A re-evaluation of auditory filter shape in delphinid odontocetes: evidence of constant-bandwidth filters.

The auditory filter shape of delphinid odontocetes was previously considered to be typically mammalian constant-quality in which filter bandwidths increase proportionally with frequency. Recent studies with porpoises demonstrate constant-bandwidth portions of the auditory filter. The critical ratios for a bottlenose dolphin were measured between 40 and 120 kHz by behaviorally determining the subject's ability to detect pure tones in the presence of white noise. Critical ratios as a function of frequency were constant, indicating the auditory filter acts as a constant-bandwidth system in this frequency range. Re-analysis of past studies supports these findings, and suggests the delphinid auditory system is best characterized as a constant-Q system below 40 kHz and a constant-bandwidth-like system between 40 kHz and 120 kHz before returning to a constant Q pattern at the highest frequencies.

Decreased echolocation performance following high-frequency hearing loss in the false killer whale (Pseudorca crassidens).

Toothed whales and dolphins possess a hypertrophied auditory system that allows for the production and hearing of ultrasonic signals. Although the fossil record provides information on the evolution of the auditory structures found in extant odontocetes, it cannot provide information on the evolutionary pressures leading to the hypertrophied auditory system. Investigating the effect of hearing loss may provide evidence for the reason for the development of high-frequency hearing in echolocating animals by demonstrating how high-frequency hearing assists in the functioning echolocation system. The discrimination abilities of a false killer whale (Pseudorca crassidens) were measured prior to and after documented high-frequency hearing loss. In 1992, the subject had good hearing and could hear at frequencies up to 100 kHz. In 2008, the subject had lost hearing at frequencies above 40 kHz. First in 1992, and then again in 2008, the subject performed an identical echolocation task, discriminating between machined hollow aluminum cylinder targets of differing wall thickness. Performances were recorded for individual target differences and compared between both experimental years. Performances on individual targets dropped between 1992 and 2008, with a maximum performance reduction of 36.1%. These data indicate that, with a loss in high-frequency hearing, there was a concomitant reduction in echolocation discrimination ability, and suggest that the development of a hypertrophied auditory system capable of hearing at ultrasonic frequencies evolved in response to pressures for fine-scale echolocation discrimination.

Audiogram of a formerly stranded long-finned pilot whale (Globicephala melas) measured using auditory evoked potentials.

Long-finned pilot whales are highly social odontocetes found in temperate and subpolar regions. This species is particularly known for its interaction with fisheries as well as its mass strandings. Recent tagging work has provided some information about pilot whales in the wild but, even though they have been successfully kept in captivity, little is known about their sensory capabilities. This study investigates the hearing abilities of a rehabilitated 2 year old male long-finned pilot whale. A complete audiogram was collected using auditory evoked potential techniques that included measurements of nine frequencies from 4 to 100 kHz presented as sinusoidally amplitude-modulated tones. The results indicated that the region of best hearing was between 11.2 and 50 kHz and the subject had relatively poor high frequency hearing compared with other odontocete species. This study emphasizes the importance of collecting basic hearing measurements from new species, understanding diagnostic life histories as well as continuously increasing the sample size of audiometry measurements within and between odontocete species as animals become available.

Shipboard measurements of the hearing of the white-beaked dolphin Lagenorhynchus albirostris.

This is the first report of an underwater audiogram from a dolphin in a capture-and-release scenario. Two bow-riding white-beaked dolphins Lagenorhynchus albirostris (a female and a male) were captured using the hoop-net technique in Faxaflói Bay, Iceland. The dolphins were transferred to a stretcher and hoisted into a plastic research tank on board a small fishing vessel. Two underwater transducers were used to cover the frequency range from 16 to 215 kHz. Two human EEG electrodes mounted in suction cups, one placed near the blow hole and the other on the dorsal fin, picked up bioelectrical responses to acoustic stimuli. Responses to about 1000 sinusoidal amplitude modulated stimuli for each amplitude/frequency combination were averaged and analyzed using a fast Fourier transform to obtain an evoked auditory response. Threshold was defined as the zero crossing of the response using linear regression. Two threshold frequencies at 50 kHz and 64 kHz were obtained from the female. An audiogram ranging from 16 to 181 kHz was obtained from an adult male and showed the typical ;U' shaped curve for odontocetes. The thresholds for both white-beaks were comparable and demonstrated the most sensitive high frequency hearing of any known dolphin and were as sensitive as the harbor porpoise.

