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Perceptual Strategies in Active and Passive Hearing of Neotropical Bats

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Perceptual Strategies in Active and Passive Hearing of Neotropical Bats Perceptual strategies in active and passive hearing of neotropical bats Holger R. Goerlitz March 2008 München Perceptual strategies in active and passive hearing of neotropical bats Holger R. Goerlitz Dissertation an der Fakultät für Biologie der Ludwig–Maximilians–Universität München zur Erlangung des Grades eines Doktors der Naturwissenschaften vorgelegt von Holger R. Goerlitz aus Stuttgart München, den 27.03.2008 Erstgutachter: PD. Dr. Lutz Wiegrebe Zweitgutachter: Prof. Dr. Benedikt Grothe Tag der mündlichen Prüfung: 22.10.2008 Contents List of figures and tables ix List of abbreviations xi Zusammenfassung xiii Summary xv 1 General introduction3 1.1 Bats........................................3 1.2 Hearing and echolocation............................6 1.3 Perception and action in the context of sensory physiology and sensory ecology 13 2 Comparing Passive and Active Hearing: Spectral Analysis of Transient Sounds 19 2.1 Introduction.................................... 19 2.2 Methods...................................... 21 2.2.1 Phyllostomus discolor .......................... 21 2.2.2 Animal housing and training procedure................ 22 2.2.3 Experiment 1: passive hearing..................... 22 2.2.4 Experiment 2: echolocation....................... 25 2.2.5 Echolocation call recording and analysis................ 26 2.2.6 Analysis and statistics.......................... 27 2.3 Results....................................... 27 2.3.1 Training performance.......................... 27 2.3.2 Experiment 1: perceptual compensation during passive hearing... 29 2.3.3 Experiment 2: spectral processing during echolocation........ 30 2.3.4 Effect of the roving level......................... 31 2.3.5 Echolocation calls............................ 32 2.4 Discussion..................................... 32 2.4.1 Experiment 1: perceptual compensation during passive hearing... 34 2.4.2 Experiment 2: spectral processing during echolocation........ 34 2.4.3 Effect of the roving level......................... 35 2.4.4 Echolocation calls............................ 36 2.4.5 Conclusion................................ 37 Acknowledgments.................................... 37 Appendix: Transponder................................ 38 3 Detection of Jittering Real Targets by Passive Listening and Echolocation 41 3.1 Introduction.................................... 41 3.1.1 Experiments on temporal accuracy................... 42 3.1.2 Objections to the jitter experiments.................. 44 3.1.3 Aim of this study............................. 45 3.2 Methods...................................... 46 3.2.1 Glossophaga soricina ........................... 46 3.2.2 Housing.................................. 47 3.2.3 Experimental setup............................ 47 3.2.4 Description of the stimuli........................ 49 3.2.5 Experimental paradigm......................... 50 3.2.6 Call recordings and simulation of the perceived temporal jitter... 52 3.2.7 Statistical methods............................ 54 3.3 Results....................................... 55 3.3.1 Echo-acoustic detection of temporal jitter in Glosso2 and Glosso3.. 55 3.3.2 Detection of jitter in Glosso1...................... 55 3.3.3 The use of passive acoustic cues.................... 57 3.3.4 Simulation of the perceived temporal jitter.............. 60 3.4 Discussion..................................... 62 3.4.1 Potential cues in the current jitter detection experiment....... 63 3.4.2 Discussion of the measured thresholds................. 66 3.4.3 The use of echolocation versus passive listening............ 68 3.4.4 Conclusion................................ 69 Acknowledgments.................................... 69 4 Orientation Response of Wild Free-Flying Bats to Real and Virtual Objects 73 4.1 Introduction.................................... 73 4.2 Methods...................................... 77 4.2.1 Study site and animals.......................... 77 4.2.2 Playback and video-recording setup.................. 77 4.2.3 Test and control stimuli......................... 80 4.2.4 Experimental procedure and stimulus presentation.......... 81 4.2.5 Video analysis and flight path extraction............... 82 4.2.6 Evaluation of flyways and statistics................... 83 4.3 Results....................................... 84 4.3.1 Behavioural observations........................ 84 4.3.2 Control recordings of undisturbed flight paths............. 85 4.3.3 Orientation response to real objects.................. 88 4.3.4 Orientation response to virtual objects................. 92 4.4 Discussion..................................... 97 4.4.1 Real objects................................ 