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$ pt 2: sensory input $ ch 2: echolocation in bats $ bat behavior $ decoding the acoustic environment $ hunting bats $ neural mechanisms $ moth responses to predation $ summary PART 2: SENSORY WORLDS #06: ECHOLOCATION IN BATS II
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$ pt 2: sensory input $ ch 2: echolocation in bats $ bat behavior $ decoding the acoustic environment $ hunting bats $ neural mechanisms $ moth responses to predation $ summary PART 2: SENSORY WORLDS #06: ECHOLOCATION IN BATS II
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$ 2 polar views of brain structure-function relationships... $ ball of wool... $ anatomical homogeneity $ cell function diversity $ network & structure... $ anatomical diversity $ cell function homogeneity $ both... BRAIN STRUCTURE – FUNCTION
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$ brain functions determined by... $ experimentation $ proximity & connectivity known inputs & outputs ( eg, olfaction) BRAIN STRUCTURE – FUNCTION
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$ Drosophila brain BRAIN STRUCTURE – FUNCTION
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antenna antennal lobe antennal gl. tract mushroom body lateral horn $ Drosophila olfactory BRAIN STRUCTURE – FUNCTION
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$ human brain BRAIN STRUCTURE – FUNCTION
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$ human auditory NEURAL MECHANISMS
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$ human auditory NEURAL MECHANISMS
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$ bat auditory $ ear drum vibration $ hair displacement $ amplitude $ frequency $ 1° auditory neurons in spiral ganglion $ CNS ) ) ) ) ) ) ) NEURAL MECHANISMS
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$ human auditory ( b ) $ hair cells on basilar membrane = basilar fibers $ shorter @ base, frequency $ longer @ apex, frequency $ excitation... $ inhibition... NEURAL MECHANISMS
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$ ranges of auditory perception 1101001K10K100K FREQUENCY (Hz) HUMAN BAT NEURAL MECHANISMS
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$ bat auditory specializations echolocation ) ) ) ) ) ) ) $ frequency range $ CF bats... acoustic fovea on membrane $ thicker $ longer $ 1° auditory neurons $ more of them NEURAL MECHANISMS
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$ bat auditory specializations echolocation $ frequency range $ CF bats... acoustic fovea on membrane $ thicker $ longer $ 1° auditory neurons $ more of them $ tuned to acoustic fovea NEURAL MECHANISMS
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$ bat auditory specializations echolocation $ frequency range $ CF bats... acoustic fovea on membrane $ thicker $ longer $ 1° auditory neurons $ more of them $ tuned to acoustic fovea ? NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ bat auditory specializations echolocation $ FM bats... sensitivity to emitted call during call $ contract middle ear muscles $ attenuation in nucleus of lateral lemniscus* $ CF bats... remain sensitive to signal during call $ ~ deaf to narrow band of call, membrane sensitivity NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ inferior colliculus encoding FM signal-echo delay $ inferior colliculus neurons fixed spike rate ~ $ call $ echo delay $ not ~ intensity $ frequency tuned $ low threshold NEURAL MECHANISMS
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$ inferior colliculus encoding CF Doppler effect $ # of frequency tuned neurons at acoustic fovea $ target information $ encode insect wing beats NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex auditory cortex sensory interneuron sensory interneuron auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus CF-CFFM-FMDSCF auditory cortex auditory cortex sensory interneuron NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus CF-CFFM-FMDSCF auditory cortex auditory cortex auditory cortex lower centers NEURAL MECHANISMS
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$ auditory cortex lower centers $ individual parameters $ parallel centers NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus auditory cortex NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus CF-CFFM-FMDSCF NEURAL MECHANISMS
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$ mustached bat $ CF & FM components $ pulse & echo $ 4 harmonic frequencies $ H 1 = fundamental $ 4 amplitudes (volumes) $ H 2 > H 3 > H 4 > H 1 $ map onto different cortical areas NEURAL MECHANISMS
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$ mustached bat $ FM-FM area distance & size coding NEURAL MECHANISMS
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$ FM-FM area distance coding $ respond best to FM call + delayed FM echo $ FM-FM neurons delay tuned, in columns $ FM 1 -FM 2, FM 1 -FM 3 & FM 1 -FM 4 zones NEURAL MECHANISMS
