Numerical analysis of biosonar beamforming mechanisms and strategies in bats.

Beamforming is critical to the function of most sonar systems. The conspicuous noseleaf and pinna shapes in bats suggest that beamforming mechanisms based on diffraction of the outgoing and incoming ultrasonic waves play a major role in bat biosonar. Numerical methods can be used to investigate the relationships between baffle geometry, acoustic mechanisms, and resulting beampatterns. Key advantages of numerical approaches are: efficient, high-resolution estimation of beampatterns, spatially dense predictions of near-field amplitudes, and the malleability of the underlying shape representations. A numerical approach that combines near-field predictions based on a finite-element formulation for harmonic solutions to the Helmholtz equation with a free-field projection based on the Kirchhoff integral to obtain estimates of the far-field beampattern is reviewed. This method has been used to predict physical beamforming mechanisms such as frequency-dependent beamforming with half-open resonance cavities in the noseleaf of horseshoe bats and beam narrowing through extension of the pinna aperture with skin folds in false vampire bats. The fine structure of biosonar beampatterns is discussed for the case of the Chinese noctule and methods for assessing the spatial information conveyed by beampatterns are demonstrated for the brown long-eared bat.

[1]  A. Grinnell,et al.  Directional sensitivity of echolocation in the horseshoe bat,Rhinolophus ferrumequinum , 2004, Journal of comparative physiology.

[2]  S Carlile,et al.  The auditory periphery of the ferret. I: Directional response properties and the pattern of interaural level differences. , 1990, The Journal of the Acoustical Society of America.

[3]  J. Simmons,et al.  Spectral cues and perception of the vertical position of targets by the big brown bat, Eptesicus fuscus. , 2000, The Journal of the Acoustical Society of America.

[4]  N. Suga,et al.  Directional sensitivity of echolocation system in bats producing frequency-modulated signals. , 1974, The Journal of experimental biology.

[5]  D. P. Skinner,et al.  Broadband target classification using a bionic sonar , 1977 .

[6]  H. Schnitzler,et al.  Echolocation by Insect-Eating Bats , 2001 .

[7]  H. Peremans,et al.  A helical biosonar scanning pattern in the Chinese noctule, Nyctalus plancyi. , 2006, The Journal of the Acoustical Society of America.

[8]  Rolf Müller,et al.  Biosonar-inspired technology: goals, challenges and insights , 2007, Bioinspiration & biomimetics.

[9]  R. Altes Angle estimation and binaural processing in animal echolocation. , 1978, The Journal of the Acoustical Society of America.

[10]  John F. Canny,et al.  A Computational Approach to Edge Detection , 1986, IEEE Transactions on Pattern Analysis and Machine Intelligence.

[11]  Ramani Duraiswami,et al.  Fast head-related transfer function measurement via reciprocity. , 2006, The Journal of the Acoustical Society of America.

[12]  C. Moss,et al.  The bat head-related transfer function reveals binaural cues for sound localization in azimuth and elevation. , 2004, The Journal of the Acoustical Society of America.

[13]  J. C. Middlebrooks,et al.  Individual differences in external-ear transfer functions of cats. , 2000, The Journal of the Acoustical Society of America.

[14]  C. Moss,et al.  The role of the external ear in vertical sound localization in the free flying bat, Eptesicus fuscus. , 2007, The Journal of the Acoustical Society of America.

[15]  J. Flint A biomimetic antenna in the shape of a bat's ear , 2006, IEEE Antennas and Wireless Propagation Letters.

[16]  J A Simmons,et al.  Echolocation in bats: the external ear and perception of the vertical positions of targets. , 1982, Science.

[17]  R. Rübsamen,et al.  Foraging behaviour and echolocation in the rufous horseshoe bat (Rhinolophus rouxi) of Sri Lanka , 2004, Behavioral Ecology and Sociobiology.

[18]  Omar M. Ramahi,et al.  Near- and far-field calculations in FDTD simulations using Kirchhoff surface integral representation , 1997 .

[19]  Rolf Müller,et al.  Sound-diffracting flap in the ear of a bat generates spatial information. , 2008, Physical review letters.

