Sound Production. Phonotaxis in crickets. What is sound? Recognition and Localization. What happens over time at a single point in space?
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1 Behaviour Sound Production Phonotaxis in crickets Recognition and Localization scraper close scraper file open Males open and close wings rhythmically. On each closing stroke, scraper contacts file causing vibration of the wing which generates sound. Crickets produce a pure frequency. NROC34 212:2a 1 NROC34 212:2a 2 file What happens over time at a single point in space? It s a wave... What is sound? in a fluid medium. Pressure or Displacement Animations courtesy of Dr. Dan Russell, Kettering University NROC34 212:2a 3 NROC34 212:2a 4 Time 1
2 Basic idea of an ear -something has to be moved by the sound waves Frequency Displacement Pressure NROC34 212:2a 5 Period = time for one complete cycle of wave. Frequency = 1/Period (# waves per unit time) NROC34 212:2a 6 Amplitude Subjective qualities Amplitude= height of waves different frequencies (pitch) different s (loudness) How much does the pressure increase & decrease over the course of a wave? NROC34 212:2a 7 NROC34 212:2a 8 2
3 The mysterious Decibel Units Pressure (Pa) db SPL decibel (db) = 2 log (A 1 / A 1 ) The same two sounds shown in units of absolute pressure and db SPL. db SPL = 2 log (A/2μPa) NROC34 212:2a 9 Frequency cycles/sec Hertz (Hz) Kilohertz (khz) Audio 2-2 Hz Ultrasound >2 khz Amplitude magnitude of pressure variation measured in db SPL increase of 6 db means doubles, no matter where you are on the scale NROC34 212:2a 1 More on frequency More on frequency relative time (ms) relative time (ms) frequency (Hz) Fourier analysis - two sounds NROC34 212:2a 11 frequency (Hz) Fourier analysis: sum of two sounds NROC34 212:2a 12 3
4 Not all frequencies are pure Masking by noise relative time (ms) frequency (Hz) Sum of two sounds NROC34 212:2a NROC34 212:2a Mechanisms of hearing a neuron with its dendrite stuck to a stiff rod movement of the rod excites the cell general purpose sensory element, insects have them all over, often in groups (chordotonal organs), and these have been modified in many taxa to form ears Scolopidia NROC34 212:2a 15 Mechanisms of hearing Tympanal Organs sound waves eardrum deflected back and forth with pressure oscillations movements of eardrum stimulate neurons in chordotonal organ NROC34 212:2a 16 4
5 Where different bugs put them 1) Lepidoptera: Sphingoidea 2) Orthoptera: Ensifera 3) Diptera: Tachinidae, Sarcophagidae Coleoptera: Scarabaidae 4) Mantodea: Mantidae 5) Lepidoptera: Geometridae, Pyraloidea 6) Orthoptera: Acrididae, Pneumoridae 7) Hemiptera: Cidadidae 8) Lepidoptera: Noctuidae 9) Hemiptera: Corixidae 1) Neuroptera: Chysopidae ear Cricket auditory system acoustic trachea brain prothoracic ganglion NROC34 212:2a Fullard & Yack NROC34 212:2a 18 Behaviour A syllable, or pulse, is the sound produced in one single cycle of wing movement (on the closing stroke). Rhythmic movement of wings leads to a series of regularly spaced pulses. A few pulses in a row is a chirp. A continuous train is a trill. Song temporal pattern 1 ms 5 ms syllable or pulse duration syllable period interpulse interval no. syllables in chirp chirp duration NROC34 212:2a 19 Behaviour Female cricket follows a meandering path towards a male. Male must be same species as female and must be singing. Female is guided by sound. Cricket songs are species-specific. Phonotaxis Phonotaxis: oriented movement in response to sound. NROC34 212:2a 2 5
