A view of the world through the bat's ear: The formation of acoustic images in echolocation
Abstract
Echolocating bats perceive objects as acoustic images derived from echoes of the ultrasonic sounds they emit. They can detect, track, identify, and intercept flying insects using sonar. Many species, such as the big brown bat, Eptesicus fuscus, emit frequency-modulated sonar sounds and perceive the distance to targets, or target range, from the delay of echoes. For Eptesicus, a point-target's image has a sharpness along the range axis that is determined by the acuity of echo-delay perception, which is about 10 ns under favorable conditions. The image as a whole has a fine range structure that corresponds to the cross-correlation function between emissions and echoes. A complex target- which has reflecting points, called "glints", located at slightly different distances and reflects echoes containing overlapping components with slightly different delays--is perceived in terms of its range profile. The separation of the glints along the range dimension is encoded by the shape of the echo spectrum created by interference between overlapping echo components. However, Eptesicus transforms the echo spectrum back into an estimate of the original delay separation of echo components. The bat thus converts spectral cues into elements of an image expressed in terms of range. The absolute range of the nearest glint is encoded by the arrival time of the earliest echo component, and the spectrally encoded range separation of additional glints is referred to this time-encoded reference range for the image as a whole. Each individual glint is represented by a cross-correlation function for its own echo component, the nearest of which is computed directly from arrival-time measurements while further ones are computed by transformation of the echo spectrum. The bat then sums the cross-correlation functions for multiple glints to form the entire image of the complex target. Range and shape are two distinct features of targets that are separately encoded by the bat's auditory system, but the bat perceives unitary images that require fusion of these features to create a synthetic psychological dimension of range. 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".
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The active process of emission and comparison enables the bat to assemble an image of the external world and thus successfully navigate and predate.
“Like the mormyrid, this active process of emission and comparison enables the bat to assemble an image of the external world55–57 and thus successfully navigate and predate.”
From Corollary discharge across the animal kingdom (2008) · cites this paper as [55]
Cites
Detection, Estimation, And Modulation Theory
Trees, H. L. V. (1968) · Cited by 5,964 · see citing works
Detection, Estimation, and Modulation Theory, Part I
Trees, H. L. V. (2001) · John Wiley & Sons, Inc. eBooks · doi:10.1002/0471221082 · Cited by 3,923 · see citing works
Probability and Information Theory with Applications to Radar
(1953) · Elsevier eBooks · doi:10.1016/c2013-0-05390-x · Cited by 1,287 · see citing works
Griffin, D. R., & Lindsay, R. B. (1958) · Cited by 855 · see citing works
Journal of Comparative Physiology A
(2013) · Journal of Comparative Physiology A · doi:10.1007/359.1432-1351 · Cited by 618 · see citing works
Morphology, Echolocation and Resource Partitioning in Insectivorous Bats
Aldridge, H. D. J. N., & Rautenbach, I. L. (1987) · Journal of Animal Ecology · doi:10.2307/4947 · Cited by 579 · see citing works
Detection, Estimation, and Modulation Theory, Part III
Trees, H. L. V. (2001) · John Wiley & Sons, Inc. eBooks · doi:10.1002/0471221090 · Cited by 550 · see citing works
(1973) · Elsevier eBooks · doi:10.1016/b978-0-125-04250-5.x5001-2 · Cited by 527 · see citing works
The resolution of target range by echolocating bats
Simmons, J. A. T. (1973) · The Journal of the Acoustical Society of America · doi:10.1121/1.1913559 · Cited by 403 · see citing works
Self-citationLawrence, B. D., & Simmons, J. A. T. (1982) · The Journal of the Acoustical Society of America · doi:10.1121/1.387529 · Cited by 396 · see citing works
Self-citationAdvances in Vertebrate Neuroethology
Ewert, J.-P., Capranica, R. R., & Ingle, D. (1983) · Springer eBooks · doi:10.1007/978-1-4684-4412-4 · Cited by 394 · see citing works
