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.     

Echo features used by human listeners to discriminate among objects that vary in material or wall thickness: implications for echolocating dolphins

Echolocating dolphins extract object feature information from the acoustic parameters of echoes. To gain insight into which acoustic parameters are important for object discrimination, human listeners were presented with echoes from objects used in two discrimination tasks performed by dolphins: Hollow cylinders with varying wall thicknesses (+/-0.2, 0.3, 0.4, and 0.8 mm), and spheres made of different materials (steel, aluminum, brass, nylon, and glass). The human listeners performed as well or better than the dolphins at the task of discriminating between the standard object and the comparison objects on both the cylinders (humans=97.1%; dolphin=82.3%) and the spheres (humans= 86.6%; dolphin= 88.7%). The human listeners reported using primarily pitch and duration to discriminate among the cylinders, and pitch and timbre to discriminate among the spheres. Dolphins may use some of the same echo features as the humans to discriminate among objects varying in material or structure. Human listening studies can be used to quickly identify salient combinations of echo features that permit object discrimination, which can then be used to generate hypotheses that can be tested using dolphins as subjects.     

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.     

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.     

Neural time and movement time in choice of whistle or pulse burst responses to different auditory stimuli by dolphins

Echolocating dolphins emit trains of clicks and receive echoes from ocean targets. They often emit each successive ranging click about 20 ms after arrival of the target echo. In echolocation, decisions must be made about the target--fish or fowl, predator or food. In the first test of dolphin auditory decision speed, three bottlenose dolphins (Tursiops truncatus) chose whistle or pulse burst responses to different auditory stimuli randomly presented without warning in rapid succession under computer control. The animals were trained to hold pressure catheters in the nasal cavity so that pressure increases required for sound production could be used to split response time (RT) into neural time and movement time. Mean RT in the youngest and fastest dolphin ranged from 175 to 213 ms when responding to tones and from 213 to 275 ms responding to pulse trains. The fastest neural times and movement times were around 60 ms. The results suggest that echolocating dolphins tune to a rhythm so that succeeding pulses in a train are produced about 20 ms over target round-trip travel time. The dolphin nervous system has evolved for rapid processing of acoustic stimuli to accommodate for the more rapid sound speed in water compared to air.     

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.     

Acoustic features of objects matched by an echolocating bottlenose dolphin

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.     

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.     

Detection of complex echoes in noise by an echolocating dolphin

Dolphins echolocate with short broadband acoustic signals that have good time resolution properties. Received echoes are often complex, with many resolvable highlights or components caused by reflection of the incident signal from external and internal boundaries of a target and from different propagational modes within a target. A series of experiments was performed to investigate how dolphins perceive complex echoes. Echoes were produced by a microprocessor-controlled electronic target simulator that captured each emitted click and retransmitted the signal back to the animal after an appropriate time delay. The use of this "phantom" target allowed for precise control of the number of highlights, the time separation between highlights, and the relative amplitudes of highlights in the simulated echoes. An echolocating dolphin was trained to perform a target detection task in the presence of masking noise using these phantom echoes. The properties of simulated echoes were systematically varied, and corresponding shifts in the dolphin's detection threshold were observed, allowing for inferences of how the dolphin perceived echoes. The dolphin performed like an energy detector with an integration time of approximately 264 microseconds.     

Vocal control of acoustic information for sonar discriminations by the echolocating bat, Eptesicus fuscus

This study aimed to determine whether bats using frequency modulated (FM) echolocation signals adapt the features of their vocalizations to the perceptual demands of a particular sonar task. Quantitative measures were obtained from the vocal signals produced by echolocating bats (Eptesicus fuscus) that were trained to perform in two distinct perceptual tasks, echo delay and Doppler-shift discriminations. In both perceptual tasks, the bats learned to discriminate electronically manipulated playback signals of their own echolocation sounds, which simulated echoes from sonar targets. Both tasks utilized a single-channel electronic target simulator and tested the bat's in a two-alternative forced choice procedure. The results of this study demonstrate changes in the features of the FM bats' sonar sounds with echolocation task demands, lending support to the notion that this animal actively controls the echo information that guides its behavior.     

