Week 8 - Special Senses

Introduction
Last week you learnt about the receptive structures that permit the transmission of sensory information
related to taste and smell. This week we will focus on hearing and vision. In order for us to have a
conscious perception of these senses this information needs to be relayed to the central nervous system.

We will focus on the central pathways that transmit light and sound information and learn about the key
structures that are common to the transmission and processing of this sensory information.

Learning Outcomes
By the end of this lecture, you should be able to describe the following:
●​ LO1: the peripheral structures involved in vision
●​ LO2: how visual information is transduced
●​ LO3: the central pathways involved in the processing of visual information
●​ LO4: the peripheral structures involved in hearing
●​ LO5: how auditory information is transduced
●​ LO6: the central pathways involved in the processing of auditory information

LO1: Describe the peripheral structures involved in vision

In order to "see", we need to convert electromagnetic energy into neural signals. To understand how this
occurs, you need to know about the anatomy of the eye and how that enables electromagnetic rays to hit
the retina, where these signals then get converted into neural signals. Light is a form of electromagnetic
radiation that is visible to our eyes and can be described as a wave of energy. Light, therefore, has
properties associated with any form of wave:

These properties are wavelength, frequency and amplitude. Wavelength is the distance between
successive peaks or troughs. Frequency is the number of waves per second, and amplitude is the 'height'
difference between the trough of a wave and the peak of a wave.

The frequency of electromagnetic radiation indicates the amount of energy contained within that source.
For example, high-frequency radiation has the greatest energy content, whereas low-frequency radiation
has the lowest content. Our visual nervous system is only capable of detecting electromagnetic radiation
that occurs at the lower end of the spectrum. Visible light has a wavelength of between 400 and 700 nm.
Different wavelengths of light emit energies that are associated with different colours. For example,
wavelengths of light that are low frequency and therefore emit lower energy are visualised as red or
orange, whereas wavelengths of light that are higher frequency and emit higher energy are visualised as
blue or violet. In an idealised empty space where there is nothing to interact with, the light would travel in
a straight line. In the real world, this is not the case, and there is so much within the environment, for
example, atmospheric gases through to physical structures such as household furniture, that light can
interact with.

WHY AM I LEARNING THIS? The eye is an “object” within the environment that light encounters.
Knowing how light travels and encounters objects in our environment is therefore important for
understanding how light interacts with the eye.

Light will travel in a straight line until it encounters these substances or objects, and when it does, it will
do one of three things: it will either reflect off an object, be absorbed or refract through the object.
Reflection is characterised by the bouncing of light off a surface. How light rays reflect off a surface
depends on the angle that it hits the surface. For example, if light hits a surface perpendicularly, it will
back at an angle of 180 degrees, essentially travelling back in the direction it came from. Whereas if it hits
at an angle of 45 degrees, it is reflected at 90 degrees. When we see light, what we are actually seeing is
light that is reflected off objects within our environment.

Absorption refers to the transfer of light energy to a particular object, i.e. that object has absorbed the
light. The amount of light that an object absorbs determines what colour we see that object to be. For
example, if an object absorbs all the light rays that hit it and absolutely no light rays are reflected back off
the object, then the colour will be seen as black:

If instead, that object absorbed all wavelengths of light except for those wavelengths that emit a light
energy that is visualised as blue, we will see that object as blue due to those blue wavelengths of light
being reflected off the object:

