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Explore the physical attributes of sound, auditory sensory transduction, and neural pathways in the human ear. Learn about pitch, frequency coding, and the mysteries of auditory masking in this comprehensive overview of the auditory system.
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Psy280: Perception Prof. Anderson Department of Psychology Audition 1 & 2
Hearing: What’s it good for? • Remote sensing • Not restricted like visual field • Can sense object not visible
Hearing: The sound of silence • A tree in the forest • Physical signal but no perception • One hand clapping • No physical signal, no perception • Separate physical quantity from perceptual quality • Sound is the perceptual correlate of the physical changes in air pressure • Or water pressure when under water • John Cage’s 4:33 No. 2, 1962
What are the physical attributes associated with sound? • Loudness • Amplitude or height of pressure wave • Pitch • Frequency of times per second (Hz) a pressure wave repeats itself
What is sound quality? • Pure tones • Single frequency (f) • Rarely exist in real world • Complex tones • More than one f • Due to resonance • Air pressure causes reverberations • E.g., tuning forks • E.g., Plucking the A string on a guitar • Fundamental frequency 440 Hz (cycles/s) • Harmonics • Reverberations at multiples of the fundamental • E.g., 880, 1320 • Creates fullness of complex sounds • Timbre is the relative amplification of harmonics
The human ear • Outer ear • Focusing of sound • Resonance amplifies 2000-5000 Hz range • Converts from air to mechanical vibration • Middle ear • Amplification • Fluid denser than air • Focus vibrations onto stapes/oval window • Increased leverage from ossicles • Inner ear • Sensory transduction • Physical to neural energy • Fluid pressure changes • Bending of hair cells
Auditory sensory transduction: The inner ear • Cochlea • Coiled and liquid filled • 3 layers • Cochlear partition • Contains organ of corti • Organ of corti • Cilia (hair) cells • Between basilar and tectorial membranes • Transduction • Movement of cilia between membranes
Auditory transduction • Bending—>physical energy • Converted to neural signals • Bend one direction —> depolarization • More likely to fire AP • Other direction —> hyperpolarization • Less likely to fire AP
Audition: What and where • What is it? • *Pitch • Identification • Surprisingly, little is known beyond speech • Where is it? • *location
What: Pitch • How does neural firing signal different pitches? • 1) Timing codes • 2) Place codes
Pitch: Temporal coding • Idea: Diff f’s signaled by rate of neuronal firing • Hair cell response • Bend one direction —> depolarization • Other direction —> hyperpolarization • Result? • Bursting pattern of neural response related to frequency of oscillation
Problems with temporal coding • Problem: A single neuron can’t fire at the rate necessary to represent higher f tones • E.g., 1000-20,000 Hz (i.e., 1000-20000 per second) • Max neuron firing rate: 500-800 per second • Solution: volley principle • No single neuron represents f • Coding across many neurons with staggered firing rates • Evidence: Phase locking • Diff neurons respond to diff peaks • Not every peak • Pool across multiple neurons to represent high f’s
Pitch: Place coding Owl brainstem • Related to doctrine of specific nerve energies • What is pitch? • Activation of different places in auditory system • Frequency specific • Tonotopy • Cochlear • Brainstem • Cortical • Stimulate these regions • Should result in pitch perception Human auditory cortex
Place coding starts in cochlea • Von Bekesy studied basilar membrane in cadavers • Base more narrow and stiffer • Apex wider and more flexible • Observed traveling waves • Diff frequencies (f) result in waves w/ diff envelopes • Higher f: Peak closer to base • Lower f: Peak closer to apex • Thus, f related to “place” where peak fluctuation occurs
Frequency tuning: Neural place coding • Tonotopic arrangement of hair cell nerves • Diff nerves innervate diff parts of basilar membrane • Allows for “place” code for frequency Frequency tuning curves of single hair cells
Complex tones: Fourier decomposition • Basilar membrane acts as f analyzer • Breaks down complex f inputs into constituent pure tone components
Auditory masking: Evidence for cochlear place coding • Auditory masking • Presence of certain tones decreases perception of nearby tones • Similar f result in greater masking • Asymmetry in spread of masking • Consistent with basilar vibrational overlap • E.g. 400 Hz mask overlaps more with 800 than 200 Hz 400 Hz mask Increases threshold for 800 more than 200 Hz
