The sonar beam pattern of a flying bat as it tracks tethered insects

This paper describes measurements of the sonar beam pattern of flying echolocating bats, Eptesicus fuscus, performing various insect capture tasks in a large laboratory flight room. The beam pattern is deduced using the signal intensity across a linear array of microphones. The positions of the bat and insect prey are obtained by stereoscopic reconstruction from two camera views. Results are reported in the form of beam-pattern plots and estimated direction of the beam axis. The bat centers its beam axis on the selected target with a standard deviation (sigma) of 3 degrees. The experimental error is +/- 1.4 degrees. Trials conducted with two targets show that the bat consistently tracks one of the targets with its beam. These findings suggest that the axis of the bat sonar beam is a good index of selective tracking of targets, and in this respect is analogous to gaze in predominantly visual animals.     

Auditory scene analysis by echolocation in bats.

Echolocating bats transmit ultrasonic vocalizations and use information contained in the reflected sounds to analyze the auditory scene. Auditory scene analysis, a phenomenon that applies broadly to all hearing vertebrates, involves the grouping and segregation of sounds to perceptually organize information about auditory objects. The perceptual organization of sound is influenced by the spectral and temporal characteristics of acoustic signals. In the case of the echolocating bat, its active control over the timing, duration, intensity, and bandwidth of sonar transmissions directly impacts its perception of the auditory objects that comprise the scene. Here, data are presented from perceptual experiments, laboratory insect capture studies, and field recordings of sonar behavior of different bat species, to illustrate principles of importance to auditory scene analysis by echolocation in bats. In the perceptual experiments, FM bats (Eptesicus fuscus) learned to discriminate between systematic and random delay sequences in echo playback sets. The results of these experiments demonstrate that the FM bat can assemble information about echo delay changes over time, a requirement for the analysis of a dynamic auditory scene. Laboratory insect capture experiments examined the vocal production patterns of flying E. fuscus taking tethered insects in a large room. In each trial, the bats consistently produced echolocation signal groups with a relatively stable repetition rate (within 5%). Similar temporal patterning of sonar vocalizations was also observed in the field recordings from E. fuscus, thus suggesting the importance of temporal control of vocal production for perceptually guided behavior. It is hypothesized that a stable sonar signal production rate facilitates the perceptual organization of echoes arriving from objects at different directions and distances as the bat flies through a dynamic auditory scene. Field recordings of E. fuscus, Noctilio albiventris, N. leporinus, Pippistrellus pippistrellus, and Cormura brevirostris revealed that spectral adjustments in sonar signals may also be important to permit tracking of echoes in a complex auditory scene.     

The role of the external ear in vertical sound localization in the free flying bat, Eptesicus fuscus

The role of the external ear in sonar target localization for prey capture was studied by deflecting the tragus of six big brown bats, Eptesicus fuscus. The prey capture performance of the bat dropped significantly in the tragus-deflection condition, compared with baseline, control, and recovery conditions. Target localization error occurred in the tragus-deflected bat, and mainly in elevation. The deflection of the tragus did not abolish the prey capture ability of the bat, which suggests that other cues are available used for prey localization. Adaptive vocal and motor behaviors were also investigated in this study. The bat did not show significant changes in vocal behaviors but modified its flight trajectories in response to the tragus manipulation. The tragus-deflected bat tended to attack the prey item from above and had lower tangential velocity and larger bearing from the side, compared with baseline and recovery conditions. These findings highlight the contribution of the tragus to vertical sound localization in the free-flying big brown bat and demonstrate flight adaptations the bat makes to compensate altered acoustic cues.     

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. 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.     