Echolocation and Pursuit of Prey by Bats
Simmons, J. A., Fenton, M. B., & O'Farrell, M. J. (1979) · Science · doi:10.1126/science.758674 · Cited by 391 · see citing works
Self-citationEcholocation and Pursuit of Prey by Bats
Pontecorvo, B., & Álvarez, W. A. (1979) · Cited by 352 · see citing works
Acoustic imaging in bat sonar: Echolocation signals and the evolution of echolocation
Simmons, J. A. T., & Stein, R. A. (1980) · Journal of Comparative Physiology A · doi:10.1007/bf00660182 · Cited by 336 · see citing works
Self-citationHill, J. E., Hill, J. E., Smith, H. J. D., et al. (1984) · Cited by 300 · see citing works
The importance of atmospheric attenuation for the echolocation of bats (Chiroptera)
Griffin, D. R. (1971) · Animal Behaviour · doi:10.1016/s0003-3472(71)80134-3 · Cited by 279 · see citing works
Probability and information theory, with applications to radar
SMITH–ROSE, R. L. (1954) · Journal of Atmospheric and Terrestrial Physics · doi:10.1016/0021-9169(54)90059-8 · Cited by 246 · see citing works
Perception of Echo Phase Information in Bat Sonar
Simmons, J. A. T. (1979) · Science · doi:10.1126/science.451543 · Cited by 214 · see citing works
Self-citationAutomatic gain control in the bat's sonar receiver and the neuroethology of echolocation
Kick, S., & Simmons, J. A. (1984) · Journal of Neuroscience · doi:10.1523/jneurosci.04-11-02725.1984 · Cited by 192 · see citing works
Self-citationNeuroethology and Behavioral Physiology: Roots and Growing Points
Huber, F., & Markl, H. S. (1983) · Cited by 191 · see citing works
Foraging, echolocation and audition in bats
Neuweiler, G. (1984) · Die Naturwissenschaften · doi:10.1007/bf00455897 · Cited by 188 · see citing works
Simmons, J. A. T., Ferragamo, M. J., Moss, C. F., Stevenson, S. B., & Altes, R. A. (1990) · Journal of Comparative Physiology A · doi:10.1007/bf00192654 · Cited by 183 · see citing works
Self-citationPerformance of Airborne Animal Sonar Systems: I. Microchiroptera
Schnitzler, H., & Henson, O. W. (1980) · Springer eBooks · doi:10.1007/978-1-4684-7254-7_6 · Cited by 181 · see citing works
The echolocation and hunting behavior of the bat,Pipistrellus kuhli
Schnitzler, H. U., Kalko, E. K. V., Miller, L. A., & Surlykke, A. (1987) · Journal of Comparative Physiology A · doi:10.1007/bf00615246 · Cited by 168 · see citing works
Functional properties of auditory neurones in the cortex of echo‐locating bats.
Suga, N. (1965) · The Journal of Physiology · doi:10.1113/jphysiol.1965.sp007791 · Cited by 161 · see citing works
Target-detection by the echolocating bat,Eptesicus fuscus
Kick, S. A. (1982) · Journal of Comparative Physiology A · doi:10.1007/bf00612808 · Cited by 154 · see citing works
Information content of bat sonar echoes.
Simmons, J. A. T., Howell, D. J., & Suga, N. (1975) · PubMed · Cited by 129 · see citing works
Self-citationTarget Structure and Echo Spectral Discrimination by Echolocating Bats
Simmons, J. A. T., Lavender, W. A., Lavender, B. A., et al. (1974) · Science · doi:10.1126/science.186.4169.1130 · Cited by 129 · see citing works
Self-citationAcuity of horizontal angle discrimination by the echolocating bat,Eptesicus fuscus
Simmons, J. A., Kick, S. A., Lawrence, B. D., et al. (1983) · Journal of Comparative Physiology A · doi:10.1007/bf00612586 · Cited by 117 · see citing works
Self-citationFEATURE EXTRACTION IN THE AUDITORY SYSTEM OF BATS
Suga, N. (1973) · Elsevier eBooks · doi:10.1016/b978-0-12-504250-5.50027-8 · Cited by 112 · see citing works
Discrimination of surface-structured targets by the echolocating batMyotis myotis during flight
Habersetzer, J., & Vogler, B. (1983) · Journal of Comparative Physiology A · doi:10.1007/bf00611192 · Cited by 105 · see citing works
Covey, E., & Casseday, J. (1986) · Journal of Neuroscience · doi:10.1523/jneurosci.06-10-02926.1986 · Cited by 104 · see citing works
Neural representation of target distance in auditory cortex of the echolocating bat Myotis lucifugus
Sullivan, W. E. (1982) · Journal of Neurophysiology · doi:10.1152/jn.1982.48.4.1011 · Cited by 103 · see citing works
The Acoustical Image of Fluttering Insects in Echolocating Bats
Schnitzler, H., Menne, D., Kober, R., & Heblich, K. (1983) · Springer eBooks · doi:10.1007/978-3-642-69271-0_18 · Cited by 101 · see citing works
