Detection of targets colocalized in clutter by big brown bats (Eptesicus fuscus)

Echolocating big brown bats (Eptesicus fuscus) frequently catch insects during aerial pursuits in open spaces, but they also capture prey swarming on vegetation, and from substrates. To evaluate perception of targets on cluttered surfaces, big brown bats were trained in a two-alternative forced-choice task to locate a target, varying in height, that was embedded partway in holes (clutter) cut in a foam surface. The holes were colocalized with the possible positions of the target at distances ranging from 25 to 35 cm. For successful perception of the target, the bat had to detect the echoes contributed by the target in the same time window that contained echoes from the clutter. Performance was assessed in terms of target reflective strength relative to clutter strength in the same time window. The bats detected the target whenever the target strength was greater than 1-2 dB above the clutter.     

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.     

Evidence for spatial representation of object shape by echolocating bats (Eptesicus fuscus)

Big brown bats were trained in a two-choice task to locate a two-cylinder dipole object with a constant 5 cm spacing in the presence of either a one-cylinder monopole or another two-cylinder dipole with a shorter spacing. For the dipole versus monopole task, the objects were either stationary or in motion during each trial. The dipole and monopole objects varied from trial to trial in the left-right position while also roving in range (10-40 cm), cross range separation (15-40 cm), and dipole aspect angle (0 degrees -90 degrees ). These manipulations prevented any single feature of the acoustic stimuli from being a stable indicator of which object was the correct choice. After accounting for effects of masking between echoes from pairs of cylinders at similar distances, the bats discriminated the 5 cm dipole from both the monopole and dipole alternatives with performance independent of aspect angle, implying a distal, spatial object representation rather than a proximal, acoustic object representation.     

Transformation of external-ear spectral cues into perceived delays by the big brown bat,Eptesicus fuscus

The external-ear transfer function for big brown bats (Eptesicus fuscus) contains two prominent notches that vary from 30 to 55 kHz and from 70 to 100 kHz, respectively, as sound-source elevation moves from -40 to +10 degrees. These notches resemble a higher-frequency version of external-ear cues for vertical localization in humans and other mammals. However, they also resemble interference notches created in echoes when reflected sounds overlap at short time separations of 30-50 micros. Psychophysical experiments have shown that bats actually perceive small time separations from interference notches, and here we used the same technique to test whether external-ear notches are recognized as a corresponding time separation, too. The bats' performance reveals the elevation dependence of a time-separation estimate at 25-45 micros in perceived delay. Convergence of target-shape and external-ear cues onto echo spectra creates ambiguity about whether a particular notch relates to the object or to its location, which the bat could resolve by ignoring the presence of notches at external-ear frequencies. Instead, the bat registers the frequencies of notches caused by the external ear along with notches caused by the target's structure and employs spectrogram correlation and transformation (SCAT) to convert them all into a family of delay estimates that includes elevation.     

Spectral cues and perception of the vertical position of targets by the big brown bat, Eptesicus fuscus

Big brown bats (Eptesicus fuscus) were trained to discriminate between vertical angles subtended by paired beads suspended from fishing line. Bats were rewarded for choosing the smaller of the two angles presented. The difference between the angles was changed systematically using a transformed up-down procedure and the bats' ability to detect the difference was measured at different vertical locations. When the beads were centered at +20 degrees (above the horizon), at 0 degree (the horizon), and at -20 degrees (below the horizon), vertical angle acuity (VAA) was maintained between 2.9 degrees and 4.1 degrees. At more extreme vertical positions both bats showed loss of acuity; when the beads were centered around -40 degrees, VAA was 6.7 degrees or 8.3 degrees and at +40, VAA was worse than 21 degrees (the largest difference tested). When the tragi of both ears were bent down and glued to the side of the face, bats showed severe loss of acuity for beads centered at -20 degrees (VAA 18.3 degrees and 20.1 degrees), but maintained their angle acuity for beads centered at +20 degrees (VAA 3.8 degrees and 4.9 degrees). The results are consistent with the spectral cues created by the filtering of the external ear.     