Classification of electronically generated phantom targets by an Atlantic bottlenose dolphin (Tursiops truncatus).

Animal behavior experiments require not only stimulus control of the animal's behavior, but also precise control of the stimulus itself. In discrimination experiments with real target presentation, the complex interdependence between the physical dimensions and the backscattering process of an object make it difficult to extract and control relevant echo parameters separately. In other phantom-echo experiments, the echoes were relatively simple and could only simulate certain properties of targets. The echo-simulation method utilized in this paper can be used to transform any animal echolocation sound into phantom echoes of high fidelity and complexity. The developed phantom-echo system is implemented on a digital signal-processing board and gives an experimenter fully programmable control over the echo-generating process and the echo structure itself. In this experiment, the capability of a dolphin to discriminate between acoustically simulated phantom replicas of targets and their real equivalents was tested. Phantom replicas were presented in a probe technique during a materials discrimination experiment. The animal accepted the phantom echoes and classified them in the same manner as it classified real targets.

Dolphin hearing: relative sensitivity as a function of point of application of a contact sound source in the jaw and head region.

The auditory input area of the dolphin head was investigated in an unrestrained animal trained to beach itself and to accept noninvasive electroencephalograph (EEG) electrodes for the recording of the auditory brain-stem response (ABR). The stimulus was a synthetic dolphin click, transmitted from a piezo-electric transducer and coupled to the skin via a small volume of water. The results conform with earlier experiments on acute preparations that show best auditory sensitivity at the middle of the lower jaw. Minimum latency was found at the rear of the lower jaw. A shaded receiver configuration for the dolphin ear is proposed.

Acoustic effects of the ATOC signal (75 Hz, 195 dB) on dolphins and whales.

The Acoustic Thermometry of Ocean Climate (ATOC) program of Scripps Institution of Oceanography and the Applied Physics Laboratory, University of Washington, will broadcast a low-frequency 75-Hz phase modulated acoustic signal over ocean basins in order to study ocean temperatures on a global scale and examine the effects of global warming. One of the major concerns is the possible effect of the ATOC signal on marine life, especially on dolphins and whales. In order to address this issue, the hearing sensitivity of a false killer whale (Pseudorca crassidens) and a Risso's dolphin (Grampus griseus) to the ATOC sound was measured behaviorally. A staircase procedure with the signal levels being changed in 1-dB steps was used to measure the animals' threshold to the actual ATOC coded signal. The results indicate that small odontocetes such as the Pseudorca and Grampus swimming directly above the ATOC source will not hear the signal unless they dive to a depth of approximately 400 m. A sound propagation analysis suggests that the sound-pressure level at ranges greater than 0.5 km will be less than 130 dB for depths down to about 500 m. Several species of baleen whales produce sounds much greater than 170-180 dB. With the ATOC source on the axis of the deep sound channel (greater than 800 m), the ATOC signal will probably have minimal physical and physiological effects on cetaceans.

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.

Object representation in the bottlenose dolphin (Tursiops truncatus): integration of visual and echoic information.

A dolphin performed a 3-alternative matching-to-sample task in different modality conditions (visual/echoic, both vision and echolocation: visual, vision only; echoic, echolocation only). In Experiment 1, training occurred in the dual-modality (visual/echoic) condition. Choice accuracy in tests of all conditions was above chance without further training. In Experiment 2, unfamiliar objects with complementary similarity relations in vision and echolocation were presented in single-modality conditions until accuracy was about 70%. When tested in the visual/echoic condition, accuracy immediately rose (95%), suggesting integration across modalities. In Experiment 3, conditions varied between presentation of sample and alternatives. The dolphin successfully matched familiar objects in the cross-modal conditions. These data suggest that the dolphin has an object-based representational system.

