Echolocating Big Brown Bats, Eptesicus fuscus, Modulate Pulse Intervals to Overcome Range Ambiguity in Cluttered Surroundings.

Big brown bats (Eptesicus fuscus) emit trains of brief, wideband frequency-modulated (FM) echolocation sounds and use echoes of these sounds to orient, find insects, and guide flight through vegetation. They are observed to emit sounds that alternate between short and long inter-pulse intervals (IPIs), forming sonar sound groups. The occurrence of these strobe groups has been linked to flight in cluttered acoustic environments, but how exactly bats use sonar sound groups to orient and navigate is still a mystery. Here, the production of sound groups during clutter navigation was examined. Controlled flight experiments were conducted where the proximity of the nearest obstacles was systematically decreased while the extended scene was kept constant. Four bats flew along a corridor of varying widths (100, 70, and 40 cm) bounded by rows of vertically hanging plastic chains while in-flight echolocation calls were recorded. Bats shortened their IPIs for more rapid spatial sampling and also grouped their sounds more tightly when flying in narrower corridors. Bats emitted echolocation calls with progressively shorter IPIs over the course of a flight, and began their flights by emitting shorter starting IPI calls when clutter was denser. The percentage of sound groups containing 3 or more calls increased with increasing clutter proximity. Moreover, IPI sequences having internal structure become more pronounced when corridor width narrows. A novel metric for analyzing the temporal organization of sound sequences was developed, and the results indicate that the time interval between echolocation calls depends heavily on the preceding time interval. The occurrence of specific IPI patterns were dependent upon clutter, which suggests that sonar sound grouping may be an adaptive strategy for coping with pulse-echo ambiguity in cluttered surroundings.

Effective biosonar echo-to-clutter rejection ratio in a complex dynamic scene.

Biosonar guidance in a rapidly changing complex scene was examined by flying big brown bats (Eptesicus fuscus) through a Y-shaped maze composed of rows of strongly reflective vertical plastic chains that presented the bat with left and right corridors for passage. Corridors were 80-100 cm wide and 2-4 m long. Using the two-choice Y-shaped paradigm to compensate for left-right bias and spatial memory, a moveable, weakly reflective thin-net barrier randomly blocked the left or right corridor, interspersed with no-barrier trials. Flight path and beam aim were tracked using an array of 24 microphones surrounding the flight room. Each bat flew on a path centered in the entry corridor (base of Y) and then turned into the left or right passage, to land on the far wall or to turn abruptly, reacting to avoid a collision. Broadcasts were broadly beamed in the direction of flight, smoothly leading into an upcoming turn. Duration of broadcasts decreased slowly from 3 to 2 ms during flights to track the chains' progressively closer ranges. Broadcast features and flight velocity changed abruptly about 1 m from the barrier, indicating that echoes from the net were perceived even though they were 18-35 dB weaker than overlapping echoes from surrounding chains.

Support for the beam focusing hypothesis in the false killer whale.

The odontocete sound production system is complex and composed of tissues, air sacs and a fatty melon. Previous studies suggested that the emitted sonar beam might be actively focused, narrowing depending on target distance. In this study, we further tested this beam focusing hypothesis in a false killer whale. Using three linear arrays of hydrophones, we recorded the same emitted click at 2, 4 and 7 m distance and calculated the beamwidth, intensity, center frequency and bandwidth as recorded on each array at every distance. If the whale did not focus her beam, acoustics predicts the intensity would decay with range as a function of spherical spreading and the angular beamwidth would remain constant. On the contrary, our results show that as the distance from the whale to the array increases, the beamwidth is narrower and the received click intensity is higher than that predicted by a spherical spreading function. Each of these measurements is consistent with the animal focusing her beam on a target at a given range. These results support the hypothesis that the false killer whale is 'focusing' its sonar beam, producing a narrower and more intense signal than that predicted by spherical spreading.

Mouth gape angle has little effect on the transmitted signals of big brown bats (Eptesicus fuscus).

Bats perform high-resolution echolocation by comparing temporal and spectral features of their transmitted pulses to the received echoes. In complex environments with moving prey, dynamically adapting the transmitted pulses can increase the probability of successful target representation and interception. This study further investigates the adaptive vocal-motor strategies of big brown bats (Eptesicus fuscus). During stationary target detection experiments, echolocation sounds were simultaneously recorded with high-speed, infrared video to examine the relationship of mouth position and movement to pulse characteristics among bats. All three bats produced strobe groups, but the proportion and frequency characteristics of the strobe group pulses differed for individual bats. Additionally, mouth gape angle had little effect on the emitted pulse characteristics, which suggests that laryngeal mechanisms drive changes in emitted pulses.

High resolution acoustic measurement system and beam pattern reconstruction method for bat echolocation emissions.

Measurements of the transmit beam patterns emitted by echolocating bats have previously been limited to cross-sectional planes or averaged over multiple signals using sparse microphone arrays. To date, no high-resolution measurements of individual bat transmit beams have been reported in the literature. Recent studies indicate that bats may change the time-frequency structure of their calls depending on the task, and suggest that their beam patterns are more dynamic than previously thought. To investigate beam pattern dynamics in a variety of bat species, a high-density reconfigurable microphone array was designed and constructed using low-cost ultrasonic microphones and custom electronic circuitry. The planar array is 1.83 m wide by 1.42 m tall with microphones positioned on a 2.54 cm square grid. The system can capture up to 228 channels simultaneously at a 500 kHz sampling rate. Beam patterns are reconstructed in azimuth, elevation, and frequency for visualization and further analysis. Validation of the array measurement system and post-processing functions is shown by reconstructing the beam pattern of a transducer with a fixed circular aperture and comparing the result with a theoretical model. To demonstrate the system in use, transmit beam patterns of the big brown bat, Eptesicus fuscus, are shown.

Multi-component separation and analysis of bat echolocation calls.

The vast majority of animal vocalizations contain multiple frequency modulated (FM) components with varying amounts of non-linear modulation and harmonic instability. This is especially true of biosonar sounds where precise time-frequency templates are essential for neural information processing of echoes. Understanding the dynamic waveform design by bats and other echolocating animals may help to improve the efficacy of man-made sonar through biomimetic design. Bats are known to adapt their call structure based on the echolocation task, proximity to nearby objects, and density of acoustic clutter. To interpret the significance of these changes, a method was developed for component separation and analysis of biosonar waveforms. Techniques for imaging in the time-frequency plane are typically limited due to the uncertainty principle and interference cross terms. This problem is addressed by extending the use of the fractional Fourier transform to isolate each non-linear component for separate analysis. Once separated, empirical mode decomposition can be used to further examine each component. The Hilbert transform may then successfully extract detailed time-frequency information from each isolated component. This multi-component analysis method is applied to the sonar signals of four species of bats recorded in-flight by radiotelemetry along with a comparison of other common time-frequency representations.