Stabilization of perceived echo amplitudes in echolocating bats. II. The acoustic behavior of the big brown bat, Eptesicus fuscus, when tracking moving prey.

Big brown bats, Eptesicus fuscus, can be trained to use echolocation to track a small microphone with a food reward attached when it is moved rapidly toward them. This situation mimics prey interception in the wild while allowing very precise recording of the sonar pulses emitted during tracking behavior. The results show that E. fuscus intensity compensates, reducing emitted intensity by 6 dB per halving of target range so that the intensity incident upon the target is constant and echo intensity increases by 6 dB per halving of range. This increase in echo intensity is effectively canceled by the reduction in auditory sensitivity due to automatic gain control (AGC) of 6 to 7 dB per halving of range. Intensity compensation behavior and AGC therefore form a dual-component, symmetrical system that stabilizes perceived echo amplitudes during target approach. The same system is present in the fishing bat, Noctilio leporinus, suggesting that it may be widespread in echolocating bats. Correlation analysis shows that, despite large changes in the duration of the pulses emitted by E. fuscus during an approach, the pulse frequency structure is such that the spatial image of the target perceived along the range axis is highly stable. Pulse duration is not reduced in the manner theoretically necessary to eliminate potential echo distortion effects due to AGC, but is reduced in such a way that this distortion is insignificant. During the terminal buzz, a high degree of temporal overlap (relative to pulse duration) occurs between emitted pulse and returning echo.     

Flying big brown bats emit a beam with two lobes in the vertical plane

The sonar beam of an echolocating bat forms a spatial window restricting the echo information returned from the environment. Investigating the shape and orientation of the sonar beam produced by a bat as it flies and performs various behavioral tasks may yield insight into the operation of its sonar system. This paper presents recordings of vertical and horizontal cross sections of the sonar beam produced by Eptesicus fuscus (big brown bats) as they fly and pursue prey in a laboratory flight room. In the horizontal plane the sonar beam consists of one large lobe and in the vertical plane the beam consists of two lobes of comparable size oriented frontally and ventrally. In level flight, the bat directs its beam such that the ventral lobe is pointed forward and down toward the ground ahead of its flight path. The bat may utilize the downward directed lobe to measure altitude without the need for vertical head movements.     

Stabilization of perceived echo amplitudes in echolocating bats. I. Echo detection and automatic gain control in the big brown bat, Eptesicus fuscus, and the fishing bat, Noctilio leporinus.

Previous research on echo detection in bats has suggested that the effective threshold is a function of the acoustic clutter in the experimental environment, as might be expected given the low ambient noise levels typical of such psychophysical research. This paper demonstrates that theory of signal detectability (TSD) methodology is applicable to bats and uses it to show that an important element of clutter limiting in Eptesicus fuscus and Noctilio leporinus is backward masking of phantom targets by the real echo from the loudspeakers used to generate them. This information suggests that a previous estimate of the magnitude of automatic gain control (AGC) is too high, due to variable backward masking inherent in the experimental method used. A re-examination of gain control using a masking-free method shows that it reduces auditory sensitivity by 6 to 7 dB per halving of target range, rather than 11 dB as previously thought.     

Accuracy of distance measurement in the bat Eptesicus fuscus: theoretical aspects and computer simulations

Behavioral experiments of Simmons [J. Acoust. Soc. Am. 54, 157-173 (1973) and Science 204, 1336-1338 (1979)] on the ranging accuracy in the bat Eptesicus fuscus have led to far-reaching postulates on the existence of optimal and phase-conserving processing mechanisms in the bat. In this paper, the results of computer simulations of these experiments are presented. Two receiver types are investigated: the fully coherent cross-correlation receiver and the cross-correlation receiver with envelope processing (semicoherent). It is shown that Simmons' experiments cannot be treated as a simple estimation of distance, but require at least two (range difference experiment; see Simmons, 1973) or four (range jitter experiment; see Simmons, 1979) echolocation sounds for one decision. The performance of the bat in both experiments is much worse than predicted for a coherent and a semicoherent receiver type. The bat's accuracy in Simmons' range difference experiment is at least 18 dB worse than predicted for an optimal receiver. The results of the jitter experiment cannot be interpreted in a simple way as proof that bats are able to evaluate phase information as in a fully coherent cross-correlation receiver.     