Refraction is the bending of light rays that occurs when light rays travel from one transparent medium to
another. This process forms the basis of how images are formed in the eye. Light will bend when it travels
from one medium to another due to the difference in speed that light can travel through those two
mediums. For example, if there is little difference in the speed at which light can travel through the two
mediums, then there will be little bending of light. If we think about how this applies to the eye now.
Light can travel through the air much faster than it can travel through the fluid that is contained within the
eye. When the light rays hit this fluid, they refract due to the reduction in speed with which they are
travelling. This is similar for when light encounters a medium such as water, which slows down the speed
the light can travel - it will bend. The greater the difference in the speed that light can travel, the more
light will bend.
To understand this more, lets look at the key structures of the eye. There are a number of key gross
anatomical features of the eye that you need to be aware of to understand how we see. Interact with the
image below to learn more.
●​ The ciliary muscles are a set of smooth muscles in the eye responsible for controlling the shape of
the lens to facilitate focusing (accommodation).
●​ The cornea is the glassy transparent external surface of the eye that covers the pupil and iris.
●​ The iris surrounds the pupil and contains pigment that gives the eye its colour and helps to control
light levels.
●​ The pupil is the opening that allows light to enter the eye and reach the retina.
●​ Aqueous humor is a watery fluid contained in a compartment (anterior chamber) between the
cornea and the lens.
●​ The lens is a transparent structure, located directly behind the iris, that along with the cornea
transmits light to the retina.
●​ The zonule fibres are ligaments that are attached to the ciliary muscle and suspend the lens.
●​ Three pairs of extraocular muscles are inserted into the sclera and move the eyeball in the eye's
orbit.
●​ The sclera is the "white of the eye". It is continuous with the cornea and forms the tough wall of
the eyeball.
●​ At the back of the eye is the retina, which contains the sensory cells that convert light energy into
neural signals that can be communicated via the optic nerve to the central nervous system. Within
the retina itself, there are two important structures to be aware of:
●​ The first is the macula. This is the part of the retina that is responsible for what is termed central
vision. This is located in the middle of each retina and can be identified by its yellowish hue.
●​ The second region is the fovea. This is a dark spot that is about 2 mm in diameter found in the
centre of the retina and is the thinnest part of the retina.
●​ Cells within the retina are organised in layers that are named in reference to the middle of the
eyeball. We will look at these layers in order, starting with the layer closest to the middle of the
eyeball. This innermost layer is the ganglion cell layer which contains the cell bodies of ganglion
cells. The next layer is the inner plexiform layer which contains the synaptic contacts between
bipolar cells, amacrine cells, and ganglion cells. Beneath this is the inner nuclear layer which
contains the cell bodies of the bipolar cells, horizontal cells, and amacrine cells,. Followed by the
outer nuclear layer, which contains the cell bodies of the photo-receptors The outermost layer of
the retina contains the outer segment of the photo-receptors, which is where the photo-pigments
that ultimately respond to light that enters the eye are contained. This means that the retina is
organised in an inside out fashion, such that light has to travel through all layers as well as the
cell bodies of the photo-receptors before it hits the sensory element of the photo-receptor and can
initiate any transduction processes. Beneath the photo-receptor layer is a layer of pigmented
epithelium. This layer plays a critical role in the maintenance of the photo-receptors and
photo-pigments, and absorbs any light that passes entirely through the retina, which helps
minimise the scattering of light within the eye that would otherwise blur the image formed on the
retina.
Now that we have discussed how light travels and the structure of the eye, lets look at how light interacts
with the eye. The eye collects light rays emitted by or reflected off objects and focusses them onto the
retina in order to form images. In order to bring images into focus light needs to be bent so that converges
on the retina. This mainly occurs as light hits the cornea. The angle at which light hits the retina
determines how the light rays bend. If light hits the cornea at 90 degrees, it will pass straight through the
aqueous humour without bending. If light hits the cornea at 45 degrees, it will bend and enter the aqueous
humour at 90 degrees. This bending of light ensures that regardless of the angle that light hits the cornea,
all light rays converges on the same point in the retina. The reason that light refracts off the cornea and
converges on the retina is because the cornea is curved.
The distance that this refracted light has to travel between the cornea to the retina is called the focal
distance.

CLINICAL CONNECTION. What do you think would occur if the curvature of the cornea changed?
The curvature of the cornea can be altered in some visual conditions, such as astigmatism. This increase
in curvature of the cornea shortens the focal distance, and therefore light converges on a single point in
front of the retina, causing blurred vision. This can be corrected through the use of glasses which act to
increase the focal length and ensure that light converges on the retina.

