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Special Senses Breakdown
All special senses have corresponding ORGANS:
Organ → special sense → stimuli → neural sensation
eye → vision → light → colour
Ear & cochlea → hearing → sound waves → pitch
Vestibular apparatus → balance → head movement → motion
Noes & olfactory epithelium → smell → airborne chemicals → smell
Tongue & taste buds → gustation → tastants → flavour
Hearing: Ear Anatomy
Large, fleshy external structure; auricle
Internal structures are protected by the skull (temporal bone)
Cochlea is also a bony shell, called the osseous spiral lamina
Protects the scala vestibuli, scala media (cochlear duct) & scala tympani
Hearing: Ear Function
Auricle catches soundwaves and funnels them into the ear canal
Sound passes down ear canal.
Soundwaves hit the eardrum/timpanic membrane, at the end of the ear canal.
Eardrum separates outer & middle/inner ear
Middle ear bones: Malleus, Incus & Stapes vibrate due the waves in that order, amplifying the soundwaves as they travel to the cochlea

Hearing: Sensation
Middle ear bones: Malleus, Incus & Stapes vibrate due the waves, amplifying the soundwaves. The last bone, stapes hits the fleshy oval window, which propagates vibrations into the cochlea
Vibrations travel up the cochlea’s spiral through the scala vestibuli, towards the helicotrema (spiral apex), and back down the spiral through the scala tympani
This order of movement causes the middle space, the cochlear duct (scala media), to move up and down, stimulating mechanoreceptors inside called ‘hair cells’.
Hair cells have branches of the cochlear nerve attached to their bases (part of Vestibulocochlear nerve). Movement of hair cells stimulate this cochlear nerve.

Hearing: Sensation: Cochlear duct (Scala media)
The cochlear duct is located as the middle of three spaces occupying the cochlea (view image), hence the alternate name scala media.
The basilar membrane is located as the border separating the lower scala tympani from the middle cochlear duct.
Within this cochlear duct, can the organ of corti be located (boxed in bottom image).
As the vibrations move the fluid within the cochlea, the basilar membrane in the cochlear duct moves, and the organ of corti is stimulated

Hearing: Sensation: Organ of corti
The organ of corti is located at the bottom of the cochlear duct. With the basilar membrane bordering the bottom of it.
Connected above the basilar membrane are mehanoreceptors, called “hair cells”.
Connected to the base of these hair cells are fibres of the cochlear nerve running through the basilar membrane.
Directly above the hair cells is the tectorial membrane.
The ‘hairs’ of some hair cells can be connected to the tectorial membrane
Vibrations travelling through the fluid in the scala vestibuli & scala tympani pushes and moves the basilar membrane up and down.
Pushes hair cells up, and the hairs are smushed into the tectorial membrane
As the hair cells are mechanoreceptors, the physical deformations in their cell membranes activate then, sending graded receptor potentials which can become action potentials down the cochlear nerve
Transduces physical sensation (movement of hair cells from vibrations) into electrical signals

Hearing: Mechanoreceptors: Hair cells
Kinocilium: Tallest hair on each hair cell
Stereocilia: Remaining hairs on each hair cell (includes kinocilium)
Kinocilium and stereocilium are connected at their tips by fibres called “tip links”.
Activation of hair cells:
Due to movement of the basilar membrane, hair cells are pushed up into the tectorial membrane.
Due to the forces when the vibration waves arrive, stereocilia bend towards tallest hair, kinocilium.
Tension forms at the tip links
Tension physically opens K+ and Ca2+ ion-gated channels
Causes rapid depolarisation of cell membranes in sensory neurons (mechanoreceptors; hair cells)
Neurotransmitters released towards afferent cochlear fibres
Action potentials propagate via afferent cochlear fibres towards the brain
Special somatic afferent nerves of the Vestibulocochlear nerve (Cranial nerve 8)
Deactivation of hair cells:
When vibration waves move away, due to the forces stereocilia bend away from the tallest hair, kinocilium
Tip links relax and ion-gated channels remain closed.
No neurotransmitters released.
Sound waves can increase or decrease cochlear nerve activity:
Tectorial membrane leaning towards kinocilium, more action potentials.
Tectorial membrane moving away from kinocilium, less action potentials.
Diagram: cochlear axon, NOT axon terminal

Hearing: Sound
Sound is a pressure disturbance, alternating areas of high and low pressure of molecules travelling through air or water.
Peaks: high pressure (compressed areas)
Troughs: low pressure (rarified areas)
Wavelengths: Distance between wave peaks