[20]  P. Schlegel,et al.  Frequency sensitivity and directional hearing in the gleaning bat,Plecotus auritus (Linnaeus 1758) , 2004, Journal of Comparative Physiology A.

[21]  G. Westheimer Center-surround antagonism in spatial vision: Retinal or cortical locus? , 2004, Vision Research.

[22]  B.D. Van Veen,et al.  Beamforming: a versatile approach to spatial filtering , 1988, IEEE ASSP Magazine.

[23]  R. Müller,et al.  Noseleaf furrows in a horseshoe bat act as resonance cavities shaping the biosonar beam. , 2006, Physical review letters.

[24]  C. Rajakumar,et al.  The Boundary Element Method: Applications in Sound and Vibration , 2004 .

[25]  Rolf Müller,et al.  Pinna-rim skin folds narrow the sonar beam in the lesser false vampire bat (Megaderma spasma). , 2009, The Journal of the Acoustical Society of America.

[26]  Klaus Hartung,et al.  Head-related transfer functions of the barn owl: measurement and neural responses , 1998, Hearing Research.

[27]  Rolf Müller,et al.  Numerical study of the effect of the noseleaf on biosonar beamforming in a horseshoe bat. , 2007, Physical review. E, Statistical, nonlinear, and soft matter physics.

[28]  Shiva R. Sinha,et al.  Neurobiology of echolocation in bats , 2003, Current Opinion in Neurobiology.

[29]  F. Ihlenburg Finite Element Analysis of Acoustic Scattering , 1998 .

[30]  Richard Barrett,et al.  Templates for the Solution of Linear Systems: Building Blocks for Iterative Methods , 1994, Other Titles in Applied Mathematics.

[31]  Hans-Ulrich Schnitzler,et al.  Classification of insects by echolocating greater horseshoe bats , 1990, Journal of Comparative Physiology A.

[32]  Rolf Müller,et al.  A numerical study of the role of the tragus in the big brown bat. , 2004, The Journal of the Acoustical Society of America.

[33]  W. Sellers,et al.  The aerodynamics of big ears in the brown long-eared bat Plecotus auritus , 2008 .

[34]  Dean A. Waters,et al.  Echolocation call design and limits on prey size: a case study using the aerial-hawking bat Nyctalus leisleri , 1995, Behavioral Ecology and Sociobiology.

[35]  O. von Estorff,et al.  Improved conditioning of infinite elements for exterior acoustics , 2003 .

[36]  J. Reijniers,et al.  Simulated head related transfer function of the phyllostomid bat Phyllostomus discolor. , 2008, The Journal of the Acoustical Society of America.

[37]  Brian F. G. Katz,et al.  Round Robin Comparison of HRTF Simulation Systems: Preliminary Results , 2007 .

[38]  R. Kuc Sensorimotor model of bat echolocation and prey capture. , 1994, The Journal of the Acoustical Society of America.

[39]  J. Ben-Arie,et al.  Optimal head related transfer functions for hearing and monaural localization in elevation: a signal processing design perspective , 1996, IEEE Transactions on Biomedical Engineering.

[40]  M. Fenton,et al.  What ears do for bats: a comparative study of pinna sound pressure transformation in chiroptera. , 1993, The Journal of experimental biology.

[41]  Rolf Müller,et al.  Acoustic effects accurately predict an extreme case of biological morphology. , 2009, Physical review letters.

[42]  Wolfgang J. R. Hoefer,et al.  The Transmission-Line Matrix Method--Theory and Applications , 1985 .

[43]  Uwe Firzlaff,et al.  Spectral directionality of the external ear of the lesser spear-nosed bat, Phyllostomus discolor , 2003, Hearing Research.

[44]  N Suga,et al.  Biosonar and neural computation in bats. , 1990, Scientific American.

[45]  Rolf Müller,et al.  Frequency-swept directivity lobes—An emerging functional principle of biosonar beamforming , 2007 .

[46]  Henry E. Bass,et al.  Atmospheric absorption of sound: Update , 1990 .

[47]  R A Altes,et al.  Computer Derivation of Some Dolphin Echolocation Signals , 1971, Science.