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Neuroethology: Sound in insects http://biolpc22.york.ac.uk/404/ References Young, D (1989) Nerve cells and animal behaviour CUP [1st edition, chapter 7] Also, chapters in: Carew Behavioral neurobiology Zupanc Behavioral neurobiology papers are on the web http://biolpc22.york.ac.uk/404/cjhe/ cricket_singing/insect.htm What is neuroethology? approach to analyse neural function in terms of evolutionary advantage Main points of lecture Sound transmission is inefficient Insects use sound in a major way Overcoming physical obstacles Modifying wings for sound production the environment for sound production cuticle to hear Private communication About sound sound is a wave of particle compression and rarefaction energy (or power) is transferred from sender to hearer sound gets quieter further away sound has to be discriminated according to species, context and from random noise Sound properties c f l speed c (330 m s-1) wavelength l (m) frequency f (Hz, cycles/s) for 330Hz, wavelength 1m for 3.3kHz, wavelength 0.1m for 33kHz, wavelength 0.01m = 10mm Power transfer - i First problem for an insect for air/water interface about 0.02% Power transfer - ii Usually other insect far away With distance, power decreases more bad news for insects! 1 2 d Power transfer - iii Usually insects small sound emitted is dependent on ratio of insect size / wavelength of sound Power transfer - iv if you are smaller than 0.2 * l more bad news! Insects use sound in a major way Mantids* Grasshoppers/crickets* bugs* beetles* diptera (true flies)* lacewings moths Overcoming physical obstacles Resonance Baffle Horn Sound production Sound reception Resonance Mass and a spring; resonant frequency 1 fo 2 s m s is stiffness, m mass below fo in phase; at fo 90o; above fo out of phase Gryllus Field cricket nice loud song, carries a long way attracts females & males territorial Gryllus EMGs sound sonogram oscillogram Females… … walk to the loudest sound Sound made by wing plectrum add wax to harp frequency reduced resonator affected remove teeth Gryllus campestris sound output 60 mW muscle work 2 mW efficiency 3% Bush crickets Higher frequency damped oscillation Baffle Bush cricket Oecanthus power * 12 Gryllotalpa mole cricket Horn shaped burrow Gryllotalpa sound output 1 mW muscle work 5 mW efficiency 20% Summary so far Modified wings for sound production Modifying the environment for sound production Hearing locust = grasshopper ears Tympanum structure Tympanum structure side view Muller’s organ has sensory endings in it Tympanum (ear drum) taut membrane top view Movement it’s the relative movement which will stretch the axons Movement peaks at 3 & 6kHz indicate resonance resonance also shown by phase plot folded body not in phase with membrane even when at 4kHz amplitude is same relative movement needed to stretch axons of Muller’s organ Cricket hearing acoustic trachea acts like horn for 5kHz sound Summary so far Modified wings for sound production Modifying the environment for sound production Modified cuticle to hear Drosophila courtship Male has black tip Olfaction Singing Detecting vibration near field sound movement detector aristae - bristles on antennae Private communication Sound movement in dB since sound is measured on a log scale, 80 << 95 Summary The neuroethological approach relates the function of the nervous system to its evolutionary advantage Many insects use sound produce it quite efficiently hear it effectively