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Sound
Comes from pressure fluctuations in the air
speed through air: 340 m/s (air pressure is related to the amplitude of the sound wave)
speed through water: 1500 m/s
Sound Pressure (Pa)
“Pascals”
measures force exerted by air molecules
Loudness
the psychological perception of sound intensity
measured in dB (decibels)
Decibels (dB)
smallest perceivable pressure
0dB ≠ no sound (represents the minimum audible level)
scale: 0 (threshold of human hearing) —> 140 (Gunshot, fireworks)
Logarithmic Scale
+10dB = 10x increase in intensity
ex. sound of a helicopter x10 more intense than a hairdryer
each 10dB increase represents a tenfold increase in sound intensity
ex. difference between 20dB & 30 dB
20 dB = 10² = 100
30 dB- = 10³ = 1000
difference = 1000 - 100 = 900
Pitch
Psychological aspect of sound related mainly to the fundamental frequency
Frequency (Hz)
The number of cycles per second
Pure tones
Sounds that only have ONE frequency
Equal loudness curve
a graph plotting sound pressure level (dB SPL) against the frequency for which a listener perceives constant loudness
“phon”: unit used which corresponds to the dB value of the curve at 1k Hz
Harmonic Spectrum
the spectrum of a complex sound in which energy is at integer multiples of the fundamental frequency
set of frequencies that make up a complex sound
acts as a “recipe” that determines its unique timber or tone color
typically caused by a simple vibrating source (ex. string of guitar)
Fundamental Frequency
the lowest-frequency component of a complex periodic sound
the lowest frequency of a vibrating object or sound wave that represent the base note or pitch we hear
Timber
psychological sensation by which a listener can judge that two sounds with the same loudness and pitch are dissimilar
timber quality is conveyed by the profile of the harmonics
Auditory canal
tube-like structure that directs sound waves from the OUTER ear to the tympanic membrane (eardrum)
Eardrum (Tympanic membrane)
thin, vibrating membrane that separates the outer ear from the middle ear
transmits sound vibrations to the ossicles
Ossicles
three small bones in the middle ear (malleus, incus, stapes)
amplify and transmit sound vibrations to the inner ear
Cochlea
spiral-shaped, fluid-filled structure in the inner ear
converts sound vibrations into neural signals for hearing
Oval window
membrane-covered opening that connects the middle ear to the cochlea
transmits vibrations from the ossicles
Round Window
flexible membrane in the cochlea
helps relieve pressure from sound waves travelling through the cochlear fluid
Cochlear (auditory) nerve
nerve that carries auditory information from the cochlea to the brain
used for sound processing
Organ of Corti
a structure on the basilar membrane of the cochlea
composed of hair cells and dendrites of auditory nerve fibers
Basilar membrane
structure within the cochlea
vibrates in response to sound
plays a key role in frequency discrimination by supporting hair cells
Tectorial membrane
gelatinous membrane in the cochlea
interacts with hair cells
aid in the conversion of mechanical sound vibrations into electrical signals
Hair cells
sensory receptor cells in the cochlea
detects sound vibrations and convert them into neural signals transmitted to the brain via the auditory nerve
Cochlear Place Code
inner cochlear APEX:
low frequency
low pitch (200Hz)
outer cochlear BASE:
high frequency
high pitch (20,000 Hz)
Temporal Coding
neural strategy where auditory information is encoded by the precise timing and patterns of neuronal spikes rather than just their firing rate
Auditory nerve firing is “phase-locked” (neurons are systematically fired at a given time point of the cycle)
BUT above 4000-5000 Hz: the refractory period of auditory nerve fibers does NOT allow fibers to fire fast enough
Volley Principle
even if individual auditory neurons cannot keep the pace, the whole population of neurons can still temporarily encode frequency
Cochlear Nucleus
first brainstem region that receives auditory signals from the cochlea
where initial sound processing occurs
Superior olive
brainstem structure
involved in sound localization by comparing timing and intensity differences between ears
Inferior colliculus
midbrain structure
integrates auditory information from various brainstem nuclei
plays a role in reflexive responses to sound
Medial Geniculate Nucleus (MGN)
relay station in the thalamus
processes and transmits auditory information to the primary auditory cortex
Primary auditory cortex
region of the cerebral cortex (temporal lobe)
responsible for processing and interpreting sound information
Tonotopy
spatial organization of sound frequency processing in the auditory system
where different frequencies are mapped to specific locations along the cochlea and auditory cortex
Belt region
secondary auditory area surrounding the primary auditory cortex
processes more complex sounds (ex. speech)
Parabelt region
higher-order auditory processing area adjacent to the belt region
processes more complex sounds (ex. speech)
Dorsal “where” pathway
connects auditory areas to the parietal lobe
helps determine the location and movement of sound
Ventral “what” pathway
connects auditory regions to temporal lobe
identifies and categorizes sounds (ex. speech, music)
Conductive hearing imparment
loss of sound conduction to the cochlea (occurs when sound is blocked from reaching the inner ear due to issues in the outer or middle ear)
Frequently caused by wax, fluid build-up, infections, etc.