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.     

Range resolution and the possible use of spectral information in the echolocating bat, Eptesicus fuscus

Individuals of the echolocating bat Eptesicus fuscus were trained to discriminate simulated two-wave-front targets with internal time delays of 0 to 100 microns between the wave fronts from a one-wave-front target. The ability of bats to discriminate between such targets can be referred to as range resolution. In Eptesicus fuscus, this ability is limited to distinct internal time delays (12, 32-40, and 52-100 microns) between the two wave fronts of a double-wave-front target. Analysis of the simulated two-wave-front echoes reveals periodic frequency minima in the spectrum. Position and separation of these spectral minima depend on the time delay between the two wave fronts. The occurrence of spectral minima within the frequency range of the first harmonic in the echo of the bats' echolocation call correlates to the bats' ability to discriminate a one-wave-front echo from two-wave-front echoes, suggesting that Eptesicus fuscus uses spectral differences within the first harmonic in echoes for range resolution.     

Target representation of naturalistic echolocation sequences in single unit responses from the inferior colliculus of big brown bats

Echolocating big brown bats (Eptesicus fuscus) emit trains of frequency-modulated (FM) biosonar signals whose duration, repetition rate, and sweep structure change systematically during interception of prey. When stimulated with a 2.5-s sequence of 54 FM pulse-echo pairs that mimic sounds received during search, approach, and terminal stages of pursuit, single neurons (N = 116) in the bat's inferior colliculus (IC) register the occurrence of a pulse or echo with an average of < 1 spike/sound. Individual IC neurons typically respond to only a segment of the search or approach stage of pursuit, with fewer neurons persisting to respond in the terminal stage. Composite peristimulus-time-histogram plots of responses assembled across the whole recorded population of IC neurons depict the delay of echoes and, hence, the existence and distance of the simulated biosonar target, entirely as on-response latencies distributed across time. Correlated changes in pulse duration, repetition rate, and pulse or echo amplitude do modulate the strength of responses (probability of the single spike actually occurring for each sound), but registration of the target itself remains confined exclusively to the latencies of single spikes across cells. Modeling of echo processing in FM biosonar should emphasize spike-time algorithms to explain the content of biosonar images.     

Asymmetry and dynamics of a narrow sonar beam in an echolocating harbor porpoise

A key component in the operation of a biosonar system is the radiation of sound energy from the sound producing head structures of toothed whales and microbats. The current view involves a fixed transmission aperture by which the beam width can only change via changes in the frequency of radiated clicks. To test that for a porpoise, echolocation clicks were recorded with high angular resolution using a 16 hydrophone array. The beam is narrower than previously reported (DI = 24 dB) and slightly dorso-ventrally compressed (horizontal -3 dB beam width: 13°, vertical -3 dB beam width: 11°). The narrow beam indicates that all smaller toothed whales investigated so far have surprisingly similar beam widths across taxa and habitats. Obtaining high directionality may thus be at least in part an evolutionary factor that led to high centroid frequencies in a group of smaller toothed whales emitting narrow band high frequency clicks. Despite the production of stereotyped narrow band high frequency clicks, changes in the directionality by a few degrees were observed, showing that porpoises can obtain changes in sound radiation.     

Echolocation call intensity and directionality in flying short-tailed fruit bats, Carollia perspicillata (Phyllostomidae)

The directionality of bat echolocation calls defines the width of bats' sonar "view," while call intensity directly influences detection range since adequate sound energy must impinge upon objects to return audible echoes. Both are thus crucial parameters for understanding biosonar signal design. Phyllostomid bats have been classified as low intensity or "whispering bats," but recent data indicate that this designation may be inaccurate. Echolocation beam directionality in phyllostomids has only been measured through electrode brain-stimulation of restrained bats, presumably excluding active beam control via the noseleaf. Here, a 12-microphone array was used to measure echolocation call intensity and beam directionality in the frugivorous phyllostomid, Carollia perspicillata, echolocating in flight. The results showed a considerably narrower beam shape (half-amplitude beam angles of approximately 16° horizontally and 14° vertically) and louder echolocation calls [source levels averaging 99 dB sound pressure level (SPL) root mean square] for C. perspicillata than was found for this species when stationary. This suggests that naturally behaving phyllostomids shape their sound beam to achieve a longer and narrower sonar range than previously thought. C. perspicillata orient and forage in the forest interior and the narrow beam might be adaptive in clutter, by reducing the number and intensity of off-axis echoes.     

