Auditory Dimensions of Acoustic Images in Echolocation
Springer handbook of auditory researchPublished 1 January 1995
James A. Simmons, Michael J. Ferragamo, Prestor A. Saillant, Tim Haresign, Janine M. Wotton, Steven P. Dear
Citations58
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TL;DR
The exaggeration of aspects of auditory function to achieve spatial imaging reflects the vital role of hearing in the lives of bats — for finding prey and perceiving obstacles to flight and highlights the mechanisms behind these functions.
Abstract
Echolocation in bats is one of the most demanding adaptations of hearing to be found in any animal. Transforming the information carried by sounds into perceptual images depicting the location and identity of objects rapidly enough to control the decisions and reactions of a swiftly flying bat is a prodigious task for the auditory system to accomplish. The exaggeration of aspects of auditory function to achieve spatial imaging reflects the vital role of hearing in the lives of bats — for finding prey and perceiving obstacles to flight (Neuweiler 1990). It also highlights the mechanisms behind these functions to make echolocation a useful model for studying how the auditory system processes information and creates auditory perceptions in the most extreme circumstances.
Keywords
Agricultural and Biological SciencesEnvironmental Science
The Journal of the Acoustical Society of AmericaThe resolution of target range by echolocating bats
403 Citations1973James A. Simmons
Comparisons between discrimination performance and autocorrelation functions of echolocation sounds used in the discriminations suggest that these bats possess some neural equivalent of a matched‐filter, ideal sonar receiver which functionally cross‐correlates a replica of the outgoing signal with the returning echo to detect the echo and determine its arrival time.
The Journal of the Acoustical Society of AmericaDetection, estimation, and classification with spectrograms
259 Citations1980Richard A. Altes
Spectrogram correlation can be used for classification as well as for estimation and detection, and for maximum likelihood parameter estimation, e.g., estimation of delay or center frequency of a signal.
Journal of NeuroscienceThe monaural nuclei of the lateral lemniscus in an echolocating bat: parallel pathways for analyzing temporal features of sound
221 Citations1991Ellen Covey, JH Casseday
The results indicate that VNLLc is specialized to encode very precisely the onset of sound; the other nuclei may encode ongoing properties of a sound.
SciencePerception of Echo Phase Information in Bat Sonar
213 Citations1979James A. Simmons
Echolocating bats (Eptesicus fuscus) can detect changes as small as 500 nanoseconds in the arrival time of sonar echoes when changes appear as jitter or alternations in arrival time from one echo to the next.
The Journal of the Acoustical Society of America<i>Animal Sonar: Processes and Performance</i>
194 Citations1990Stewart H. Hulse
A Comparative Neurophysiological and Neuroanatomical Study of Echolocation Signals in Dolphins, Whale Heads and Dolphin Forehead, and Cochlear Physiology and Anatomy in Bats.
Journal of NeuroscienceAutomatic gain control in the bat's sonar receiver and the neuroethology of echolocation
193 Citations1984SA Kick, J. A. Simmons
It is hypothesized that the bat's directional emissions, directional hearing, middle ear muscle contractions, and head aim response collectively create a three- dimensional spatial tracking filter which the bat locks onto targets to stabilize echo amplitudes during interception of prey.
Journal of Comparative Physiology ADiscrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: The shape of target images in echolocation
184 Citations1990JamesA. Simmons, Michael J. Ferragamo +3 more
The locations of both 0° and 180° phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.
Performance of Airborne Animal Sonar Systems: I. Microchiroptera
181 Citations1980Hans‐Ulrich Schnitzler, O. W. Henson
Some animals, during evolution, developed eyes which were especially adapted for good visual efficiency at low light intensities whereas others improved the remaining sensory systems.
CognitionA view of the world through the bat's ear: The formation of acoustic images in echolocation
177 Citations1989James A. Simmons
The bat's use of cross-correlation-like images reveals neural computations that achieve fusion of stimulus features and offers an example of high-level operations involved in the formation of perceptual "wholes".
The Journal of the Acoustical Society of AmericaA computational model of echo processing and acoustic imaging in frequency- modulated echolocating bats: The spectrogram correlation and transformation receiver
177 Citations1993Prestor A. Saillant, James A. Simmons +2 more
The spectrogram correlation and transformation model of the sonar receiver in the big brown bat accurately reproduces the images perceived by Eptesicus in a variety of behavioral experiments on two-glint resolution in range, echo phase sensitivity, amplitude-latency trading of range estimates, dissociation of time- and frequency-domain image components, and ranging accuracy in noise.
Journal of Comparative Physiology ATarget-detection by the echolocating bat,Eptesicus fuscus
154 Citations1982Shelley A. Kick
The results demonstrate that the maximum effective range of bat sonar is greater than previously indicated by obstacleavoidance and target-interception tasks.