Neuweiler, G., Bruns, V., & Schuller, G. (1980) · The Journal of the Acoustical Society of America · doi:10.1121/1.384812 · Cited by 100 · see citing works
Evidence for a spectral basis of texture perception in bat sonar
Schmidt, S. M. (1988) · Nature · doi:10.1038/331617a0 · Cited by 99 · see citing works
Echolocation: Discrimination of targets by the bat, Eptesicus fuscus
Simmons, J. A. T., & Vernon, J. A. (1971) · Journal of Experimental Zoology · doi:10.1002/jez.1401760307 · Cited by 99 · see citing works
Self-citationDalland, J. I. (1965) · Science · doi:10.1126/science.150.3700.1185 · Cited by 98 · see citing works
Sonar for generalized target description and its similarity to animal echolocation systems
Altes, R. A. (1976) · The Journal of the Acoustical Society of America · doi:10.1121/1.380831 · Cited by 97 · see citing works
Echolocation: Implications for Ecology and Evolution of Bats
Fenton, M. B. (1984) · The Quarterly Review of Biology · doi:10.1086/413674 · Cited by 95 · see citing works
Doppler-shift compensation in insect-catching horseshoe bats
Trappe, M., & Schnitzler, H. U. (1982) · Die Naturwissenschaften · doi:10.1007/bf00364902 · Cited by 93 · see citing works
The acoustic basis for target discrimination by FM echolocating bats
Simmons, J. A. T., & Chen, L. (1989) · The Journal of the Acoustical Society of America · doi:10.1121/1.398694 · Cited by 93 · see citing works
Self-citationThe role of the flight membranes in insect capture by bats
Webster, F. A., & Griffin, D. R. (1962) · Animal Behaviour · doi:10.1016/0003-3472(62)90056-8 · Cited by 91 · see citing works
Accuracy of target ranging in echolocating bats: acoustic information processing
Moss, C. F., & Schnitzler, H. U. (1989) · Journal of Comparative Physiology A · doi:10.1007/bf00619357 · Cited by 90 · see citing works
Sonar for generalized target description and its similarity to animal echolocation systems
Altes, R. A. (1975) · The Journal of the Acoustical Society of America · doi:10.1121/1.2001919 · Cited by 90 · see citing works
ECHOLOCATION OF FLYING INSECTS BY THE BAT, CHILONYCTERIS PARNELLII
Novick, A., & Vaišnys, J. R. (1964) · Biological Bulletin · doi:10.2307/1539249 · Cited by 89 · see citing works
Suga, N. (1988) · Medical Entomology and Zoology · Cited by 85 · see citing works
Target discrimination by the echolocation of bats
Griffin, D. R., Friend, J. H., & Webster, F. A. (1965) · Journal of Experimental Zoology · doi:10.1002/jez.1401580204 · Cited by 82 · see citing works
Sonar Tracking of Horizontally Moving Targets by the Big Brown Bat Eptesicus fuscus
Masters, W. M., Moffat, A. J. M., & Simmons, J. A. T. (1985) · Science · doi:10.1126/science.4001947 · Cited by 74 · see citing works
Self-citationSimmons, J. A. T., Freedman, E. G., Stevenson, S. B., et al. (1989) · The Journal of the Acoustical Society of America · doi:10.1121/1.398693 · Cited by 70 · see citing works
Self-citationEcholocation Signals and Echoes in Air
Pye, J. D. (1980) · Springer eBooks · doi:10.1007/978-1-4684-7254-7_14 · Cited by 62 · see citing works
Time and frequency domain processing in the inferior colliculus of echolocating bats
Bodenhamer, R. D., & Pollak, G. D. (1981) · Hearing Research · doi:10.1016/0378-5955(81)90055-1 · Cited by 62 · see citing works
The Determination of Distance by Echolocating Bats
Cahlander, D. A., McCue, J. J. G., & Webster, F. A. (1964) · Nature · doi:10.1038/201544a0 · Cited by 58 · see citing works
The Performance of Echolocation: Acoustic Images Perceived by Echolocating Bats
Simmons, J. A. T., & Grinnell, A. D. (1988) · Springer eBooks · doi:10.1007/978-1-4684-7493-0_40 · Cited by 57 · see citing works
Self-citationR�bsamen, R., Neuweiler, G., & Sripathi, K. (1988) · Journal of Comparative Physiology A · doi:10.1007/bf00612436 · Cited by 50 · see citing works
Acoustic Radiation Patterns for the Echolocating Bats Chilonycteris rubiginosa and Eptesicus fuscus
Simmons, J. A. T. (1969) · The Journal of the Acoustical Society of America · doi:10.1121/1.1911804 · Cited by 48 · see citing works
Self-citationFunctional zones in the auditory cortex of the echolocating bat, Myotis lucifugus
Wong, D., & Shannon, S. L. (1988) · Brain Research · doi:10.1016/0006-8993(88)90176-x · Cited by 48 · see citing works
Discrimination of fluttering targets by the FM-batPipistrellus stenopterus?