Phantom echo highlight amplitude and temporal difference resolutions of an echolocating dolphin, Tursiops truncatus

A dolphin's ability to discriminate targets may depend greatly on the relative amplitudes and the time separations of echo highlights within the received signal. Previous experiments with dolphins varied the physical parameters of targets, but did not fully investigate how changes in these parameters correspond with the scattered acoustic wave forms and the dolphin's subsequent response. This experiment utilizes a phantom echo system to test a dolphin's detection response to relative amplitude differences of secondary and trailing echo highlights and the time separation differences of all the echo highlights both within and outside the animal's integration window. By electronically manipulating the amplitude and temporal separation of the echo highlights, the underlying acoustic classification cues are more efficiently investigated. The animal successfully discriminated between a standard echo signal and one with the secondary highlight amplitude lowered by 7 dB from the standard. Furthermore, the animal successfully discriminated between a standard echo signal and one with the trailing highlight amplitude lowered by 3 dB from the standard and also a standard echo signal and one with a time separation of 150 mus between the secondary and trailing highlights.     

Classification of broadband echoes from prey of a foraging Blainville's beaked whale

Blainville's beaked whales (Mesoplodon densirostris) use broadband, ultrasonic echolocation signals with a -10 dB bandwidth from 26 to 51 kHz to search for, localize, and approach prey that generally consist of mid-water and deep-water fishes and squid. Although it is well known that the spectral characteristics of broadband echoes from marine organisms vary as a function of size, shape, orientation, and anatomical group, there is little evidence as to whether or not free-ranging toothed whales use spectral cues in discriminating between prey and nonprey. In order to study the prey-classification process, a stereo acoustic tag was deployed on a Blainville's beaked whale so that emitted clicks and the corresponding echoes from targets in the water could be recorded. A comparison of echoes from targets apparently selected by the whale and those from a sample of scatterers that were not selected suggests that spectral features of the echoes, target strengths, or both may have been used by the whale to discriminate between echoes. Specifically, the whale appears to favor targets with one or more nulls in the echo spectra and to seek prey with higher target strengths at deeper depths.     

An echolocation visualization and interface system for dolphin research

The present study describes the development and testing of a tool for dolphin research. This tool was able to visualize the dolphin echolocation signals as well as function as an acoustically operated "touch screen." The system consisted of a matrix of hydrophones attached to a semitransparent screen, which was lowered in front of an underwater acrylic panel in a dolphin pool. When a dolphin aimed its sonar beam at the screen, the hydrophones measured the received sound pressure levels. These hydrophone signals were then transferred to a computer where they were translated into a video image that corresponds to the dynamic sound pressure variations in the sonar beam and the location of the beam axis. There was a continuous projection of the image back onto the hydrophone matrix screen, giving the dolphin an immediate visual feedback to its sonar output. The system offers a whole new experimental methodology in dolphin research and since it is software-based, many different kinds of scientific questions can be addressed. The results were promising and motivate further development of the system and studies of sonar and cognitive abilities of dolphins.     

Assessment of dolphin (Tursiops truncatus) auditory sensitivity and hearing loss using jawphones

Devices known as jawphones have previously been used to measure interaural time and intensity discrimination in dolphins. This study introduces their use for measuring hearing sensitivity in dolphins. Auditory thresholds were measured behaviorally against natural background noise for two bottlenose dolphins (Tursiops truncatus); a 14-year-old female and a 33-year-old male. Stimuli were delivered to each ear independently by placing jawphones directly over the pan bone of the dolphin's lower jaw, the assumed site of best reception. The shape of the female dolphin's auditory functions, including comparison measurements made in the free field, favorably matches that of the accepted standard audiogram for the species. Thresholds previously measured for the male dolphin at 26 years of age indicated a sensitivity difference between the ears of 2-3 dB between 4-10 kHz, which was considered unremarkable at the time. Thresholds for the male dolphin reported in this study suggest a high-frequency loss compared to the standard audiogram. Both of the male's ears have lost sensitivity to frequencies above 55 kHz and the right ear is 16-33 dB less sensitive than the left ear over the 10-40 kHz range, suggesting that males of the species may lose sensitivity as a function of age. The results of this study support the use of jawphones for the measurement of dolphin auditory sensitivity.     