Not a neural hearing loss, but rather a temporary one
Sensorineural hearing impairment
hearing loss due to damage to the cochlea
ex. congenital (condition from birth), drugs, age, chronic/phasic exposure to loud noise
Interaural Time Difference (ITD)
The difference in time between a sound arriving at one ear versus the other
Listeners can detect interaural delays of as little as 0.01 ms.
Azimuth
the angle of sound source relative to the center of the head
very precise
Physiology of ITD
specialized, high-precision neural circuits in the auditory brainstem (mainly superior olive) that detect the subtle differences in arrival time of a sound wave between the two
with the help of coincidence detector neurons
Interaural Level Difference (ILD)
The difference in level (intensity) between a sound arriving at one ear versus the other
frequency does not matter
longest ms. —> where the sound is coming from
coincidence detector neurons
specialized cells that fire only when they receive multiple excitatory inputs within a very narrow, synchronized time window
Inverse Square Law
the intensity of sound decreases as a function of the inverse of the square of the distance
formula: 𝐼𝑛𝑡𝑒𝑛𝑠𝑖𝑡𝑦_𝑑𝑖𝑠𝑡𝑎𝑛𝑐𝑒 𝑡 = 𝐼𝑛𝑡𝑒𝑛𝑠𝑖𝑡𝑦_𝑠𝑜𝑢𝑟𝑐𝑒 /𝑑𝑖𝑠𝑡𝑎𝑛𝑐𝑒²
sound decreases with distance
harder to tell small differences in distance between 2 objects if they’re both far away than closer
Sound localization Problem
How can you tel if a source is loud and far away vs. close but quiet?
Solution: the spectral composition of sound changes with distance
long wavelengths are always more resistant to obstacles (sound, lights, etc.)
sources that are far away are likely to have encountered more obstacles
air also has “sound-absorbing” qualities
—> therefore, the intensity of higher frequencies decreases as a function of distance (distal sounds have more reverberated than direct energy)
Cones of confusion
Problem: Auditory localization issue where sounds originating from different locations are perceived as coming from the same place
have identical ITD and ILD
Often makes it hard to distinguish if a sound is in front, behind, or above
Solutions:
moving your head (this will change cones of confusion & the only point that will retain its ITD and ILD is the “real” source)
the pinna (also ear canal, head, and torso) slightly distorts the amplitude of certain frequencies as a function of elevation
Directional Transfer function
A specialized type of Head-Related transfer function that describes how the pinna, head, and torso modify sound from a specific direction before it reaches the eardrums
focuses on spectral cues for sound localization by removing the average spectral content
Auditory Stream Segregation
the perceptual organization of a complex acoustic signal into separate auditory events for which each stream is heard as a separate event
Segregation cues
the location of sounds
the frequency (pitch) of the sounds
the timing of the sounds
the timbre of the sounds
the onset of the sounds
rule of “good continuation” (continuity effects)
higher-order information (restoration effects)
Grouping by frequency (pitch)
tones that have similar frequencies will tend to be grouped together
Grouping by timber
tones that have similar timbre will tend to be grouped together
Grouping by onset
when sounds begin at different times they appear to be coming from different sound sources
Continuity effect
Despite interruptions, one can still “hear” a continuous sound if the gap is filled with noise
In that case, the sound is perceived as continuing behind the noise
However, if the gaps aren’t filled with noise, the sound is perceived as separate “chunks”
Grouping by Time
tones that are close together in time will tend to be grouped together
Restoration effect
Despite interruptions, one can still “hear” a sentence if the gaps are filled with noise
in this case, higher-order semantic/syntactic knowledge is used to “fill the blanks”
as for continuity effects, the effect vanishes if the gaps are not filled with noise
Place Coding
Because of how the cochlea and basilar membrane are constructed, acoustic stimuli of different frequencies cause different amounts of movement along the basilar membrane
In the neural tissue, different locations encode specific aspects of sound
Specifically, frequency or pitch
The bigger vibration = lower frequency
Narrower basilar membrane = higher frequency
higher frequency sounds cause bending of the basilar membrane closest to the stapes, resulting in more hair cell activity in that area
Rate coding
low frequency sounds are processed using a rate coding system
the pattern of neurotransmitter release from the hair cells deepest in the cochlea (furthest from the stapes) determines the perception of low frequency sounds
Time coding of sound
Frequency will generate a firing of neurons at the same frequency throughout time
a process where auditory nerve fibers fire in a specific pattern in response to sound waves
Hearing Aids
Acoustic amplifiers that you put in the ears to make sounds louder and then restore audibility to softer sounds
“micro-amplifiers”
If hearing loss is severe → cochlear implants
Types of hearing aids
In-the-ear (ITE) hearing aids
Behind-the-ear (BTE) hearing aid