ZoophysiologyThe Neural Basis of Echolocation in Bats
144 Citations1989George D. Pollak, John H. Casseday
This monograph is about analysis of sound in the brainstem of echolocating bats and concerns the relationship between brain structure and brain function.
The Journal of the Acoustical Society of AmericaInformation in sonar echoes of fluttering insects available for echolocating bats
142 Citations1990Rudolf Kober, Hans‐Ulrich Schnitzler
Journal of NeurophysiologyTonotopic and functional organization in the auditory cortex of the big brown bat, Eptesicus fuscus
134 Citations1993Steven P. Dear, Jonathan B. Fritz +3 more
Journal of Comparative Physiology AConvergence of temporal and spectral information into acoustic images of complex sonar targets perceived by the echolocating bat, Eptesicus fuscus
131 Citations1990JamesA. Simmons, Cynthia F. Moss +1 more
FM echolocating bats (Eptesicus fuscus) were trained to discriminate between a two-component complex target and a one-component simple target simulated by electronically-returned echoes in a series of experiments that explore the composition of the image of the two- component target.
Proceedings in life sciencesTime Resolution in Auditory Systems
129 Citations1985Axel Michelsen
It is argued why the auditory system is capable of detecting a ,2-4 dB decrement of intensity in approx.
The Journal of PhysiologyRecovery cycles and responses to frequency modulated tone pulses in auditory neurones of echo‐locating bats
126 Citations1964Nobuo Suga
Journal of Comparative Physiology AAcuity of horizontal angle discrimination by the echolocating bat,Eptesicus fuscus
124 Citations1983J. A. Simmons, Shelley A. Kick +4 more
The observed performance of Eptesicus fuscus at horizontal-angle discrimination matches the performance to be expected if the 0.5 μs acuity of perception of the time-of-occurrence of sonar echoes measured in echo-jitter experiments can be applied to binaural echo arrivaltime perception.
NatureA possible neuronal basis for representation of acoustic scenes in auditory cortex of the big brown bat
114 Citations1993Steven P. Dear, James A. Simmons +1 more
It is reported here that response properties of delay-tuned neurons in the cortical tonotopic area of the bat, Eptesicus, transform the sequential arrival times of echoes with different delays into a concurrent, accumulating neural representation of multiple objects at different ranges.
The Journal of the Acoustical Society of AmericaThe sound emission pattern of the echolocating bat, <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>
114 Citations1989David J. Hartley, Roderick A. Suthers
Journal of NeurophysiologyInteraural time and intensity coding in superior olivary complex and inferior colliculus of the echolocating bat Molossus ater
114 Citations1985G. Harnischfeger, Gerhard Neuweiler +1 more
Single-unit responses to tonal stimulation with interaural disparities were recorded in the nuclei of the superior olivary complex (SOC) and the central nucleus of the inferior colliculus (ICC) of the echolocating bat, Molossus ater, and it is concluded that the neurons reacted toInteraural differences of stimulus-onset time (transient time difference) but not to phase differences (ongoing time difference).
The Journal of PhysiologyThe neurophysiology of audition in bats: temporal parameters
109 Citations1963Alan D. Grinnell
The electrophysiological data that exist for mammals, mostly cats, suggest a recovery time very similar to that ofhuman listeners, when comparable stimulus conditions are employed, conditions analogous to those facing an echolocating bat.
The Journal of the Acoustical Society of AmericaCoding and processing in the auditory systems of FM-signal-producing bats
107 Citations1973Nobuo Suga, Peter Schlegel
For echolocation, certain insectivorous bats emit frequency‐modulated (FM) sounds in which frequency always sweeps from high to low, and although seemingly paradoxical, such properties are easily explained by neural network models.
Hearing ResearchTime is traded for intensity in the bat's auditory system
98 Citations1988George D. Pollak
The main conclusion is that the auditory system processes interaural disparities by transforming level differences at the two ears into latency differences, and the discharge probability of each binaural neuron is determined largely by the arrival times of the discharges from the excitatory and inhibitory ears.
The Journal of the Acoustical Society of AmericaThe acoustic basis for target discrimination by FM echolocating bats
93 Citations1989James A. Simmons, Lynda Chen
The acoustic properties of the targets are measured 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.
Animal BehaviourThe role of the flight membranes in insect capture by bats
89 Citations1962F. A. Webster, Donald R. Griffin
While the use of tail and wing membranes greatly increases the potential area of contact with insect prey over the area of the opened mouth alone, the photographs almost invariably show that the bat's head is pointed at the insect well before contact with it, and these photographs strongly indicate that each insect is individually located and intercepted.