Spatial perception and adaptive sonar behavior

Bat echolocation is a dynamic behavior that allows for real-time adaptations in the timing and spectro-temporal design of sonar signals in response to a particular task and environment. To enable detailed, quantitative analyses of adaptive sonar behavior, echolocation call design was investigated in big brown bats, trained to rest on a stationary platform and track a tethered mealworm that approached from a starting distance of about 170 cm in the presence of a stationary sonar distracter. The distracter was presented at different angular offsets and distances from the bat. The results of this study show that the distance and the angular offset of the distracter influence sonar vocalization parameters of the big brown bat, Eptesicus fuscus. Specifically, the bat adjusted its call duration to the closer of two objects, distracter or insect target, and the magnitude of the adjustment depended on the angular offset of the distracter. In contrast, the bat consistently adjusted its call rate to the distance of the insect, even when this target was positioned behind the distracter. The results hold implications for understanding spatial information processing and perception by echolocation.     

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.     

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.     

The bat head-related transfer function reveals binaural cues for sound localization in azimuth and elevation

Directional properties of the sound transformation at the ear of four intact echolocating bats, Eptesicus fuscus, were investigated via measurements of the head-related transfer function (HRTF). Contributions of external ear structures to directional features of the transfer functions were examined by remeasuring the HRTF in the absence of the pinna and tragus. The investigation mainly focused on the interactions between the spatial and the spectral features in the bat HRTF. The pinna provides gain and shapes these features over a large frequency band (20-90 kHz), and the tragus contributes gain and directionality at the high frequencies (60 to 90 kHz). Analysis of the spatial and spectral characteristics of the bat HRTF reveals that both interaural level differences (ILD) and monaural spectral features are subject to changes in sound source azimuth and elevation. Consequently, localization cues for horizontal and vertical components of the sound source location interact. Availability of multiple cues about sound source azimuth and elevation should enhance information to support reliable sound localization. These findings stress the importance of the acoustic information received at the two ears for sound localization of sonar target position in both azimuth and elevation.     

基于粒子滤波的多普勒信息辅助目标定位跟踪算法

多基地声呐探测系统主要通过测量回波的时延和方位信息进行目标定位与跟踪,定位精度受声速、时延和方位测量误差的影响较大,可以通过多普勒信息辅助进一步提高定位跟踪精度.现有的多普勒信息辅助定位跟踪算法多适用于单基地声呐系统,多基地中的多普勒测量值与目标状态的关系更为复杂,需要研究新的融合方法.该文提出了一种适用于多基地声呐系...     

Echolocation and flight strategy of Japanese house bats during natural foraging, revealed by a microphone array system

Using only a microphone array system, echolocation pulses and three-dimensional flight paths in the frequency-modulated bat, Pipistrellus abramus, during natural foraging, were simultaneously examined. During the search phase, the inter-pulse interval, pulse duration, and moving distance of the bat between successive emissions were relatively constant at around 89.5 ± 18.7 ms, 6.90 ± 1.31 ms, and 0.50 ± 0.20 m, respectively. The bats started to decrease these acoustical parameters within 2-3 m of the prey capture point. For every emission along a flight path, the distance between a bat and its prey capture point was calculated as both direct distance to capture (DDC), which corresponded to the target distance, and flight distance to capture (FDC) along the flight path. The DDC matched the FDC after the start of the approach phase, indicating that foraging bats followed a straight-ahead path to the target. In addition, the duration of the quasi-constant frequency component of emitted pulses was slightly extended just before the convergence of the DDC with the FDC. These findings suggest that the bats confirm the presence of target prey by extending the duration of the pulse and then select a straight-ahead approach by forecasting the movement of the prey.     

Insights into dolphin sonar discrimination capabilities from human listening experiments

A variety of dolphin sonar discrimination experiments have been conducted, yet little is known about the cues utilized by dolphins in making fine target discriminations. In order to gain insights on cues available to echolocating dolphins, sonar discrimination experiments were conducted with human subjects using the same targets employed in dolphin experiments. When digital recordings of echoes from targets ensonified with a dolphinlike signal were played back at a slower rate to human subjects, they could also make fine target discriminations under controlled laboratory conditions about as well as dolphins under less controlled conditions. Subjects reported that time-separation-pitch and duration cues were important. They also reported that low-amplitude echo components 32 dB below the maximum echo component were usable. The signal-to-noise ratio had to be greater than 10 dB above the detection threshold for simple discrimination and 30 dB for difficult discrimination. Except for two cases in which spectral cues in the form of "click pitch" were important, subjects indicated that time-domain rather than frequency-domain processing seemed to be more relevant in analyzing the echoes.     

