Glossary

Terms used in this book.

A reference list of the technical vocabulary used in What is hearing?. Inline occurrences of these terms in the lessons are auto-tooltipped (dotted underline) so you can hover for a quick definition; for a fuller treatment with context, return here.

89 terms from this book.

A

A1
Primary auditory cortex, on Heschl’s gyrus. The cortical entry point of the auditory pathway.
acoustic impedance
The ratio of acoustic pressure to particle velocity in a propagating wave (Z = p/v). For a plane wave in a medium of density ρ and wave speed c, Z = ρc.
amplitude
The magnitude of a wave’s departure from equilibrium. For sound, the size of the pressure fluctuation.
auditory nerve
The ~30,000-fibre bundle carrying spike-train information from the cochlea to the cochlear nucleus in the brainstem.
auditory streaming
The brain’s perceptual organisation of acoustic input into one or more sources (streams). A bistable example is the van Noorden galloping triplet.
auricula
Anatomical name for the pinna.

B

basilar membrane
The membrane separating scala media from scala tympani. Its position-dependent stiffness gives different places different natural frequencies.
Bayes
Bayes’ theorem: P(M|S) = P(S|M)·P(M)/P(S). The brain’s posterior over hypotheses M given sensory data S combines the likelihood with the prior.

C

characteristic frequency
The frequency at which a given place on the basilar membrane (or auditory-nerve fibre, or cortical neuron) responds most strongly.
cochlea
The spiral, fluid-filled organ of the inner ear that performs frequency analysis on incoming sound and transduces it into neural signals.
cochlear amplifier
The active feedback process in the cochlea, driven by outer-hair-cell electromotility, that sharpens basilar-membrane tuning beyond what passive mechanics gives.
cochlear nucleus
The first central auditory structure; receives all auditory-nerve fibres on its own side. Has ventral and dorsal subdivisions.
cochlear synaptopathy
Damage to the synapses between inner hair cells and low-spontaneous-rate auditory-nerve fibres, often invisible on standard audiograms but causing difficulty hearing in noise.

D

decibel
A logarithmic unit of ratio: 20·log10(amplitude ratio) or 10·log10(power ratio). Used for sound pressure level (SPL) and hearing level (HL).
duplex theory
Rayleigh’s theory (1907) that ITDs dominate low-frequency localisation and ILDs dominate high-frequency localisation.

E

ear canal
The tube about 25 mm long running from the pinna to the eardrum. Its closed-tube resonance amplifies frequencies near 3 kHz.
eardrum
The tympanic membrane: a thin sheet at the inner end of the ear canal that vibrates in response to pressure waves and drives the ossicular chain.
electromotility
The voltage-driven length change of outer hair cells, mediated by prestin. Generates the active gain of the cochlear amplifier.
endocochlear potential
The +80 mV potential of the endolymph in scala media relative to perilymph. Powers hair-cell transduction.
endolymph
The fluid in scala media; high in K⁺, low in Na⁺. Held at +80 mV (the endocochlear potential) relative to perilymph.

F

formant
A resonant peak in the spectrum of a vowel. The configuration of mouth and tongue produces F1, F2, F3… which together identify the vowel.
free-energy principle
Friston’s proposal that any self-organising system minimises variational free energy, a bound on negative log-evidence under its generative model.
frequency
The number of oscillation cycles per second, measured in hertz (Hz). For sound, this is what the brain perceives as pitch.

G

Greenwood function
The empirical fit relating cochlear place x to characteristic frequency f: f(x) = A(10^(α(1−x/L)) − K). For humans: A=165.4, α=2.1, K=0.88, L=35 mm.

H

helicotrema
The small opening at the cochlear apex where scala vestibuli and scala tympani communicate, short-circuiting DC pressure imbalance.
Hopf bifurcation
The critical point at which a damped oscillator becomes a self-sustained one. The mammalian cochlea operates near this bifurcation.
HRTF
Head-Related Transfer Function. The full frequency-dependent filter the body applies between a sound source in space and the listener’s eardrum.

