Echo delay versus spectral cues for temporal hyperacuity in the big brown bat, Eptesicus fuscus

Big brown bats can discriminate between echoes that alternate in delay (jitter) by as little as 10–15 ns and echoes that are stationary in delay. This delay hyperacuity seems so extreme that it has been rejected in favor of an explanation in terms of artifacts in echoes, most likely spectral in nature, that presumably are correlated with delay. Using different combinations of digital, analog, and cable delays, we dissociated the overall delay of jittering echoes from the size of the analog component of delay, which alone is presumed to determine the strength of the apparatus artifact. The bats' performance remains invariant with respect to the overall delay of the jittering echoes, not with respect to the amount of analog delay. This result is not consistent with the possible use of delay-related artifacts produced by the analog delay devices. Moreover, both electronic and acoustic measurements disclose no spectral cues or impedance-mismatch reflections in delayed signals, just time-delays. The absence of artifacts from the apparatus and the failure of overlap and interference from reverberation to account for the 10-ns result means that closing the gap between the level of temporal accuracy plausibly explained from physiology and the level observed in behavior may require a better understanding of the physiology.

[1]  Cynthia F. Moss,et al.  Convergence of temporal and spectral information into acoustic images of complex sonar targets perceived by the echolocating bat, Eptesicus fuscus , 1990, Journal of Comparative Physiology A.

[2]  James A. Simmons,et al.  Auditory Dimensions of Acoustic Images in Echolocation , 1995 .

[3]  Nathan Intrator,et al.  Time-Frequency Computational Model for Echo-Delay Resolution in Sonar Images of the Big Brown Bat, Eptesicus Fuscus , 2005 .

[4]  J A Simmons,et al.  Perception of echo phase information in bat sonar. , 1979, Science.

[5]  H. Schnitzler,et al.  Range estimation by echolocation in the bat Eptesicus fuscus: trading of phase versus time cues. , 1989, The Journal of the Acoustical Society of America.

[6]  C F Moss,et al.  Echo-delay resolution in sonar images of the big brown bat, Eptesicus fuscus. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[7]  K. Beedholm,et al.  Bat sonar: an alternative interpretation of the 10-ns jitter result , 1998, Journal of Comparative Physiology A.

[8]  G. Pollak,et al.  Some comments on the proposed perception of phase and nanosecond time disparities by echolocating bats , 1993, Journal of Comparative Physiology A.

[9]  J. A. Simmons,et al.  Evidence for perception of fine echo delay and phase by the FM bat, Eptesicus fuscus , 1993, Journal of Comparative Physiology A.

[10]  W. M. Masters,et al.  Detection of jitter in intertarget spacing by the big brown bat Eptesicus fuscus , 1997, Journal of Comparative Physiology A.

[11]  Walter Heiligenberg,et al.  Neural Nets in Electric Fish , 1991 .

[12]  Alan D. Grinnell,et al.  Hearing in Bats: An Overview , 1995 .

[13]  R A Altes,et al.  Ubiquity of hyperacuity. , 1989, The Journal of the Acoustical Society of America.

[14]  George D. Pollak,et al.  Characteristics of phasic on neurons in inferior colliculus of unanesthetized bats with observations relating to mechanisms for echo ranging , 1977 .

[15]  Gareth Jones,et al.  The influence of flight speed on the ranging performance of bats using frequency modulated echolocation pulses. , 2001, The Journal of the Acoustical Society of America.

[16]  Robert F. Port,et al.  Neural Representation of Temporal Patterns , 1995, Springer US.

[17]  Hans-Ulrich Schnitzler,et al.  Range Determination by Measuring Time Delays in Echolocating Bats , 1985 .

[18]  M. Konishi,et al.  Neuronal and behavioral sensitivity to binaural time differences in the owl , 1981, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[19]  C. A. Long,et al.  Biology of Bats , 1972 .

[20]  Cynthia F. Moss,et al.  Behavioral Studies of Auditory Information Processing , 1995 .

[21]  Alan D. Grinnell,et al.  The Performance of Echolocation: Acoustic Images Perceived by Echolocating Bats , 1988 .

[22]  S Shamma,et al.  The case of the missing pitch templates: how harmonic templates emerge in the early auditory system. , 2000, The Journal of the Acoustical Society of America.

[23]  C F Moss,et al.  Acoustic image representation of a point target in the bat Eptesicus fuscus: evidence for sensitivity to echo phase in bat sonar. , 1993, The Journal of the Acoustical Society of America.

[24]  James A. Simmons,et al.  The Processing of Sonar Echoes by Bats , 1980 .

[25]  S. Stevenson,et al.  Discrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: The shape of target images in echolocation , 1990, Journal of Comparative Physiology A.

[26]  J. Simmons,et al.  Echolocation: discrimination of targets by the bat, Eptesicus fuscus. , 1971, The Journal of experimental zoology.

[27]  H. Schnitzler,et al.  Accuracy of target ranging in echolocating bats: acoustic information processing , 1989, Journal of Comparative Physiology A.

[28]  Nicola Neretti,et al.  Evaluation of an auditory model for echo delay accuracy in wideband biosonar. , 2003, The Journal of the Acoustical Society of America.

[29]  T. F. Weiss,et al.  A comparison of synchronization filters in different auditory receptor organs , 1988, Hearing Research.

[30]  D. Griffin,et al.  Listening in the Dark , 1959 .

[31]  D. Hartley,et al.  The sound emission pattern of the echolocating bat, Eptesicus fuscus , 1989 .

[32]  Nicola Neretti,et al.  Time-frequency model for echo-delay resolution in wideband biosonar. , 2003, The Journal of the Acoustical Society of America.

[33]  John H. Casseday,et al.  Mechanisms for Analysis of Auditory Temporal Patterns in the Brainstem of Echolocating Bats , 1995 .

[34]  James A. Simmons,et al.  Auditory Computations for Biosonar Target Imaging in Bats , 1996 .

[35]  C. Köppl Phase Locking to High Frequencies in the Auditory Nerve and Cochlear Nucleus Magnocellularis of the Barn Owl, Tyto alba , 1997, The Journal of Neuroscience.