f Mystery of the missing fundamental • 400 Hz fundamental plus harmonics (800, 1200, 1600, 2000) • Sounds like 400 Hz pitch with complex timbre • What if remove fundamental f (400Hz)? • Perceived pitch doesn’t change! • Hence: The missing fundamental • Problem for place coding • No direct stimulation of 400 Hz on basilar membrane • Harmonic structure determines perceived pitch • Not what is present on basilar membrane • What we hear is not what the basilar membrane tell us, but what our brain does
What does Barry White sound like on the telephone? • Telephone carries 300-3400Hz • Typical male voice • Fundamental f = 120 Hz • Barry white • 30 Hz? • Can’t speak to Barry on the telephone? • Missing fundamental allows us to hear “virtual” pitch of voice
If its too loud your too old Pain and pleasure • Db (SPL) scale • Loudness doubles about every 10 db at 1000 Hz • Audibility curves • Loudness varies with f • Low volume • Attenuated low and high f relative to midrange • High volume • Less frequency attenuation • Low volume sounds muddy • Mostly mid range • I like my music loud Each curve represents equal loudness
Otoacoustic emissions: Talking ears • Ears don’t only receive sounds, they make them! • Discovered in 1978 • Tiny microphones • Occur spontaneously and also in response to sound • It like your ears are talking back! • Created by movement of outer hair cells (ohc) • Part of auditory sensitivity is movement of ohc to change region specific flexibility of basilar membrane • Allows tuning curves to be so narrow • Hearing impairments often start with loss of ohc function
Auditory localization • Where is the sound coming from? • Distance • Elevation (vertical) • Azimuth (horizontal) • Localization not nearly as precise as vision • Localization within 2-3.5 degrees in front of head • 20 degrees behind head • Suggests important role of vision • Tunes auditory localization
Why is is auditory localization not obvious? • Vision • Stimulate different photoreceptors in eye • Audition • No such separation of sounds sources on sensory surface • Sources combine to equally stimulate ear receptors
Why have two ears? • Two aural perspectives on the world • Like vision, can be used to get different sound pictures of environment • Binaural cues • The disparities between ears is used for localization
Azimuth • Interaural (between ears) Time Difference (ITD) • Air pressure changes are very slow relative to speed of light • ITD at side = max 600 µS • ITD at front = 0 • Can induce perception of location by varying ITD using headphones • Interaural Level (intensity) Difference (ILD) • Amplitude decreases w/ distance • Head casts sound/acoustic shadow • Reduced amplitude due to reflection • Measure w/ tiny microphones • f dependent • Greater shadow for higher f
Above Level Below Elevation • ITD/ILD not very useful • Use spectral cues • Frequency information can result in different perceptual qualia • Monaural: f serves as signal for pitch • Binaural: f serves as signal for location • Pinna differentially absorb f • Result: Notches in frequency spectra
Distance • At close distances (< 1 meter) • ILD can discriminate near and far • At very close distances ILD is very large (e.g. 20 Db) • But what’s that going to do for us? • At far distances • We are very poor judges for unfamiliar sounds • Suggests that sound serves as signal for visual search • Use sound level for familiar sources • Frequency: Auditory atmospheric haze • Absorption of high f • Sound muffled • Auditory parallax • Sounds move faster across ears at near relative to far distances
Brain basis for localization Sound to right • ITD detectors • Brainstem: Superior olivary nucleus • Primary auditory cortex • Coincidence detection • Neurons fire maximally when signals arrive at same time • Thus: “coincidence” • Axonal distance create input delays Sound to left
Auditory scene analysis • How do we segregate different sounds being produced by many sources simultaneously? • How do we tell what frequencies belong to what source? • E.g., Cocktail party • Don’t perceive an unorganized jumble of frequencies • Not simply high vs low f • Most f ranges overlap • How do we segregate information as belonging to distinct auditory objects?
Principles of auditory grouping • Like gestalt visual principles • Auditory stream segregation • Similarity • Timbre • Location • Pitch • Time 1 stream 2 streams
Auditory-visual interactions: Location and pitch • Visual capture of sound • Location: Ventriloquism effect • Pitch: McGurk effect • “Ba” • “Va” • “Tha” • “Da” • Visual information is integrated with audition • Creates fused auditory visual perception
Auditory-visual interactions: Location and pitch • Auditory experience is much more than pressure level changes