Sum, Y. W., & Menne, D. (1988) · Journal of Comparative Physiology A · doi:10.1007/bf00604010 · Cited by 40 · see citing works
Altes, R. A. (1989) · The Journal of the Acoustical Society of America · doi:10.1121/1.397566 · Cited by 32 · see citing works
ECHOLOCATION OF FLYING INSECTS BY THE BAT, CHILONYCTERIS PSILOTIS
Novick, A. (1965) · Biological Bulletin · doi:10.2307/1539557 · Cited by 32 · see citing works
A New Concept of Echo Evaluation in the Auditory System of Bats
Beuter, K. (1980) · Springer eBooks · doi:10.1007/978-1-4684-7254-7_33 · Cited by 30 · see citing works
Acoustic Reflection from Surfaces and Shapes
Neubauer, W. G. (1986) · Cited by 29 · see citing works
Novick, A. C. (1977) · Zenodo (CERN European Organization for Nuclear Research) · doi:10.5281/zenodo.13528576 · Cited by 25 · see citing works
Physiological Mechanisms for Spatial Filtering and Image Enhancement in the Sonar of Bats
Simmons, J. A., & Kick, S. A. (1984) · Annual Review of Physiology · doi:10.1146/annurev.ph.46.030184.003123 · Cited by 23 · see citing works
Self-citationTarget Discrimination and Target Classification in Echolocating Bats
Ostwald, J., Schnitzler, H. U., & Schuller, G. (1988) · Springer eBooks · doi:10.1007/978-1-4684-7493-0_42 · Cited by 22 · see citing works
Localization of Acoustic Signals in the Owl
Konishi, M. (1983) · Springer eBooks · doi:10.1007/978-1-4684-4412-4_10 · Cited by 9 · see citing works
Flagged first, then by citations
Cited by
Sorted by citations
Corollary discharge across the animal kingdom
Crapse, T. B., & Sommer, M. A. (2008) · Nature Reviews Neuroscience · doi:10.1038/nrn2457 · Cited by 789 · see citing works
Human echolocation as a basic form of perception and action
Stroffregen, T. A., & Pittenger, J. B. (1995) · Ecological Psychology · doi:10.1207/s15326969eco0703_2 · Cited by 169 · see citing works
Auditory Dimensions of Acoustic Images in Echolocation
Simmons, J. A., Ferragamo, M. J., Saillant, P. A., Haresign, T., Wotton, J. M., Dear, S. P., & Lee, D. N. (1995) · Springer Handbook of Auditory Research · doi:10.1007/978-1-4612-2556-0_4 · Cited by 60 · see citing works
Self-citation
- Authors
- Simmons, J. A.
- Affiliations
- Brown University (US)
- Corresponding author
- Simmons, James A.
- ORCID
- Simmons, James A.
- Venue
- Cognition
- Publisher
- Elsevier BV
- Published
- 1989-11
- Volume
- 33
- Issue
- 1-2
- Pages
- 155-199
- ISSN
- 0010-0277 (ISSN-L)
- OpenAlex
- W2070598675
- Cited by
- 176
- References
- 66
- Citations by year
- 2026: 1 · 2025: 4 · 2024: 6 · 2023: 3 · 2022: 1 · 2021: 5 · 2020: 7 · 2019: 4 · 2018: 28 · 2017: 1 · 2016: 4 · 2015: 1 · 2014: 3 · 2013: 2 · 2012: 2
- FWCI
- 6.40
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Cognition (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (5k02mh000521-07)
- National Institute of Mental Health (7-KO2-MH00521)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Bat Biology and Ecology Studies
- Life Sciences › Neuroscience › Cognitive Neuroscience › Tactile and Sensory Interactions
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Developmental Biology › Animal Vocal Communication and Behavior
- Keywords
- echolocation, acoustic images, Eptesicus fuscus, glint detection, cross correlation function, range profile, feature fusion, auditory scene analysis, target identification, target interception
- MeSH terms
- Animals, Auditory Pathways, Brain, Brain Mapping, Chiroptera, Echolocation, Orientation, Predatory Behavior
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review