I

ILD
Interaural Level Difference. The intensity difference between the two ears, dominant at frequencies whose wavelength is smaller than the head.
impedance
In acoustics, the ratio of pressure to particle velocity. A measure of how strongly a medium resists being moved by a wave.
incus
The second middle-ear ossicle (the “anvil”); pivots between malleus and stapes.
inferior colliculus
A midbrain nucleus and obligatory waystation for ascending auditory information. Where ITDs, ILDs and other features converge into a spatial map.
inner hair cell
The cochlea’s actual transducer. ~3,500 of them along the basilar membrane, each contacting 10–30 auditory-nerve fibres.
interaural level difference
The intensity difference between the two ears, dominant at frequencies whose wavelength is smaller than the head.
interaural time difference
The arrival-time difference for a sound between the two ears. Humans resolve differences as small as 10 μs.
ITD
Interaural Time Difference. The arrival-time difference for a sound between the two ears. Humans resolve differences as small as 10 μs.

J

Jeffress model
Lloyd Jeffress (1948) proposed delay-line coincidence detectors as the neural mechanism for ITD measurement; still the canonical picture for the MSO and bird homologue.

L

lateral lemniscus
The white-matter tract carrying the ascending auditory pathway from cochlear nucleus and superior olive to the inferior colliculus.
lateral superior olive
LSO. Brainstem nucleus that computes interaural level differences from excitatory ipsilateral and inhibitory contralateral inputs.
likelihood
In Bayesian inference, P(S|M): how probable the observed sensory input is given that hypothesis M is true.
LSO
Lateral superior olive. Brainstem nucleus that computes interaural level differences.

M

malleus
The first middle-ear ossicle (the “hammer”); attached to the eardrum on one side and the incus on the other.
McGurk effect
A multisensory illusion in which the visual articulation of one syllable and the audio track of another produce a third percept.
medial geniculate body
MGB. The thalamic relay for auditory information, sitting between the inferior colliculus and the auditory cortex.
medial superior olive
MSO. Brainstem nucleus that computes interaural time differences using coincidence detection.
MET channel
Mechanoelectrical Transduction channel. A mechanically-gated ion channel on stereocilia that opens when the bundle is deflected, depolarising the hair cell.
MSO
Medial superior olive. Brainstem nucleus that computes interaural time differences.

O

organ of Corti
The sensory structure on the basilar membrane containing inner and outer hair cells, supporting cells, and the tectorial membrane.
ossicles
The three smallest bones in the body — malleus, incus, stapes — that transmit motion from the eardrum to the oval window of the cochlea.
outer hair cell
Active mechanical element of the organ of Corti. ~12,000 of them; they shorten and lengthen with voltage, sharpening basilar-membrane tuning.
oval window
A membrane-covered opening at the base of scala vestibuli, where the stapes footplate drives the cochlear fluid.

P

perilymph
The fluid in scala vestibuli and scala tympani; ionic composition resembles extracellular fluid (high Na⁺, low K⁺).
phase locking
The tendency of an auditory-nerve fibre to fire at a preferred phase of a low-frequency stimulus cycle, up to ~5 kHz.
phonemic restoration
A perceptual effect in which a phoneme replaced by noise is heard as intact, restored by context-driven priors.
pinna
The visible outer flap of the ear. Its folds and ridges encode elevation by introducing spectral notches that depend on the sound source’s vertical angle.
place coding
Encoding of frequency by which location on the basilar membrane (or fibre in the auditory nerve) is most active.
plane wave
A wave whose phase fronts are infinite parallel planes; idealisation of a wave from a distant source, valid locally near the listener.
posterior
In Bayesian inference, P(M|S): the probability of hypothesis M given the sensory input S. This is what the brain perceives.
predictive coding
A theory of cortical processing in which higher layers predict the activity of lower layers, and only the prediction errors propagate upward.
prestin
The membrane protein in outer hair cells that changes conformation with voltage, producing the cell’s electromotility.
primary auditory cortex
A1. The first cortical processing stage for sound, on Heschl’s gyrus of the superior temporal plane. Tonotopically organised.
prior
In Bayesian inference, P(M): the probability of a hypothesis before any sensory input. Context, expectation, learned experience shape this.