97 4.4.2 Virtual objects.............................. 99 4.4.3 Methodological discussion........................ 101 4.4.4 Conclusion and outlook......................... 102 Acknowledgments.................................... 103 Contents 5 General discussion 105 5.1 Comparison of methods............................. 105 5.2 Sensory physiology: Spectral and temporal processing............ 107 5.2.1 Auditory receiver models in echolocation................ 108 5.3 The context of sensory ecology......................... 110 5.4 The end. ..................................... 110 References 111 Acknowledgments 133 Curriculum vitae 135 List of publications 137 Ehrenwörtliche Erklärung 139 vii List of figures and tables List of figures 1.1 The impulse response and the generation of echoes.............. 14 1.2 Perception-action-loop.............................. 15 2.1 Echolocation call of Phyllostomus discolor ................... 21 2.2 Y-shaped training platform........................... 23 2.3 Amplitude spectra of the filters and of example stimuli............ 24 2.4 Block diagram of Experiment 1 (passive hearing)............... 25 2.5 Block diagram of Experiment 1 (echolocation)................. 26 2.6 Learning curves of Bats 4 and 5......................... 28 2.7 Experiment 1 (passive hearing): Classification of filtered impulses...... 29 2.8 Experiment 2 (echolocation): Classification of filtered calls.......... 30 2.9 Experiment 2 (echolocation): Influence of the roving level.......... 31 2.10 Call parameters of Phyllostomus discolor in Experiment 2 (echolocation).. 33 2.11 Calculated influences on the frequency centroid................ 36 3.1 Echolocation calls of Glossophaga soricina ................... 46 3.2 Experimental chamber of the jitter experiment................ 47 3.3 Sketch of the setup and the feeders....................... 48 3.4 Impulse response of the vibrating speaker membrane............. 50 3.5 Sketch of the control-setup........................... 52 3.6 Perceived jitter depending on vibration frequency and pulse interval.... 53 3.7 Temporal jitter detection functions....................... 55 3.8 Jitter detection function for Glosso1...................... 56 3.9 Initial test, control and retraining performance of Glosso2.......... 58 3.10 Sound recordings of the vibrating membranes................. 59 3.11 Training and control performance of Glosso3................. 60 3.12 Call sequences of Glossophaga soricina ..................... 61 3.13 Simulated perceived jitter at threshold..................... 63 3.14 Bat-pollinated flowers.............................. 67 4.1 Hypothetical flight paths around the virtual object.............. 76 4.2 Tamana Cave, Trinidad............................. 78 4.3 Block diagram of the experimental setup.................... 79 4.4 Camera-only control: Example flight paths without any object....... 86 List of tables 4.5 Camera-only control: Flight paths and flyways without any object..... 87 4.6 Real object: Example flight paths........................ 89 4.7 Real object: Flight paths............................ 90 4.8 Real object: Flyways............................... 91 4.9 Real object: Reaction distances......................... 92 4.10 Virtual objects: Interpolated flight paths, overview of December 09th .... 93 4.11 Virtual objects: Interpolated flight paths, overview of December 10th .... 94 4.12 Virtual objects: Interpolated flight paths, overview December of 15th .... 95 4.13 Virtual objects: Flyways of December 09th ................... 96 4.14 Virtual objects: Flyways of December 10th ................... 97 4.15 Virtual objects: Flyways of December 15th ................... 98 List of tables 1.1 Systematics of the studied species........................6 3.1 Published distance discrimination thresholds................. 42 3.2 Published jitter detection thresholds...................... 43 3.3 Pulse intervals of call sequences......................... 62 4.1 Impulse response parameters.......................... 81 x List of abbreviations 2-AFC two-alternative, forced-choice CF constant frequency dB FS decibel Full Scale dB/oct. decibel per octave dB SPL decibel Sound Pressure Level FM frequency-modulated IR impulse response log10M4 decadic logarithm of the 4. moment PI pulse interval pp peak-to-peak RMS root mean square SCAT spectrogram correlation and transformation (a sonar receiver model) SEM standard error of the mean std standard deviation Zusammenfassung Das auditorische System wertet spektrale und zeitliche Schalleigenschaften, wie z.B. Fre- quenzgehalt und Zeitverlauf aus, um eine interne Repräsentation der Umgebung zu erhalten und auditorisch gesteuertes Verhalten zu kontrollieren. Mit echoortenden Fledermäusen als Modellsystem habe ich die spektrale und zeitliche Verarbeitung während der Echoortung untersucht und im Vergleich zum passiven Hören
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