Much smaller because it is detached from the amplifier (leaves room to allow air to go through the ear canal and leaves it open)
Used when we want to address hearing loss in the high frequencies only (Frequency-specific amplification)
Hearing aid fit check
validation of acoustic output at the tympanic membrane
Electro-acoustic stimulation (EAS)
the use of a hearing aid and a cochlear implant technology together in the same ear
Output of a speakerphone (hearing aid)
A cord to receive sounds and turn them into an electrode (cochlear implant)
Stimulate the nerve for the high frequencies
—> We use the software on those machines to adjust to the needs of the patients
Hearing aids (How they work)
a microphone captures sound waves and converts them into digital signals
a computer processor amplifies and modifies these signals to match specific hearing loss needs
a speaker (receiver) converts the signals back into sound waves, delivering them into the ear canal
Cochlear implants
An electrode array that is installed along the cochlear nerve to restore hearing
Includes two pieces:
External: There is an external piece with a microphone that picks up a sound
Internal (under the skin): Then signals the sound to a subcutaneous piece that itself trans converts the sound into an electrical signal along the cochlea
Placed surgically
Candidacy for device
Hearing aid candidate:
< 60 dB of hearing loss
> 60% of speech recognition
Cochlear implant candidate:
> 60 dB of hearing loss
< 60% of speech recognition
The Superior Temporal Sulcus
auditory analogue of the fusiform face area in the visual system
region that is selective for voices
Wernicke’s area
Most important area to understand language
posterior section of the superior temporal gyrus (left hemisphere)
Wernicke’s aphasia
Fluent language but with a lack of sense/meaning (inability to recognize sounds as words)
Impairment in meaning understanding
Reading is impaired (able to read the words but lack understanding of the meaning of words)
Speech does not make sense because they do not know the literal meaning of words
Broca’s area
Controls the motor organization of speech sounds
In the inferior frontal gyrus (left hemisphere)
Broca’s Aphasia
Not fluent language (difficulty finding the words to communicate) but comprehension is intact
Opposite of wernicke’s aphasia
Trouble repeating words
Patients can recover to some degree
Phonemes
a unit of sound that distinguishes one words from another in a particular language
Ex. kill vs. kiss
Required when speaking and understanding speech
Respiration (#1)
The diaphragm pushes air out of lungs → through trachea → up to the larynx (contains vocal folds or vocal cords)
1st step to speech production
Phonation (#2)
The process through which vocal folds are made to vibrate when air pushes out of the lungs
Vocal folds will vibrate as the air pushes through these folds (on each sides)
More tension = higher-pitched sounds
Small vocal cords, high-pitched voices → children < women < men
The type of sound that is being produced by the vocal folds will have a harmonic spectrum (amplitude/frequency)
Articulation (#3)
The act or manner of producing a speech sound using the vocal tract
Oral tract (vocal and nasal) is the area above the larynx
Humans can change the shape of their vocal tract by manipulating the jaws, lips, tongue, etc. (“articulation”)
This allows the change in sounds
—> Changing size and shape of vocal tracts will increase/decrease energy at different frequencies
includes formants and spectograms
Formants
Peaks in the speech spectrum (helps identifying the phoneme)
Labelled by number (from lowest to highest)
Shorter vocal tracts = higher frequencies
Spectrograms
Represent the sentence said by someone (based on amplitude, frequency and time)
The spectrum of speech sounds changes over time
x: time
y: frequency
color: energy (amplitude)
Coarticulation
Phenomenon when our oral tract anticipate the next phoneme as we are pronouncing the current phoneme
Occurs to experienced talkers (they position their tongues in antiticipation of the next consonant or vowel) → causing a change in pronunciation (overlap)
Makes it difficult for the auditory system to identify the phoneme
Ex. “di” → “du”
Categorial Perception
Occurs when a stimulus changes continuously but we perceive it as belonging to discrete categories
Because of this → differences across category boundaries appear LARGER than equally large differences within category
We do not perceive the sounds as continuously varying BUT there are sharp categorical boundaries between stimuli
Perceived as more different
—> resolution to coarticulation
Learning to Listen
4-day old French babies prefer hearing French over Russian
sound distinctions are specific to various languages (ex. “r” and “l” are not distinguished in Japanese)
infants begin filtering out irrelevant acoustics long before they start to say speech sounds
ex. english-speaking infants <10 months can distinguish between two types of “t” sounds that are different phonemes in Hindi but lose that ability after 10 months, while Hindi infants still continue to make distinction
—> Babies aren’t just learning words—they’re reshaping how they hear the world.
