Hearing sensation levels of emitted biosonar clicks in an echolocating Atlantic bottlenose dolphin

Emitted biosonar clicks and auditory evoked potential (AEP) responses triggered by the clicks were synchronously recorded during echolocation in an Atlantic bottlenose dolphin (Tursiops truncatus) trained to wear suction-cup EEG electrodes and to detect targets by echolocation. Three targets with target strengths of -34, -28, and -22 dB were used at distances of 2 to 6.5 m for each target. The AEP responses were sorted according to the corresponding emitted click source levels in 5-dB bins and averaged within each bin to extract biosonar click-related AEPs from noise. The AEP amplitudes were measured peak-to-peak and plotted as a function of click source levels for each target type, distance, and target-present or target-absent condition. Hearing sensation levels of the biosonar clicks were evaluated by comparing the functions of the biosonar click-related AEP amplitude-versus-click source level to a function of external (in free field) click-related AEP amplitude-versus-click sound pressure level. The results indicated that the dolphin's hearing sensation levels to her own biosonar clicks were equal to that of external clicks with sound pressure levels 16 to 36 dB lower than the biosonar click source levels, varying with target type, distance, and condition. These data may be assumed to indicate that the bottlenose dolphin possesses effective protection mechanisms to isolate the self-produced intense biosonar beam from the animal's ears during echolocation.     

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena)

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I–VI), and 1.4 ms (II–IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal’s size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.     

TARGET DISCRIMINATION BY AN ECHOLOCATING FINLESS PORPOISE, NEOPHOCAENA PHOCAENOIDES

The ability of a finless porpoise (Neophocaena phocaenoides) to discriminate the material and size of a target by echolocation was investigated. The porpoise was required to choose a standard target of a 15-mm diameter solid steel cylinder from two stimuli, a standard and a comparison target, presented simultaneously. The porpoise could distinguish a standard cylinder from acrylic resin and brass targets, but had difficulties distinguishing it from an aluminum cylinder. In size discrimination, the porpoise could distinguish the standard from 12-, 18-, and 20-mm diameter cylinders, but had difficulties distinguishing it from a 14-mm diameter cylinder. Echo measurements suggest that the porpoise is able to detect: (1) time difference between two echo highlights to within approximately 1 μS, (2) frequency shifts of approximately 7 kHz in a broadband echo having a peak frequency of nearly 140 kHz, (3) time-separation pitch of approximately 30 kHz, and (4) target strength differences of approximately 1 dB.     

Gain control in the sonar of odontocetes

The sonar of odontocetes processes echo-signals within a wide range of echo levels. The level of echoes varies widely by tens of decibels depending on the level of the emitted sonar pulse, the target strength, the distance to the target, and the sound absorption by the water media. The auditory system of odontocetes must be capable of effective perception, analysis, and discrimination of echo-signals within all this variability. The sonar of odontocetes has several mechanisms to compensate for the echo-level variation (gain control). To date, several mechanisms of the biosonar gain control have been revealed in odontocetes: (1) adjustment of emitted sonar pulse levels (the longer the distance to the target, the higher the level of the emitted pulse), (2) short-term variation of hearing sensitivity based on forward masking of the echo by the preceding self-heard emitted pulse and subsequent release from the masking, and (3) active long-term control of hearing sensitivity. Recent investigations with the use of the auditory evoked-potential technique have demonstrated that these mechanisms effectively minimize the variation of the response to the echo when either the emitted sonar pulse level, or the target distance, or both vary within a wide range. A short review of these data is presented herein.     

ECHOLOCATION RATES OF TWO HARBOR PORPOISES (PHOCOENA PHOCOENA)

The rate of occurrence of click trains of two harbor porpoises ( Phocoena phocoena ) were counted during 14 nights. We developed an echolocation signal detection system that was harnessed to a porpoise and activated a light when the animal emitted an echolocation signal. This device, referred to as a click-light, detects echolocation signals above 150 dB re 1 μPa in the 28–180 kHz range. Echolocation rates, i.e., occurrences of click trains, changed frequently, ranging from 0 to 25 per minute. Echolocation rates were affected by feeding, individual difference, and enclosure type such as the net enclosure and the pool. The porpoise echolocation rates seemed to show acclimation.     

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow,Acoustically Complex Habitat

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPa_pp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPa_pp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.     