Morphology-induced information transfer in bat sonar

It has been argued that an important part of understanding bat echolocation comes down to understanding the morphology of the bat sound processing apparatus. In this Letter we present a method based on information theory that allows us to assess target localization performance of bat sonar, without a?priori knowledge on the position, size, or shape of the reflecting target. We demonstrate this method using simulated directivity patterns of the frequency-modulated bat Micronycteris microtis. The results of this analysis indicate that the morphology of this bat's sound processing apparatus has evolved to be a compromise between sensitivity and accuracy with the pinnae and the noseleaf playing different roles.     

水中目标回波亮点统计特征研究

礁石和海洋动物引起的混响是主动声纳最严重的干扰, 如何区分礁石、鱼群和水中目标一直是制约主动声纳识别技术的难点问题. 针对礁石与目标回波难以区分的问题, 从特征识别的应用角度, 研究水中复杂目标全方位回波亮点特征的有效表征和应用方式, 基于目标回波亮点模型, 提出拷贝相关器输出的目标散射函数估计方法, 给出对目标回波亮点相对关系进行定量分析的目标回波特征统计表征方式, 并基于湖上实验提取了物理机理明确的目标回波亮点统计特征, 使得目标时间-角度谱中所蕴含的目标特征信息能够很直接地转化为主动声纳易于应用的目标特征. 关键词： 水中目标 回波亮点 统计特征     

A fluid–structure interaction model of insect flight with flexible wings

We present a fluid–structure interactions (FSI) model of insect flapping flight with flexible wings. This FSI-based model is established by loosely coupling a finite element method (FEM)-based computational structural dynamic (CSD) model and a computational fluid dynamic (CFD)-based insect dynamic flight simulator. The CSD model is developed specifically for insect flapping flight, which is capable to model thin shell structures of insect flexible wings by taking into account the distribution and anisotropy in both wing morphology involving veins, membranes, fibers and density, and in wing material properties of Young’s modulus and Poisson’s ratios. The insect dynamic flight simulator that is based on a multi-block, overset grid, fortified Navier–Stokes solver is capable to integrate modeling of realistic wing-body morphology, realistic flapping-wing and body kinematics, and unsteady aerodynamics in flapping-wing flights. Validation of the FSI-based aerodynamics and structural dynamics in flexible wings is achieved through a set of benchmark tests and comparisons with measurements, which contain a heaving spanwise flexible wing, a heaving chordwise-flexible wing with a rigid teardrop element, and a realistic hawkmoth wing rotating in air. A FSI analysis of hawkmoth hovering with flapping flexible wings is then carried out and discussed with a specific focus on the in-flight deformation of the hawkmoth wings and hovering aerodynamic performances with the flexible and rigid wings. Our results demonstrate the feasibility of the present FSI model in accurately modeling and quantitatively evaluating flexible-wing aerodynamics of insect flapping flight in terms of the aerodynamic forces, the power consumption and the efficiency.     

Clutter interference along the target range axis in the echolocating bat, Eptesicus fuscus

The sensitivity of the echolocating bat, Eptesicus fuscus, for detection of a sonar target is impaired by the presence of additional targets located at similar distances. At a range of 54 cm, sensitivity to one target declines if the range separation to other targets is smaller than 8-9 cm. This loss of sensitivity is an example of clutter interference along the range axis, created by simultaneous masking of one set of echoes by another. Echoes that fall within an experimentally defined critical range band may sum together to contribute collectively to detection in that band. Echoes falling into separate bands may be detected independently. Acoustic glints within a band appear to be grouped together to be perceived as a single range-extended target of complex structure. Range bands may thus define what a "target" is by specifying within-target and between-target differences in range. The width of critical range bands appears to depend upon target range, with wider bands at greater ranges.     

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 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.     

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.     

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.