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$ FM-FM area size coding $ FM-FM neurons delay & amplitude tuned $ disruption no influence on freq. discrimination NEURAL MECHANISMS
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$ auditory pathway: mustached bat (FM & CF) cochlea auditory nerve cochlear nucleus inferior colliculus nucleus of lateral lemniscus medial geniculate nucleus superior olivary nucleus CF-CFFM-FMDSCF auditory cortex auditory cortex NEURAL MECHANISMS
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$ FM-FM area distance & size coding $ medial geniculate nuc. delays call signal FM-FM $ call + delay coincidence FM-FM auditory cortex auditory cortex NEURAL MECHANISMS
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$ mustached bat $ FM-FM area distance & size coding $ CF-CF area velocity coding NEURAL MECHANISMS
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$ CF-CF area velocity coding (Doppler shift) $ CF-CF neurons Doppler shift tuned, in columns $ CF 1 - CF 2 compares 30 vs 61 kHz tones $ CF 1 - CF 3 compares 30 vs 92 kHz tones NEURAL MECHANISMS
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$ CF-CF area velocity coding (Doppler shift) $ columns map relative velocity coordinates $ -2 +9 m/s $ most 0 +4 m/s (biologically relevant) NEURAL MECHANISMS
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$ mustached bat $ FM-FM area distance & size coding $ CF-CF area velocity coding $ DSCF area Doppler shifted CF 2 NEURAL MECHANISMS
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$ DSCF area Doppler shifted CF 2 signals $ 60.6 62.2 kHz $ signal @ 61 kHz $ neurons tuned for freq. & amplitude of echo $ cortical acoustic fovea $ disruption no influence on delay discrimination NEURAL MECHANISMS
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$ why harmonics ? $ many bats & signals $ H 2 > H 3 > H 4 > H 1 $ fundamental H 1 weak $ only heard by sender $ vs other harmonics $ ~ dual password NEURAL MECHANISMS
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$ why are there any moths left ? $ moths hear rate signals $ early stages (...6) $ moth hears pulse before bat hears echo $ fly in opposite direction MOTH RESPONSES TO PREDATION
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$ why are there any moths left ? $ moths hear rate signals $ late stages (9...) $ moth hears pulse after detection $ moths dive ground & shelter $ erratic flight confusing echo MOTH RESPONSES TO PREDATION
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$ bats echolocate to $ navigate $ locate, identify, track & capture prey (flying insects) $ 2 kinds of signals $ FM, broad band, short duration (< 5 ms) $ CF, narrow band, long duration (5 – 100 ms) $ complex analyses of emitted calls & returning echos SUMMARY
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$ complex analyses of emitted calls & returning echos $ FM call-echo delay distance $ echo amplitude (volume) angular size $ distance + amplitude absolute size $ spatial location... $ binaural comparisons azimuth $ amplitude @ different ear positions horizontal $ CF Doppler shift relative velocity of target SUMMARY
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$ acoustic fovea $ narrow CF range Doppler shift analyses $ overrepresented throughout auditory system $ Doppler shift compensation of call echo in acoustic fovea $ 3 stages of hunting ~ # pulses / s (resolution of target) $ search @ ~ 10 Hz $ approach @ ~ 50 Hz $ terminal @ ~ 100-200 Hz SUMMARY
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$ auditory specializations echolocation $ acoustic fovea Doppler shifted CF echoes $ basilar membrane (cochlea) expanded AF range $ nuc. of lateral lemniscus attenuate FM calls $ inferior colliculus responds to FM echo delays $ medial geniculate nuc. call-echo coincidence detection & relay to auditory cortex $ auditory cortex tonotopic maps SUMMARY
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$ flow diagram $ basilar membrane expanded AF $ inferior colliculus FM echo delays $ med. geniculate nuc. FM coincidence $ auditory cortex tonotopic maps SUMMARY
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$ auditory specializations echolocation $ auditory cortex tonotopic maps $ FM-FM area distance information map onto freq. & delay-sensitive columns of neurons $ CF-CF area velocity information maps onto Doppler shift-sensitive columns of neurons $ DCSF area velocity information maps onto AF range of Doppler shift-sensitive neurons SUMMARY
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$ bat-moth co-evolution... maintains balance $ moths respond to bat signals $ signal rate (approach), moths fly away from signal $ signal rate (terminal), moths either dive or fly irratically $ some emit pulses to jam bat radar SUMMARY
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