Hearing: Sound Frequency: Sensitivity
Humans can sense from 2,000 to 20,000 frequencies (measured in hertz).
more sensitive to frequencies between 1,500 to 4,000 hertz
Frequency: Number of soundwaves/wavelengths propagated in a certain period of time
More frequency, lower wavelength, vice versa
Frequency determines pitch
Smaller wavelengths → Higher frequency → higher pitch
Larger wavelengths → Lower frequency → lower pitch

Hearing: Sound intensity/loudness
The loudness of a sound is proportional to its amplitude in soundwaves.
Higher amplitude: louder
Lower amplitude: quieter
Frequency/pitch/wavelengths are irrelevant
However, usually lower amplitudes (quieter) are associated with deeper sounds with low pitches (e.g tummy grumbling)
Contrarily, higher amplitudes (louder) are associated with sharper sounds with high pitches (e.g mouse squeak)


Hearing: Cochlea: Basillar membrane - Tonotopically organised
The basilar membrane is a horizontal lining of fibres extending down towards the cochlear apex. Its structure changes as it moves towards the cochlear apex and is called tonotopical organisation.
Base of cochlea:
Shorter fibres, more taut & rigid
Thicker fibres
Less flexible
Requires more energy and force to be moved by cochlear fluid
Vibrates at shorter wavelengths, high frequencies, higher pitch, higher amplitude
Apex of cochlea:
Longer fibres, more floppy & malleable
Thinner fibres
More flexible
Requires less energy and force to be moved by cochlear fluid
Vibrates at lower wavelengths, lower frequencies, lower pitch, lower amplitude

Hearing: Neural pathways
Special somatic afferent nerves of the Vestibulocochlear nerve (Cranial nerve 8)
Where?
Action potentials propagate from hair cells
Travel along the cochlear branch of the vestibulocochlear nerve
Synapse with the cochlear nuclei within the brain stem’s medulla
Information from both ears combines within the cochlear nuclei
Fibres travel up the brain stem, some staying on their respective sides
Fibres reach the temporal lobe, and respect left & right auditory cortices (cortexes)
Hearing is perceived
Auditory info is processed by other sections of the brain to enable interpretation & integration of the image
Beginning: In the inner ear, specifically base of hair cells lining the cochlea
Ending: Primary auditory cortex, temporal lobe

Hearing: Temporal lobe, Primary & secondary auditory cortex map
Each segment of the primary auditory cortex responds to impulses arriving from specific nerve fibres. Each section responds to a certain cochlear nerve fibre that reacts to a certain pitch/frequency
From anterior to posterior, the primary auditory cortex transduces cochlear sensation from the apex, to the base.
Left & right sides respond to information received from both ears
Hence, damage to one side of the brain’s temporal lobe (e.g left temporal lobe), will cause hearing deficits in both ears

Hearing: Locating sounds: Interaural time difference (Spatial audio)
Unless positioned directly in front of a listener, sounds heard angled away from the listener will be heard microseconds apart between both ears.
This time difference is termed the Interaural time difference
Helps brain position sound sources

Hearing: Locating sounds: Interaural intensity difference (Spatial audio)
Unless positioned directly in front of a listener, sounds heard angled away from the listener will be heard a different intensities between ears, due to the head blocking some soundwaves.
This time difference is termed the Interaural intensity difference
Helps brain position sound sources

Hearing Disorders: 1 of many: Conductive hearing loss
Soundwaves entering the ear canal cannot be transmitted across the eardrum into the inner ear.
Temporary causes: Ear wax buildup, ruptured eardrum
Permanent causes: Otosclerosis - inner ear bones (Maleus, incus & stapes) become malformed and cannot vibrate
Stapes fuses to oval window
Maleus, incus and stapes become fused in some form, preventing vibration
Conductive hearing loss: Soundwaves cannot be “conducted” to the cochlea

Hearing Disorders: 1 of many: Sensorineural hearing loss
Sensorineural hearing loss: Issues with the sensory neurons or neural pathways that cause hearing loss.
Damage to the cochlea or neural pathways towards the brain
Often permenant
Hearing Disorders: Technology
Hearing aids: Amplify incoming sounds, making soundwaves more conductive and able to pass the ear canal towards the cochlea.
Cochlear implants: Directly stimulate the cochlear nerve via electrical impulses, allowing soundwaves to bypass the cochlea.
Used when substantial hearing damage is present, and hearing aids provide no effect.