Biosonar behaviour of free-ranging porpoises

Detecting objects in their paths is a fundamental perceptional function of moving organisms. Potential risks and rewards, such as prey, predators, conspecifics or non-biological obstacles, must be detected so that an animal can modify its behaviour accordingly. However, to date few studies have considered how animals in the wild focus their attention. Dolphins and porpoises are known to actively use sonar or echolocation. A newly developed miniature data logger attached to a porpoise allows for individual recording of acoustical search efforts and inspection distance based on echolocation. In this study, we analysed the biosonar behaviour of eight free-ranging finless porpoises (Neophocaena phocaenoides) and demonstrated that these animals inspect the area ahead of them before swimming silently into it. The porpoises inspected distances up to 77 m, whereas their swimming distance without using sonar was less than 20 m. The inspection distance was long enough to ensure a wide safety margin before facing real risks or rewards. Once a potential prey item was detected, porpoises adjusted their inspection distance from the remote target throughout their approach.     

A comprehensive computational model of animal biosonar signal processing

Computational models of animal biosonar seek to identify critical aspects of echo processing responsible for the superior, real-time performance of echolocating bats and dolphins in target tracking and clutter rejection. The Spectrogram Correlation and Transformation (SCAT) model replicates aspects of biosonar imaging in both species by processing wideband biosonar sounds and echoes with auditory mechanisms identified from experiments with bats. The model acquires broadband biosonar broadcasts and echoes, represents them as time-frequency spectrograms using parallel bandpass filters, translates the filtered signals into ten parallel amplitude threshold levels, and then operates on the resulting time-of-occurrence values at each frequency to estimate overall echo range delay. It uses the structure of the echo spectrum by depicting it as a series of local frequency nulls arranged regularly along the frequency axis of the spectrograms after dechirping them relative to the broadcast. Computations take place entirely on the timing of threshold-crossing events for each echo relative to threshold-events for the broadcast. Threshold-crossing times take into account amplitude-latency trading, a physiological feature absent from conventional digital signal processing. Amplitude-latency trading transposes the profile of amplitudes across frequencies into a profile of time-registrations across frequencies. Target shape is extracted from the spacing of the object’s individual acoustic reflecting points, or glints, using the mutual interference pattern of peaks and nulls in the echo spectrum. These are merged with the overall range-delay estimate to produce a delay-based reconstruction of the object’s distance as well as its glints. Clutter echoes indiscriminately activate multiple parts in the null-detecting system, which then produces the equivalent glint-delay spacings in images, thus blurring the overall echo-delay estimates by adding spurious glint delays to the image. Blurring acts as an anticorrelation process that rejects clutter intrusion into perceptions.     

Spatial unmasking in the echolocating Big Brown Bat, <Emphasis Type="Italic">Eptesicus fuscus</Emphasis>

Masking affects the ability of echolocating bats to detect a target in the presence of clutter targets. It can be reduced by spatially separating the targets. Spatial unmasking was measured in a two-alternative-forced-choice detection experiment with four Big Brown Bats detecting a wire at 1 m distance. Depth dependent spatial unmasking was investigated by the bats detecting a wire with a diameter of 1.2 mm in front of a masker with a threshold distance of 11 cm behind the wire. For angle dependent spatial unmasking the masker was turned laterally, starting from its threshold position at 11 cm. With increasing masker angles the bats could detect thinner wires with diameters decreasing from 1.2 mm (target strength −36.8 dB) at 0° to 0.2 mm (target strength −63.0 dB) at 22°. Without masker, the bats detected wire diameters of 0.16 mm (target strength −66.2 dB), reached with masker positions beyond 23° (complete masking release). Analysis of the sonar signals indicated strategies in the echolocation behavior. The bats enhanced the second harmonics of their signals. This may improve the spatial separation between wire and masker due to frequency-dependent directionality increase of sound emission and echo reception.     

Synthetic communication signals influence wild harbour porpoise (Phocoena phocoena) behaviour

We used our novel and programmable Porpoise Alarm (PAL, patd.) to synthesize life-like, electronic harbour porpoise communication signals based on those described for captive animals. In the Little Belt, Denmark, we employed PAL (source level 158 ± 1 dB p–p re 1 μPa@1 m; centroid frequency 133 ± 8.5 kHz) to synthesize three aggressive click train types termed “A”, “F3” and “M1” to naive, free-living harbour porpoises. Via theodolite tracking (372 h of total visual effort spread over 10 expeditions) we found that, depending on signal type, porpoises either avoid or become attracted to PAL: Signal types “A” and “F3” are slight deterrents, porpoises increasing minimum range (+23 to 32 m, respectively), whereas “M1” attracts porpoises, reducing range (by ? 29 m). As determined via archival acoustic detectors (AADs), both signals “F3” and “M1” led the animals to significantly intensify their click rate (by +10% and 68%, respectively) while signal “A” led to a significant reduction ( ? 59%). We propose that equipping fishing gear with PAL emitting signal “F3” could potentially reduce porpoise by-catch by increasing (1) awareness through enhanced echolocation and (2) distance to the nets. Detection probability and radius of PAL/AAD tandems could be improved by emitting signal “M1” to focus porpoise echolocation signals on the AAD. The signal may also be useful in luring animals away from hazards, which may be helpful for conservation measures prior to the onset of harmful acoustic activities such as pile-driving, seismic exploration or ammunition clearance.     