At first: broad, flexible perception
Over time: specialized for their native language
“Motor theory” of speech perception:
Motor processes used to produce speech sounds are used in reverse to understand the acoustic speech signal
Supported by the McGurk Effect: showed that what someone sees can affect what they hear
Problem: Speech production is as complex, if not more complex, than speech perception
solution: reduce the number of phonetic categories
Tone height
a sound quality corresponding to the level of pitch
Tone height is monotonically related to frequency
Tone Chroma
a sound quality shared by notes that have the same octave interval
Each note on the musical scale (A-G) has a different chroma
Octave
the interval between two sound frequencies having a ratio of 2:1
Western music has 13 notes separated by equally spaced pitch intervals (semitones)
There are 10 octaves within the audible range (piano has 7 octaves)
Consonance
When two or more notes are played simultaneously (a chord) or sequentially
Consonance: the combination of sounds is pleasant, as if the notes “go together”
Happens when the fundamental frequency of the two notes has a simple ratio
Many harmonics of the two sounds will coincide
→ study shows that even people as young as 2-month old infants prefer consonant conditions over dissonant ones
Dissonance
When two or more notes are played simultaneously (a chord) or sequentially
Dissonance: the combination of sounds is unpleasant or “off”
Happens when the fundamental frequencies of the two notes have a complex ratio
Very few harmonics will coincide
Scale
A particular subset of the notes in an octave
Key
the scale that functions as the basis of a musical composition
Ex. a composition in the key of C major contains notes mostly from the C major scale
Major and Minor scales
Major and minor scales are differentiated by the pattern of intervals (number of semitones) between successive notes
Major scales: 2-2-1-2-2-2-1
Sound “happy”
Minor scales: 2-1-2-2-1-2-2
Sound “sad”
Tonic
the root note of the key
Acts as the gravity point of the key
Moving away from and back at resting point → what makes music interesting and why it has a pleasing effect on us
Melody
a sequence of notes or chords perceived as a single coherent structure
Ex. row, row, row your boat
defined by contours
can change octaves or keys and still be the same melody even if they have completely different notes
Contours
The pattern of rises and declines in pitch (rather than by an exact sequence of sound frequencies)
Right auditory cortex
Music is mostly processed in the RIGHT auditory cortex
speech is mostly processed in the left auditory cortex
Music and Brain study
Study:
Presented two different pitches to participants:
1st group: stable, still pitch
2nd group: changing pitch
Result:
Fixed spitch sequence vs silence → primary auditory cortex is active
Changing pitch sequence vs. fixed pitch sequence → melodies are being processed by the right belt and para belt regions of the right auditory cortex
Temporal resolution
Speech requires fine temporal resolution
Production is very fast (10-15 consonants & vowels per second)
Can be doubled if you talk fast
Experienced talkers position tongue in anticipation of next consonant or vowel → causing coarticulation
Coarticulation will cause overlap in articulatory or speech patterns
Spectral Resolution
music requires fine spectral resolution
The left auditory cortex is able to distinguish the different lyrics but not the different melodies
The right auditory cortex is able to distinguish the different melodies but not the different lyrics
Congenital Amusia
inability to perceive music
perfectly normal otherwise, only have a problem with identifying music
Unaware of how they sing
Fail to recognize popular tunes, without the help of lyrics
Akin to dyslexia, prosopagnosia, etc.
Seems to have problems with their arcuate fasciculus (on the right side)
Congenital Amusia Diagnosis
Developed a battery of musical tests
In our brains we have different specific modules/regions for processing the melody.
When we process music → we combine melody and the temporal structure to recognize the musical excerpt
Arcuate Fasciculus
bundle of white matter that connects the auditory cortex to frontal regions (inferior frontal gyrus and superior temporal gyrus)
Absolute Pitch
capacity to actually name or produce a pitch without comparisons to other notes
1/1500 have it;
Heritable (has genetic basis)
requires exposure to musical training in early life (environment)
Reward Prediction Error
concept used to describe how dopamine neurons in your brain (specifically in the ventral tagmental area of brainstem) signal the difference between the reward you expect and the reward you actually get
Works in 3 main ways:
Positive surprise: if a reward happens and you didnt see it coming → dopamine neurons show a sudden burst of activity
No surprise: if you see a cue that predicts a reward is coming → dopamine neurons fire when they see that cue
However when the reward arrives there is no extra bursts
Negative surprise: if you expect a reward but it does not happen → your dopamine neurons fire at the initial cue but activity drops or pauses at the moment the reward was supposed to appear
→ Reward circuitry is interested by learning what causes the reward and not necessarily the reward itself