Time-frequency analysis and modeling of the backscatter of categorized dolphin echolocation clicks for target discrimination

A set of dolphin echolocation clicks collected from an Atlantic bottlenose dolphin in Kaneohe Bay, Hawai'i from a previous experiment is examined in terms of their time and frequency characteristics. The center frequency and rms bandwidth are calculated for the clicks and these are clustered into four classes by using a model based on the Bayesian information criterion. The echo signatures are attained from a solid, elastic homogeneous sphere for each class of clicks from an acoustic scattering model. The results from the scattering model are compared to experimental values. The joint time-frequency content of the resulting echo signals is obtained by the reduced interference distribution (RID). The RIDs are plotted and examined for each signal class for four spherical targets of different material compositions. RID correlation values are obtained for a standard target versus comparison targets by using a time-frequency correlator. The results suggest that dolphins may discriminate by auditory inspection of the time-frequency information returned by the targets. The modification of the outgoing clicks and examination of time-frequency target information may be fundamental to a dolphin's ability to identify and discriminate targets.     

Range discrimination by big brown bats (Eptesicus fuscus) using altered model echoes: implications for signal processing

The sonar emissions of two big brown bats (Eptesicus fuscus) were modeled to create a "normal" echolocation signal for each bat which was then used as an artificial echo to synthesize a phantom target. The bat's task was to indicate which of two phantom targets (presented singly) was the "near" target and which the "far" target. Threshold range discrimination at a nominal target distance of 80 cm was about 0.6 cm for both bats. The normal signal was then modified to change the relative energy in each harmonic, the signal duration, the curvature of the frequency sweep, the absolute frequency, the phase of the second and third harmonics relative to the first, or the Doppler shift of the signal. To determine which modifications affected ranging performance, the altered models were used in tests of range discrimination that were interleaved on a day-to-day basis with tests using the normal model. Of the 12 modifications tested, only those changing the curvature of the frequency sweep affected performance. This result appears not to be predicted by current models of echo processing in FM bats. Eptesicus may be able to compensate for certain types of distortions of a returning echo, an ability possibly related to Doppler tolerance or to the characteristics of the natural variation in a bat's emissions.     

Off-axis sonar beam pattern of free-ranging finless porpoises measured by a stereo pulse event data logger

The off-axis sonar beam patterns of eight free-ranging finless porpoises were measured using attached data logger systems. The transmitted sound pressure level at each beam angle was calculated from the animal's body angle, the water surface echo level, and the swimming depth. The beam pattern of the off-axis signals between 45 degrees and 115 degrees (where 0 degrees corresponds to the on-axis direction) was nearly constant. The sound pressure level of the off-axis signals reached 162 dB re 1 microPa peak-to-peak. The surface echo level received at the animal was over 140 dB, much higher than the auditory threshold level of small odontocetes. Finless porpoises are estimated to be able to receive the surface echoes of off-axis signals even at 50-m depth. Shallow water systems (less than 50-m depth) are the dominant habitat of both oceanic and freshwater populations of this species. Surface echoes may provide porpoises not only with diving depth information but also with information about surface direction and location of obstacles (including prey items) outside the on-axis sector of the sonar beam.     

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.     

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.     

Echolocation signals of wild dolphins

Most of our understanding of dolphin echolocation has come from studies of captive dolphins performing various echolocation tasks. Recently, measurements of echolocation signals in the wild have expanded our understanding of the characteristics of these signals in a natural setting. Measuring undistorted dolphin echolocation signals with free swimming dolphins in the field can be a challenging task. A four hydrophone array arranged in a symmetrical star pattern was used to measure the echolocation signals of four species of dolphins in the wild. Echolocation signals of the following dolphins have been measured with the symmetrical star array: white-beaked dolphins in Iceland, Atlantic spotted dolphins in the Bahamas, killer whales in British Columbia, and dusky dolphins in New Zealand. There are many common features in the echolocation signals of the different species. Most of the signals had spectra that were bimodal: two peaks, one at low frequencies and another about an octave higher in frequency. The source level of the sonar transmission varies as a function of 20logR, suggesting a form of time-varying gain but on the transmitting end of the sonar process rather than the receiving end. The results of the field work call into question the issue of whether the signals used by captive dolphins may be shaped by the task they are required to perform rather than what they would do more naturally.