Biological BulletinTHE SENSITIVITY OF ECHOLOCATION IN BATS
88 Citations1958Alan D. Grinnell, Donald R. Griffin
Thedistance at which bats (Myotis lucifugus) react to the presence of a row of small wires has been measured by a photographic determination of the distance at which the pulse repetition rate first increases as the bats fly towards the wires.
Hearing ResearchDirectionality of sound pressure transformation at the pinna of echolocating bats
83 Citations1988Philip H.-S. Jen, De‐Mao Chen
The directionality of sound pressure transformation at the pinna of three species of bats was studied by measuring the sound pressure level of a tone at the tympanic membrane as a function of azimuth and elevation of the sound source under free-field conditions.
Journal of Experimental ZoologyTarget discrimination by the echolocation of bats
83 Citations1965Donald R. Griffin, J. H. Friend +1 more
These bats seem capable of some qualitative discrimination based on the “fine structure” of echoes from small moving targets, as evidenced by the overlap in intensity both in the peak amplitudes and in each of the ten 8 kc bands from 100–20 kc.
Journal of Comparative Physiology ACommon principle of guidance by echolocation and vision
79 Citations1992David N. Lee, F. R. van der Weel +3 more
Using echolocation, bats move as gracefully as birds through the cluttered environment, suggesting common principles of optic and acoustic guidance, and simpler control is possible using the tau function of any sensory variable S that is a power function of distance to aperture.
The Journal of the Acoustical Society of AmericaPerception of structured phantom targets in the echolocating bat, <i>M</i> <i>e</i> <i>g</i> <i>a</i> <i>d</i> <i>e</i> <i>r</i> <i>m</i> <i>a</i> <i>l</i> <i>y</i> <i>r</i> <i>a</i>
78 Citations1992Sabine Schmidt
The results presented here show that the discrimination of target surface structure can be explained as a discrimination of echo spectra.
The Journal of the Acoustical Society of AmericaStabilization of perceived echo amplitudes in echolocating bats. II. The acoustic behavior of the big brown bat, <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>, when tracking moving prey
76 Citations1992David J. Hartley
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.
ScienceSonar Tracking of Horizontally Moving Targets by the Big Brown Bat <i>Eptesicus fuscus</i>
74 Citations1985W. Mitch Masters, Anne J. M. Moffat +1 more
When following a moving target, echolocating bats (Eptesicus fuscus) keep their heads aimed at the target's position, and the bat obtains frequent position updates by emitting sonar signals at a high rate.
The Journal of the Acoustical Society of AmericaClutter interference and the integration time of echoes in the echolocating bat, <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>
70 Citations1989James A. Simmons, Edward G. Freedman +3 more
The ability of the echolocating bat to detect a sonar target is affected by the presence of other targets along the same axis at slightly different ranges, and clutter interference occurs and one set of echoes masks the other.
Hearing ResearchTime and frequency domain processing in the inferior colliculus of echolocating bats
63 Citations1981R. Bodenhamer, George D. Pollak
In response to FM signals, the cells exhibited a much greater frequency selectivity than that observed following tone burst stimulation, and the potential role of this dual frequency coding in spectral and temporal processing of biosonar signals and echoes is discussed.
The Journal of the Acoustical Society of AmericaAcoustic information available to bats using frequency- modulated sounds for the perception of insect prey
62 Citations1994Cynthia F. Moss, Mark Zagaeski
It has been proposed that the FM bat receives stroboscopic-like glimpses of fluttering prey whose spatial representation depends on the operation of the bat's sonar receiver.
Echolocation Signals and Echoes in Air
62 Citations1980J. D. Pye
Since the Frascati meeting in 1966, there has been considerable expansion in understanding of the acoustic structures used by bats and birds for echolocation in air, due to the development of instrumental techniques which were already undergoing something of a revolution at the time.
The Journal of the Acoustical Society of AmericaSensorimotor model of bat echolocation and prey capture
62 Citations1994Roman Kuc
This model indicates that successful nonpredictive tracking of an ideal prey can be accomplished with a very simple system of bat sensorimotor system, using the big brown bat Eptesicus fuscus as a model.
Journal of Comparative Physiology AEvidence for perception of fine echo delay and phase by the FM bat, Eptesicus fuscus
57 Citations1993James A. Simmons
Amplitude-latency trading of perceived delay confirms that the big brown bat's fine delay and phase perception depends on a temporal neural code, which depends on stimulus delays, not the delays of extraneous sounds.
The Performance of Echolocation: Acoustic Images Perceived by Echolocating Bats
57 Citations1988James A. Simmons, Alan D. Grinnell
The performance of echolocation in bats is a measure of the effectiveness of this system of orientation for representing significant features of objects in perceptual images.