Recognition of aspect-dependent three-dimensional objects by an echolocating Atlantic bottlenose dolphin.

We examined the ability of a bottlenose dolphin (Tursiops truncatus) to recognize aspect-dependent objects using echolocation. An aspect-dependent object such as a cube produces acoustically different echoes at different angles relative to the echolocation signal. The dolphin recognized the objects even though the objects were free to rotate and sway. A linear discriminant analysis and nearest centroid classifier could classify the objects using average amplitude, center frequency, and bandwidth of object echoes. The results show that dolphins can use varying acoustic properties to recognize constant objects and suggest that aspect-independent representations may be formed by combining information gleaned from multiple echoes.

Neural network modeling of a dolphin's sonar discrimination capabilities.

The capability of an echolocating dolphin to discriminate differences in the wall thickness of cylinders was previously modeled by a counterpropagation neural network using only spectral information from the echoes. In this study, both time and frequency information were used to model the dolphin discrimination capabilities. Echoes from the same cylinders were digitized using a broadband simulated dolphin sonar signal with the transducer mounted on the dolphin's pen. The echoes were filtered by a bank of continuous constant-Q digital filters and the energy from each filter was computed in time increments of 1/bandwidth. Echo features of the standard and each comparison target were analyzed in pairs by a counterpropagation neural network, a backpropagation neural network, and a model using Euclidean distance measures. The backpropagation network performed better than both the counterpropagation network, and the Euclidean model, using either spectral-only features or combined temporal and spectral features. All models performed better using features containing both temporal and spectral information. The backpropagation network was able to perform better than the dolphins for noise-free echoes with Q values as low as 2 and 3. For a Q of 2, only temporal information was available. However, with noisy data, the network required a Q of 8 in order to perform as well as the dolphin.

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)

Matching-to-sample by an echolocating dolphin (Tursiops truncatus).

An adult male dolphin was trained to perform a three-alternative delayed matching-to-sample task while wearing eyecups to occlude its vision. Sample and comparison stimuli consisted of a small and a large PVC plastic tube, a water-filled stainless steel sphere, and a solid aluminum cone. Stimuli were presented under water and the dolphin was allowed to identify the stimuli through echolocation. The echolocation clicks emitted by the dolphin to each sample and each comparison stimulus were recorded and analyzed. Over 48 sessions of testing, choice accuracy averaged 94.5% correct. This high level of accuracy was apparently achieved by varying the number of echolocation clicks emitted to various stimuli. Performance appeared to reflect a preexperimental stereotyped search pattern that dictated the order in which comparison items were examined and a complex sequential-sampling decision process. A model for the dolphin's decision-making processes is described.

The critical interval in dolphin echolocation: what is it?

A backward masking function relating target detection to masker delay was generated for a bottlenosed dolphin in an active echolocation target detection task. The masker was triggered by each outgoing echolocation click and could be temporally adjusted from coincidence with the target echo to delays of 700 microseconds. The animal reported target condition using a go/no-go response procedure. A modified method of constants was used to present the four masking delay intervals. Results indicated that 700- and 500-microseconds delays had little effect on target detection. However, as the delay was reduced to 100 microseconds, detection dropped to chance performance. The calculated 70% detection threshold corresponded to a delay of 265 microseconds. The results are discussed in support of the view that time separation pitch (TSP) may be an analytic mechanism used by the dolphin to discern various within-echo target attributes rather than an analytic mechanism for determining target range.

Necrotic stomatitis in a dolphin.

Necrotic stomatitis of undetermined etiology was found in an Atlantic bottlenosed dolphin, Tursiops truncatus. The case history, treatment and hematologic findings are described. The animal remained anorectic throughout the course of the illness, and was force fed on a daily basis for approximately 5 weeks.