Q

quality factor
Q. A dimensionless measure of resonance sharpness. Equal to the peak amplitude (relative to DC) of a damped driven oscillator.

R

rate coding
Encoding of stimulus intensity (or frequency above the phase-locking limit) by the firing rate of a neuron.
reflection
When a wave hits a boundary between two media, part of its energy turns back into the first medium. The reflection coefficient R = (Z2 − Z1)/(Z1 + Z2).
resonance
The condition where a driving frequency matches a system’s natural frequency, producing maximum response amplitude.
reverberation
The collection of reflections that follows a direct sound in a room, gradually decaying. Characterised by reverberation time, T60.
ribbon synapse
A specialised presynaptic structure in inner hair cells (and a few other sensory cells) that releases vesicles continuously at high rates with sub-millisecond precision.
round window
A flexible membrane at the base of scala tympani that bulges out to give the incompressible cochlear fluid somewhere to go.

S

scala media
The middle cochlear chamber, filled with endolymph; contains the organ of Corti.
scala tympani
The lower of the three cochlear chambers, filled with perilymph; terminates at the round window.
scala vestibuli
The upper of the three cochlear chambers, filled with perilymph; the stapes pushes into it at the oval window.
Shepard tone
An auditory illusion of a continuously rising (or falling) pitch, created by stacking octaves with a fading amplitude envelope.
speed of sound
The propagation speed of small-amplitude pressure disturbances. ≈343 m/s in air at room temperature, ≈1480 m/s in water.
spiral ganglion
The collection of cell bodies whose axons form the auditory nerve. Each spiral-ganglion neuron contacts one or a few inner hair cells.
stapedius reflex
A bilateral reflex that contracts the stapedius muscle in response to loud sounds, attenuating ossicular transmission at low frequencies by 10–15 dB.
stapes
The third middle-ear ossicle (the “stirrup”); its footplate seats in the oval window and drives the cochlear fluid.
stereocilia
Actin-filled rod-like protrusions on the apical surface of hair cells, arranged in a graded array with tip links connecting neighbours.
STRF
Spectro-Temporal Receptive Field. The 2-D function (frequency × time) characterising what stimulus features drive a cortical neuron to fire.
superior olivary complex
A brainstem region where information from both ears first meets. Contains the MSO (ITDs) and LSO (ILDs).
superior temporal gyrus
STG. A region of temporal cortex extending forward from A1; contains regions specialised for processing the phonemic structure of speech.

T

tectorial membrane
A gelatinous strip overhanging the organ of Corti. Couples basilar-membrane motion to the stereocilia of the hair cells.
thalamus
A central diencephalic structure that relays most sensory information to cortex. The medial geniculate body is its auditory subdivision.
tip link
A molecular thread connecting the top of one stereocilium to the side of its taller neighbour. Stretching opens the MET channel.
tonotopy
Spatial organisation in which neighbouring elements respond to neighbouring frequencies. Preserved from cochlea through brainstem to cortex.
traveling wave
A wave that propagates while maintaining its shape (in a uniform medium) or that varies its shape as it moves (in a non-uniform one like the cochlea).
tympanic membrane
The eardrum: a thin sheet that vibrates in response to pressure waves and drives the ossicular chain.

V

vector strength
A measure of phase-locking strength: |⟨exp(iφ)⟩| averaged over many spikes. 1 = perfect, 0 = random.

W

wave equation
A second-order partial differential equation describing how a disturbance propagates. For pressure in air: ∂²p/∂t² = c²∇²p.
WKB approximation
A method for solving wave equations with slowly-varying coefficients. Gives a solution of the form A(x)·exp(i·∫κ(x)dx) for a position-dependent wavenumber.