The Journal of the Acoustical Society of AmericaRange estimation by echolocation in the bat <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>: Trading of phase versus time cues
47 Citations1989Dieter Menne, Ingrid Kaipf +3 more
Bats of the species Eptesicus fuscus have been trained to discriminate a stationary simulated target from a target with a virtual distance that jitters from sound to sound, and were able to detect a pure phase jitter, e.g., polarity inversion of the signal.
The Journal of the Acoustical Society of AmericaRange resolution and the possible use of spectral information in the echolocating bat, <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>
41 Citations1990Joachim Mogdans, Hans‐Ulrich Schnitzler
The occurrence of spectral minima within the frequency range of the first harmonic in the echo of thebats' 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 thefirst harmonic in echoes for range resolution.
Journal of Comparative Physiology ADiscrimination of fluttering targets by the FM-batPipistrellus stenopterus?
40 Citations1988Y. W. Sum, D. Menne
Five bats of the species Pipistrellus stenopterus were trained in a two-alternative forced-choice procedure to discriminate between two fluttering targets, comparable to that of bats emitting longer sounds with constant-frequency (CF) components and a higher duty cycle.
Springer handbook of auditory researchAuditory Computations for Biosonar Target Imaging in Bats
37 Citations1996James A. Simmons, Prestor A. Saillant +5 more
Bats are nocturnal flying mammals classified in the order Chiroptera that have evolved a biological sonar, called echolocation, to orient in darkness—to guide their flight around obstacles and to detect their prey.
Journal of Comparative Physiology ASome comments on the proposed perception of phase and nanosecond time disparities by echolocating bats
36 Citations1993George D. Pollak
The purpose of this review is to point out a number of technical problems and deficiencies in those experiments which undermine the veracity of the purported demonstration of phase perception and nanosecond time resolution by bats.
The Journal of the Acoustical Society of AmericaUltrasonic spectroscopy in bovine serum albumin solutions
34 Citations1990P.-K. Choi, J.-R. Bae +1 more
The Journal of the Acoustical Society of AmericaAcoustic image representation of a point target in the bat <i>E</i> <i>p</i> <i>t</i> <i>e</i> <i>s</i> <i>i</i> <i>c</i> <i>u</i> <i>s</i> <i>f</i> <i>u</i> <i>s</i> <i>c</i> <i>u</i> <i>s</i>: Evidence for sensitivity to echo phase in bat sonar
32 Citations1993Cynthia F. Moss, James A. Simmons
The results suggest that Eptesicus is sensitive to the phase reversal of echoes and thus have implications for assessing receiver models of echolocation.
Journal of Comparative Physiology ADiscrimination of wingbeat motion by bats, correlated with echolocation sound pattern
31 Citations1991RoaldC. Roverud, Volker Nitsche +1 more
Behavioral evidence is provided supporting the hypothesis that bats that use CF/FM echolocation sounds have adaptations for the perception of insect wingbeat motion and that long-CF/FM species are more specialized for this task than short-CF-FM species.
A New Concept of Echo Evaluation in the Auditory System of Bats
30 Citations1980Karl Beuter
Bats use their echolocation system very effectively for spatial orientation and prey hunting and important tasks are the detection of weak target echoes or echoes superimposed by clutter-interference, the precise localization of targets, and the identification of target structure.
Proceedings in life sciencesRange Determination by Measuring Time Delays in Echolocating Bats
24 Citations1985Hans‐Ulrich Schnitzler, Dieter Menne +1 more
Annual Review of PhysiologyPhysiological Mechanisms for Spatial Filtering and Image Enhancement in the Sonar of Bats
23 Citations1984J. A. Simmons, Shelley A. Kick
The Journal of the Acoustical Society of AmericaImpulse-response method to predict echo responses from targets of complex geometry. Part II. Computer implementation and experimental validation
20 Citations1994Alain Lhémery, Raphaële Raillon
IEEE Transactions on Sonics and UltrasonicsTexture Analysis with Spectrograms
15 Citations1984Richard A. Altes
The Journal of the Acoustical Society of AmericaStroboscopic hearing as a mechanism for prey discrimination in frequency-modulated bats?
11 Citations1994Albert S. Feng, Curtis J. Condon +1 more
FM bats can potentially employ a "stroboscopic hearing" strategy for discriminating insects on the basis of the wing-beat frequency of the prey, and data indicate that the across-pulse amplitude modulation becomes undetectable when the AM frequency is integer multiples of the pulse repetition rate.
Proceedings in life sciencesTheoretical Limits of Time Resolution in Narrow Band Neurons
4 Citations1985Dieter Menne
The apparent time delay between two peaks of a signal corrupted by noise will not be the same when measured repeatedly, but will show some statistical variance, so at low noise levels the variance will be low and therefore the measurement will be more accurate than at high noise levels.