The Diel Rhythms of Biosonar Behavior in the Yangtze Finless Porpoise (Neophocaena asiaeorientalis asiaeorientalis) in the Port of the Yangtze River: The Correlation between Prey Availability and Boat Traffic

Information on the habitat use of the critically endangered Yangtze finless porpoise (Neophocaena asiaeorientalis asiaeorientalis) is critical for its conservation. The diel biosonar behavior of the porpoise in the port areas of the Yangtze River was examined along with simultaneous observations of fish density and boat traffic. Biosonar pulses from the porpoises were detected for 1233 min (5.77%) over a 21,380 min duration of effective observations. In total, 190 (5.63%) buzzes (an indication of prey capture attempts) were recorded among the 3372 identified click trains. Of the 168 echolocation encounters (bouts of click trains less than eight min apart), 150 (89.3%) involved single animals, indicating that solitary porpoises were frequently present and feeding in the port areas. Significant diel patterns were evident involving the biosonar behavior of the porpoises (including click trains and buzzes), fish density and boat traffic. The frequencies of the click trains and buzzes were significantly lower during the day than in the evening and at night, which suggests that porpoises in this region are primarily engaged in crepuscular and nocturnal foraging. The lack of a significant diel pattern in the echolocation encounters indicates the importance of the port in porpoise conservation. A forced feeding schedule may be associated with the lack of a significant correlation between porpoise acoustics and boat traffic. Overall, prey availability appears to be the primary factor that attracts porpoises. Additionally, porpoises tend to migrate or remain downstream in the morning and migrate or remain upstream in the evening, most likely to follow their prey. The findings of this study can be used to improve the conservation of the Yangtze finless porpoise.     

Echolocation constraints of Daubenton's Bat foraging over water

1. Daubenton's Bats ( Myotis daubentonii ) foraging over a stream concentrated their activity over calm surfaces, avoiding an adjacent area with small ripples (< 3 cm high). Aerial insects were most abundant over the ripples, so insect distribution could not explain why the bats avoided this area.
2. The bats flew low over water and always ( N = 22) directed the head forwards, presumably emitting the echolocation beam parallel to the surface, thus minimizing clutter. At an angle of incidence of 30° there was significantly more clutter from the rippled water.
3. The ripples produced ultrasonic noises in the form of transient pulses at an average rate of 6·2 per second. In the present case, such pulses were common enough potentially to interfere with target detection by the bats. Transient noises and echo clutter from moving ripples may be the principal reason why bats generally avoid foraging low over turbulent water.
4. The target strength of a potential insect prey at the water surface and the source levels of the bats' searching signals were measured to use in estimating the echo level at the bat when it detects the prey. The echo level at detection (+ 38 dB sound pressure level) was about the same as the clutter level extrapolated to the detection distance. This suggests that Daubenton's Bat operates at very low signal-to-noise ratios when foraging for insects near the water surface.     

The degradation of distance discrimination in big brown bats (Eptesicus fuscus) caused by different interference signals

The ability of two big brown bats (Eptesicus fuscus) to discriminate the distance to an electronically synthesized phantom target by echolocation was tested in the presence of interfering signals presented slightly before the target echo. Interfering signals were chosen to have differing degrees of similarity to the typical echolocation emission used by the bat in this task (which was the signal used to create the phantom target), and we predicted that the degree of disruption of ranging would be proportional to the similarity of the interference to the target echo. This prediction was not confirmed; rather, all interference signals not identical to the target echo increased the threshold to about twice that found with no interference. When the interference was identical to the target echo, the threshold increased to about 4 times that with no interference. When each bat was presented with phantom target echoes appropriate for the other bat, its range discrimination threshold increased about ten fold, and in this case the degree of interference of different signals was related to their similarity to the target echo, not to their similarity to the bat's normal signal. We suggest that Eptesicus may suppress interference by a more sophisticated strategy than simple linear matched filtering.Abbreviations E exemplar signal - M _f foreign model signal - M _r reversed self-model signal - M _s self-model signal - N noise signal - SPL sound pressure level     

PORPOISE CLICKS FROM A SPERM WHALE NOSE—CONVERGENT EVOLUTION OF 130 KHZ PULSES IN TOOTHED WHALE SONARS?

