on natural dolphin echolocation pulses. Six synthetic signals are generated and used to drive the dolphin based sonar (DBS) developed through the Biosonar Program office at the SPAWAR Systems Center, San Diego, CA. Analyses of the detailed echo structure for these pulses ensonifying two solid copper spherical targets indicate differences in discriminatory potential between the signals. It is suggested that target discrimination could be improved through the transmission of a signal packet in which the chirp structure is varied between pulses. Evidence that dolphins may use such a strategy themselves comes from observations of variations in the transmissions of dolphins carrying out target detection and identification tasks.

Using a binaural biomimetic array to identify bottom objects ensonified by echolocating dolphins.

The development of a unique dolphin biomimetic sonar produced data that were used to study signal processing methods for object identification. Echoes from four metallic objects proud on the bottom, and a substrate-only condition, were generated by bottlenose dolphins trained to ensonify the targets in very shallow water. Using the two-element ('binaural') receive array, object echo spectra were collected and submitted for identification to four neural network architectures. Identification accuracy was evaluated over two receive array configurations, and five signal processing schemes. The four neural networks included backpropagation, learning vector quantization, genetic learning and probabilistic network architectures. The processing schemes included four methods that capitalized on the binaural data, plus a monaural benchmark process. All the schemes resulted in above-chance identification accuracy when applied to learning vector quantization and backpropagation. Beam-forming or concatenation of spectra from both receive elements outperformed the monaural benchmark, with higher sensitivity and lower bias. Ultimately, best object identification performance was achieved by the learning vector quantization network supplied with beam-formed data. The advantages of multi-element signal processing for object identification are clearly demonstrated in this development of a first-ever dolphin biomimetic sonar.

Instrumenting free-swimming dolphins echolocating in open water.

Dolphins within the Navy Marine Mammal Program use echolocation to effectively locate underwater mines. They currently outperform manmade systems at similar tasks, particularly in cluttered environments and on buried targets. In hopes of improving manmade mine-hunting sonar systems, two instrumentation packages were developed to monitor free-swimming dolphin motion and echolocation during open-water target detection tasks. The biosonar measurement tool (BMT) is carried by a dolphin and monitors underwater position and attitude while simultaneously recording echolocation clicks and returning echoes through high-gain binaural receivers. The instrumented mine simulator (IMS) is a modified bottom target that monitors echolocation signals arriving at the target during ensonification. Dolphin subjects were trained to carry the BMT in open-bay bottom-object target searches in which the IMS could serve as a bottom object. The instrumentation provides detailed data that reveal hereto-unavailable information on the search strategies of free-swimming dolphins conducting open-water, bottom-object search tasks with echolocation.

Echolocation characteristics of free-swimming bottlenose dolphins during object detection and identification.

A biosonar measurement tool (BMT) was created to investigate dolphin echolocation search strategies by recording echolocation clicks, returning echoes, and three-dimensional angular motion, velocity, and depth of free-swimming dolphins performing open-water target detections. Trial start and stop times, locations determined from a differential global positioning system (DGPS), and BMT motion and acoustic data were used to produce spatial and acoustic representations of the searches. Two dolphins (LUT, FLP) searched for targets lying on the seafloor of a bay environment while carrying the BMT. LUT searched rapidly (< 10 s), produced few clicks, and varied click-peak frequency (20-120 kHz); FLP searched relatively slowly (tens of seconds) and produced many hundreds of clicks with stereotypical frequency-dependent energy distributions dominating from 30-60 kHz. Dolphins amplified target echo returns by either increasing the click source level or reducing distance to the target but without reducing source level. The distribution of echolocation click-peak frequencies suggested a bias in the dominant frequency components of clicks, possibly due to mechanical constraints of the click generator. Prior training and hearing loss accommodation potentially explain differences in the search strategies of the two dolphins.

Multiecho processing by an echolocating dolphin.

Bottlenose dolphins (Tursiops truncatus) use short, wideband pulses for echolocation. Individual waveforms have high-range resolution capability but are relatively insensitive to range rate. Signal-to-noise ratio (SNR) is not greatly improved by pulse compression because each waveform has small time-bandwidth product. The dolphin, however, often uses many pulses to interrogate a target, and could use multipulse processing to combine the resulting echoes. Multipulse processing could mitigate the small SNR improvement from pulse compression, and could greatly improve range-rate estimation, moving target indication, range tracking, and acoustic imaging. All these hypothetical capabilities depend upon the animal's ability to combine multiple echoes for detection and/or estimation. An experiment to test multiecho processing in a dolphin measured detection of a stationary target when the number N of available target echoes was increased, using synthetic echoes. The SNR required for detection decreased as the number of available echoes increased, as expected for multiecho processing. A receiver that sums binary-quantized data samples from multiple echoes closely models the N dependence of the SNR required by the dolphin. Such a receiver has distribution-tolerant (nonparametric) properties that make it robust in environments with nonstationary and/or non-Gaussian noise, such as the pulses created by snapping shrimp.

Discrimination of complex synthetic echoes by an echolocating bottlenose dolphin.

Bottlenose dolphins (Tursiops truncatus) detect and discriminate underwater objects by interrogating the environment with their native echolocation capabilities. Study of dolphins' ability to detect complex (multihighlight) signals in noise suggest echolocation object detection using an approximate 265-micros energy integration time window sensitive to the echo region of highest energy or containing the highlight with highest energy. Backscatter from many real objects contains multiple highlights, distributed over multiple integration windows and with varying amplitude relationships. This study used synthetic echoes with complex highlight structures to test whether high-amplitude initial highlights would interfere with discrimination of low-amplitude trailing highlights. A dolphin was trained to discriminate two-highlight synthetic echoes using differences in the center frequencies of the second highlights. The energy ratio (delta dB) and the timing relationship (delta T) between the first and second highlights were manipulated. An iso-sensitivity function was derived using a factorial design testing delta dB at -10, -15, -20, and -25 dB and delta T at 10, 20, 40, and 80 micros. The results suggest that the animal processed multiple echo highlights as separable analyzable features in the discrimination task, perhaps perceived through differences in spectral rippling across the duration of the echoes.

Discrimination of amplitude-modulated synthetic echo trains by an echolocating bottlenose dolphin.

Bottlenose dolphins (Tursiops truncatus) have an acute ability to use target echoes to judge attributes such as size, shape, and material composition. Most target recognition studies have focused on features associated with individual echoes as opposed to information conveyed across echo sequences (feature envelope of the multi-echo train). One feature of aspect-dependent targets is an amplitude modulation (AM) across the return echoes in the echo train created by relative movement of the target and dolphin. The current study examined whether dolphins could discriminate targets with different AM envelopes. "Electronic echoes" triggered by a dolphin's outgoing echolocation clicks were manipulated to create sinusoidal envelopes with varying AM rate and depth. Echo trains were equated for energy, requiring the dolphin to extract and retain information from multiple echoes in order to detect and report the presence of AM. The dolphin discriminated amplitude-modulated echo trains from those that were not modulated. AM depth thresholds were approximately 0.8 dB, similar to other published amplitude limens. Decreasing the rate of modulation from approximately 16 to 2 cycles per second did not affect the dolphin's AM depth sensitivity. The results support multiple-echo processing in bottlenose dolphin echolocation. This capability provides additional theoretical justification for exploring synthetic aperture sonar concepts in models of animal echolocation that potentially support theories postulating formation of images as an ultimate means for target identification.