6 Behaviour Phonotaxis female cricket must be able to: recognize the song as that of a conspecific localize the source frequency, temporal and directional information from the acoustic stimulus (song) Behaviour Kramer Treadmill switch speaker NROC34 212:2a 21 NROC34 212:2a 22 Behaviour Open-loop Behaviour Closed Loop: animal s response generates feedback that can update ongoing behaviour. Open Loop: stimulus conditions remain unchanged no matter what the animal does. Behaviour Phonotaxis: preference for natural syllable rate 1 8 response sensory feedback response sensory feedback NROC34 212:2a 23 tracking behaviour (percent total time) NROC34 212:2a syllable rate (number per second) 24 6
7 Anatomy Cricket auditory system Anatomy Insect tracheal system ear acoustic trachea Gas exchange by direct diffusion to/from tissue. System of air tubes connecting spiracles (openings body surface) with all internal tissues. Typically one pair of spiracles per body segment. brain prothoracic ganglion NROC34 212:2a 25 NROC34 212:2a 26 Anatomy Acoustic trachea Hearing Direction: pressure difference spiracle wavelength ( ) = cf physical distance between one wave and the next d 2 -d 1 d 1 d 2 tympanum spiracle d 2 -d 1 /2 tympanum (eardrum) acoustic trachea NROC34 212:2a 27 frequency-dependent NROC34 212:2a 28 7
8 Hearing Peripheral directionality Hearing Peripheral frequency tuning Eardrum will only vibrate within a range of frequencies. Auditory receptors will only respond to sounds of certain frequencies. 27?? 9 left ear 18 right ear Response of the ear varies with the location of the source. NROC34 212:2a 29 receptors eardrum NROC34 212:2a cricket ear 3 Peripheral Filtering Properties of peripheral sensory apparatus limit sensitivity to relevant range of stimuli. sound frequency light wavelength Sensitivity of peripheral sensory apparatus varies according to some parameter of interest. auditory directionality polarization sensitivity (maybe a later topic?) More generally... Neural Filtering Properties of neurons that limit their responses to certain stimuli, or classes of stimuli. Networks of neurons that are selective for certain stimuli (feature detectors). NROC34 212:2a 31 NROC34 212:2a 32 8
9 Hearing Central auditory processing prothoracic ganglion - first stage of central processing several identified neurons with known functions (others whose functions remain mysterious) Hearing Prothoracic auditory interneurons omega neuron 1 ascending neuron 1 NROC34 212:2a 33 ascending neuron 2 (next week) NROC34 212:2a 34 Localization Omega neurons Localization Reciprocal Inhibition input: from ear output stimulate ipsilateral ear output: inhibition input stimulate contralateral ear The two ON1 s inhibit each other. Peripheral directionality means that input to one of them will be stronger if sound source is not directly ahead. More strongly activated ON1 will inhibit the other, and directional signal is amplified. ON1 also inhibits contralateral AN1 (Schildberger and Hörner paper). NROC34 212:2a 35 NROC34 212:2a 36 9
10 Recognition Prothoracic interneurons Recognition Brain Neurons ON1 input output AN1 output AN2 (next week) Auditory pathway through the brain: AN1 - BNC1 - BNC2... NROC34 212:2a No evidence of temporal filtering in prothorax. 37 NROC34 212:2a 38 Recognition Brain Neuron Responses BNC2 BNC1 AN1 latency chirp Increasing latency at each stage (evidence of sequential connectivity). Decreasing accuracy of temporal pattern. spikes per chirp syllable interval (ms) BNC2 neurons are selctive for syllable repetition rate. NROC34 212:2a 39 BNC2 BNC1 AN1 Recognition Brain Neurons Two classes: BNC1 & BNC2 based on connectivity BNC1 receive input from AN1 BNC2 receive input from BNC1 Several subclasses of each based on response properties. NROC34 212:2a 4 1
11 Recognition Temporal Filtering by brain neurons Recognition A Model for Recognition prothorax low-pass brain low-pass chirp band-pass relative response (% max) high-pass band-pass ON1, AN1 high-pass BNC1 & 2 BNC2 syllable rate (#/sec) NROC34 212:2a 41 NROC34 212:2a 42 11
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