ABSTRACT

Small toothed whales of the family Phocoenidae and delphinid genus Cephalorhynchus produce long-duration, narrowband biosonar clicks above 100 kHz, that might be seen as an adaptation for short range echolocation in shallow water. This paper presents data showing that the distantly related, and larger pygmy sperm whale Kogia breviceps (Kogiidae), that is a deep-diving, cephalopod-eating toothed whale, produce narrow-banded high frequency (NBHF) clicks identical to those of Phocoena and Cephalorhynchus (f₀ = 130 kHz, Q_3dB>10, duration > 80 msec). Thus, NBHF biosonar signals have evolved on three independent occasions in the odontocete suborder, but the apparent functional convergence does not relate to anatomical or niche similarity. Rather, it is suggested that a biosonar strategy adapting to a minimum in ocean ambient noise above 100 kHz in concert with high Q auditory filters have led to convergent evolution of the NBHF biosonar clicks. Since these biosonar signals carry all their energy at frequencies above the upper hearing limit of the killer whale Orcinus orca, predator avoidance may also have been a evolutionary shaping factor of the sonar signals from these non-whistling odontocetes.     

Echo SPL influences the ranging performance of the big brown bat,Eptesicus fuscus

Four bats of the species Eptesicus fuscus were trained in a two-alternative forced-choice procedure to discriminate between two phantom targets that differed in range. The rewarded stimulus was located at a distance of 52.7 cm, while the other unrewarded stimulus was further away. Only one target was presented at a time.In the first experiment we measured the range discrimination performance at an echo SPL of –28 dB relative to the bat's sonar transmission. A 75% correct performance level was arbitrarily defined as threshold and was obtained at a delay difference of 80 s, corresponding to a range difference of 13.8 mm.In the second experiment the delay difference was fixed at 150 s and the echo SPL varied between –8 and –48 dB relative to sonar emissions. The performance of the bats depended on the relative echo SPL. At –28 dB the bats showed the best performance. It deteriorated at an increase of the relative echo SPL to –18 dB and –8 dB. The performance also deteriorated when the relative echo SPL was reduced to –38 dB and –48 dB. Only at low relative echo SPLs did the bats partially compensate for the reduction in echo SPL and increased the SPL of their emitted signals by a few dB.Our results support the hypothesis that neurons exhibiting paradoxical latency shift may be involved in encoding target range. This hypothesis predicts a decrease in performance at high echo SPLs as we found it in our experiments. The observed reduction in performance at very low echo SPLs may be due to a decrease in S/N ratio.     

Click-based echolocation in bats: not so primitive after all

Echolocating bats of the genus Rousettus produce click sonar signals, using their tongue (lingual echolocation). These signals are often considered rudimentary and are believed to enable only crude performance. However, the main argument supporting this belief, namely the click’s reported long duration, was recently shown to be an artifact. In fact, the sonar clicks of Rousettus bats are extremely short, ~50–100 μs, similar to dolphin vocalizations. Here, we present a comparison between the sonar systems of the ‘model species’ of laryngeal echolocation, the big brown bat (Eptesicus fuscus), and that of lingual echolocation, the Egyptian fruit bat (Rousettus aegyptiacus). We show experimentally that in tasks, such as accurate landing or detection of medium-sized objects, click-based echolocation enables performance similar to laryngeal echolocators. Further, we describe a sophisticated behavioral strategy for biosonar beam steering in clicking bats. Finally, theoretical analyses of the signal design—focusing on their autocorrelations and wideband ambiguity functions—predict that in some aspects, such as target ranging and Doppler-tolerance, click-based echolocation might outperform laryngeal echolocation. Therefore, we suggest that click-based echolocation in bats should be regarded as a viable echolocation strategy, which is in fact similar to the biosonar used by most echolocating animals, including whales and dolphins.     

Auditory temporal resolution and evoked responses to pulsed sounds for the Yangtze finless porpoises (<Emphasis Type="Italic">Neophocaena phocaenoides asiaeorientalis</Emphasis>)

Temporal cues are important for some forms of auditory processing, such as echolocation. Among odontocetes (toothed whales, dolphins, and porpoises), it has been suggested that porpoises may have temporal processing abilities which differ from other odontocetes because of their relatively narrow auditory filters and longer duration echolocation signals. This study examined auditory temporal resolution in two Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis) using auditory evoked potentials (AEPs) to measure: (a) rate following responses and modulation rate transfer function for 100 kHz centered pulse sounds and (b) hearing thresholds and response amplitudes generated by individual pulses of different durations. The animals followed pulses well at modulation rates up to 1,250 Hz, after which response amplitudes declined until extinguished beyond 2,500 Hz. The subjects had significantly better hearing thresholds for longer, narrower-band pulses similar to porpoise echolocation signals compared to brief, broadband sounds resembling dolphin clicks. Results indicate that the Yangtze finless porpoise follows individual acoustic signals at rates similar to other odontocetes tested. Relatively good sensitivity for longer duration, narrow-band signals suggests that finless porpoise hearing is well suited to detect their unique echolocation signals.     

