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746 Lecture 2. Echolocation in Bats. Aim. Outline properties of sound Hunting behaviour of bats Types of Echolocation sounds Specialisation of Ear CNS Auditory behaviour of moths. Properties of sound. Sound is wave of rarefaction and compression has speed 330m/s, c = f * l
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746 Lecture 2 Echolocation in Bats
Aim • Outline properties of sound • Hunting behaviour of bats • Types of Echolocation sounds • Specialisation of • Ear • CNS • Auditory behaviour of moths
Properties of sound • Sound is wave of rarefaction and compression • has speed 330m/s, c = f * l • wavelength l - determines whether objects will reflect or diffract sound • frequency f • intensity - • measured in dB
Harmonics • multiples of frequency • usually less intense
Hunting behaviour of bats Taphozous Pipistrellus Megaderma Hipposideros
Echolocation sounds • all bats use “ultrasonic” sounds • CF - • constant frequency • long tone, often with some harmonics • velocity • FM - • frequency modulated • short burst of sound • range determination
CF in free air
FM • near ground or vegetation
CF-FM • CF then droop • depends on place
What will bat hear? • itself ? • reflection ? • quieter • more variable? • Doppler shift in frequency ?
Doppler shift (i) • emitted sound
Doppler shift (ii) • Reflected sound sometimes in phase and sometimes out of phase in out
Doppler shift (iii) • If reflected and emitted sound have similar intensity, Doppler echo will generate beats • Production of new frequency from old!
Doppler summary • New frequency – depends on ratio of outgoing sound and incoming sound • Incoming sound is reflected off ground/trees • Difference in frequency therefore tells how fast the bat is flying • fnew = fout (v + s)/v • v speed of sound • s speed of bat
Echoes • From stationary insect • head on- symmetrical • sideways on asymmetric Echoes from fixed Tipula
Summary so far • Ultrasonic sound • CF FM • habitat dependent • Echoes return information • moving insects • time to return • frequency spectrum • shifted • broadened
Behaviour to physiology • Specialisation of • Ear • CNS
Bat ear (i) • Large pinnae • directional sensitivity • extra gain • Tragus • elevation
Bat ears (ii) • middle ear muscles • reduce sensitivity while emitting? flying bat
Bat ear (iii) • More of cochlea tuned to high frequencies than in other mammals
Tuning curve • auditory nerve • tuned to “best” frequency of emitted CF • actually to just above (Why?)
CF lowered in flight • Doppler shift as fly towards object raises return sound frequency
CNS outline AC IC CN
CNS • Auditory cortex neurons sensitive to pairs of stimuli • load/quiet • delay time crucial • time map
mechanisms of delay • coincidence detection • inhibition of sound • delay line • slow axon • synapse • control with vocalisation • inferior colliculus neurons respond only 30/40ms after vocalisation
Summary - audition • Ear and CNS both highly specialised • more sensitive to ultrasonic frequencies • achieve increase in sensitivity to echo • respond to pairs of stimuli
Moth Auditory system • 2 axons in ear low and high threshold
Behaviour • low threshold - fly fast • high threshold - stop flying and fall • ?like a leaf • Emit clicks - • jam bat sonar - phantom echo returns at wrong time? • warning of unpalatability? • moths (Euproctis) emit clicks in mimicry of distasteful moths
Conclusion • co-evolution of bats and moths • defence reactions • escape • auditory camouflage • auditory