Echo-intensity compensation in echolocating bats (Pipistrellus abramus) during flight measured by a telemetry microphone

An onboard microphone (Telemike) was developed to examine changes in the basic characteristics of echolocation sounds of small frequency-modulated echolocating bats, Pipistrellus abramus. Using a dual high-speed video camera system, spatiotemporal observations of echolocation characteristics were conducted on bats during a landing flight task in the laboratory. The Telemike allowed us to observe emitted pulses and returning echoes to which the flying bats listened during flight, and the acoustic parameters could be precisely measured without traditional problems such as the directional properties of the recording microphone and the emitted pulse, or traveling loss of the sound in the air. Pulse intensity in bats intending to land exhibited a marked decrease by 30 dB within 2 m of the target wall, and the reduction rate was approximately 6.5 dB per halving of distance. The intensity of echoes returning from the target wall indicated a nearly constant intensity (-42.6 +/- 5.5 dB weaker than the pulse emitted in search phase) within a target distance of 2 m. These findings provide direct evidence that bats adjust pulse intensity to compensate for changes in echo intensity to maintain a constant intensity of the echo returned from the approaching target at an optimal range.     

The detection by sonar of difficult targets (including centimetre-scale plastic objects and optical fibres) buried in saturated sediment

This paper reports on a laboratory study into the use of sonar to detect objects, two of which exhibit a poor acoustic impedance mismatch with the water-saturated sediment in which they are buried to depth of about 30 cm. The targets are solid cylinders having diameters of 20-25 mm and 50 cm length, made of polyethylene, of telecommunications optical fibre, and of steel. Steel spheres are included for comparison. A poor acoustic impedance mismatch between the target and the host sediment is one factor that can make buried targets difficult to detect with sonar, but such detection is increasingly becoming an issue in a range of applications from archaeology to defence to telecommunications. Attention is paid to those signal processing techniques which could be of potential benefit. For this range of test objects, comparisons are made between use of optimal filtering and synthetic aperture sonar. In addition, the potential of a range of acousto-optical effects (optical time domain reflectometry, Raman and Brillouin scattering, and fibre optic hydrophones) is assessed in the Appendix for the particular application of detecting non-metallised fibre optic telecommunications cables. A web page dedicated to this paper hosts movies and reports at http://www.isvr.soton.ac.uk/fdag/uaua/target_in_sand.htm.     

Second-harmonic generation in Ge(20)As(25)S(55) glass irradiated by an electron beam

Second-harmonic generation was observed in Ge(20)As(25)S(55) chalcogenide glass irradiated by an electron beam. The second-harmonic intensity increased with increasing electron-beam current and accelerating voltage. The second-harmonic generation in Ge(20)As(25)S(55) glass was caused by the space-charge electrostatic field that was generated by irradiation of an electron beam. Second-order nonlinearity X((2)) as great as 0.8 pm/V was obtained. The results of measurements of thermally stimulated depolarization current indicated that the glass was poled in the thin layers of its surface (several micrometers) and that the nonlinearity was stable.     

Dimension reduction in heterogeneous neural networks: Generalized Polynomial Chaos (gPC) and ANalysis-Of-VAriance (ANOVA)

We propose, and illustrate via a neural network example, two different approaches to coarse-graining large heterogeneous networks. Both approaches are inspired from, and use tools developed in, methods for uncertainty quantification (UQ) in systems with multiple uncertain parameters – in our case, the parameters are heterogeneously distributed on the network nodes. The approach shows promise in accelerating large scale network simulations as well as coarse-grained fixed point, periodic solution computation and stability analysis. We also demonstrate that the approach can successfully deal with structural as well as intrinsic heterogeneities.