Highly Directional Sonar Beam of Narwhals (Monodon monoceros) Measured with a Vertical 16 Hydrophone Array

Recordings of narwhal (Monodon monoceros) echolocation signals were made using a linear 16 hydrophone array in the pack ice of Baffin Bay, West Greenland in 2013 at eleven sites. An average -3 dB beam width of 5.0° makes the narwhal click the most directional biosonar signal reported for any species to date. The beam shows a dorsal-ventral asymmetry with a narrower beam above the beam axis. This may be an evolutionary advantage for toothed whales to reduce echoes from the water surface or sea ice surface. Source level measurements show narwhal click intensities of up to 222 dB pp re 1 μPa, with a mean apparent source level of 215 dB pp re 1 μPa. During ascents and descents the narwhals perform scanning in the vertical plane with their sonar beam. This study provides valuable information for reference sonar parameters of narwhals and for the use of acoustic monitoring in the Arctic.     

Labile cochlear tuning in the mustached bat

Summary Acoustic stimuli near 60 kHz elicit pronounced resonance in the cochlea of the mustached bat (Pteronotus parnellii parnellii). The cochlear resonance frequency (CRF) is near the second harmonic, constant frequency (CF2) component of the bat's biosonar signals. Within narrow bands where CF2 and third harmonic (CF3) echoes are maintained, the cochlea has sharp tuning characteristics that are conserved throughout the central auditory system. The purpose of this study was to examine the effects of temperature-related shifts in the CRF on the tuning properties of neurons in the cochlear nucleus and inferior colliculus.Eighty-two single and multi-unit recordings were characterizedin 6 awake bats with chronically implanted cochlear microphonic electrodes. As the CRF changed with body temperature, the tuning curves of neurons sharply tuned to frequencies near the CF2 and CF3 shifted with the CRF in every case, yielding a change in the unit's best frequency. The results show that cochlear tuning is labile in the mustached bat, and that this lability produces tonotopic shifts in the frequency response of central auditory neurons. Furthermore, results provide evidence of shifts in the frequency-to-place code within the sharply tuned CF2 and CF3 regions of the cochlea. In conjunction with the finding that biosonar emission frequency and the CRF shift concomitantly with temperature and flight, it is concluded that the adjustment of biosonar signals accommodates the shifts in cochlear and neural tuning that occur with active echolocation.Abbreviations BF best frequency - CF characteristic frequency - CF2, CF3 second and third harmonic, constant frequency components of the biosonar signal - CM cochlear microphonic - CN cochlear nucleus - CRF cochlear resonance frequency - IC inferior colliculus - MT minimum threshold - OAE otoacoustic emission - Q_10dB BF (or CF) divided by the response bandwidth at 10 dB above MT     

CHANGES IN CLICK SOURCE LEVELS WITH DISTANCE TO TARGETS: STUDIES OF FREE-RANGING WHITE-BEAKED DOLPHINS LAGENORHYNCHUS ALBIROSTRIS AND CAPTIVE HARBOUR PORPOISES PHOCOENA PHOCOENA

ABSTRACT

Probably all odontocetes use echolocation for spatial orientation and detection of prey. We used a four hydrophone “Y” array to record the high frequency clicks from free-ranging White-beaked Dolphins Lagenorhynchus albirostris and captive Harbour Porpoises Phocoena phocoena. From the recordings we calculated distances to the animals and source levels of the clicks. The recordings from White-beaked Dolphins were made in Iceland and those from Harbour Porpoises at Fjord & Baelt, Kerteminde, Denmark during prey capture. We used stringent criteria to determine which clicks could be defined as being on the acoustic axis. Two dolphin and nine porpoise click series could be used to track individual animals, which presumably focused on the array hydrophones or a fish right in front of the array. The apparent source levels of clicks in the individual tracks increased with range. One individual White-beaked Dolphin and three Harbour Porpoises regulate their output signal level to nearly compensate for one-way transmission loss while approaching a target. The other dolphin regulated the output differently. For most of the recordings the sound level at the target remains nearly constant and the echo level at the animal increases as it closes on the target.