The lens of the eye also causes some additional light refraction, which aids in increasing the sharpness of
an image, which is particularly important for near vision. The lens achieves this through a process called
accommodation. During accommodation, the ciliary muscle contracts and swells in size. This causes the
lens to become rounder and therefore increases the refractive power of the lens. Light rays that would
have otherwise converged behind the retina now have a shorter focal length and converge on the retina.
The opposite occurs if the ciliary muscles relax and the curvature of the lens decreases, reducing the
refractive power of the lens and increasing the focal distance.

Key messages:
●​ We 'see' light (i.e., electromagnetic radiation) that is reflected off objects in our environment
●​ Light is refracted by the eye, passes through the pupil and converges on the retina
●​ The retina contains sensory cells - photoreceptors - that convert light energy into neural signals
(phototransduction)

LO2: Describe how visual information is transduced

Photoreceptors located within the retina are responsible for converting light energy into neural signals that
are ultimately transmitted to the nervous system. This process is termed phototransduction which you will
learn about in this chapter.
The most direct pathway for visual information to exit the eye is from photoreceptors to bipolar cells to
ganglion cells. The photoreceptors respond to light, and influence the membrane potential of the bipolar
cells connected to them. This causes ganglion cells to fire action potentials and these impulses propagate
along the optic nerve to the rest of the brain:

Retinal processing is also influenced by two additional cell types:

Horizontal cells - these cells receive input from the photo-receptors, which influences the activity of
surrounding bipolar cells and photo-receptors.
Amacrine cells - these cells generally receive input from bipolar cells and influence surrounding ganglion,
bipolar, and other amacrine cells.
There are three important points to remember: (1) photoreceptors are the only cells involved in visual
perception that respond to light, all other cells in the retina are influenced by light via either direct or
indirect synaptic connections with photo-receptors; (2) The ganglion cells are the only source of output
from the retina; and (3) Ganglion cells are the predominant retinal neurons that fire action potentials. All
other retinal cells depolarize or hyperpolarize, with a rate of neurotransmitter release that is proportional
to the membrane potential, but they do not fire action potentials.

The conversion of electromagnetic radiation into neural signals occurs in the two types of photo-receptors
at the back of the retina, the rods and cones. Each photoreceptor has four regions. An outer segment, an
inner segment, a cell body and a synaptic terminal. The outer segment contains a stack of membranous
disks that contain light sensitive photopigments which absorb light and trigger changes in the
photo-receptor membrane potential. The structure of the outer segment differs between rod
photo-receptors and cone photoreceptors, with rod photoreceptors possessing a long, cylindrical outer
segment that contains many disks. While cone photo-receptors have a shorter, tapering outer segment with
fewer membranous disks. Because rods have a greater number of disks, and thus greater levels of
photo-pigment, they are about 1000 times more sensitive to light than cones. In addition to this, there are
significantly more rods than cones.

The structural differences between rods and cones correlate with important functional differences. Firstly,
only rods contribute to vision in night-time lighting; whereas in daytime lighting, cones do the bulk of the
work. While in intermediate light levels, for example indoor lighting, both rods and cones are responsible
for vision. The other key difference between rods and cones is in their photopigment. While all rods
contain the same type of photopigment, there are three different types of cones which each contain a
different pigment. These pigments are each sensitive to different wavelengths of light and therefore cones
are responsible for colour vision. Finally there are differences in the location of cone and rod
photoreceptors. Most of the 5 million cones are located in the fovea and the proportion diminishes
significantly as you move out into the peripheral retina. In contrast there are no rods in the central fovea
and the ratio of rods to cones in the peripheral retina favour rods.

Photoreceptors are responsible for converting light energy into changes in membrane potential. Watch the
video below which explains how this occurs.

The process of phototransduction in cones is virtually the same as in rods. The only major difference is in
the type of opsins in the membranous disks of the cone outer segments. The cones in our retinas contain
one of three opsins that each respond to different wavelengths of light. Because of this we refer to cones
as detecting different colours. Those cones that we refer to as detecting blue colours actually respond to
short wavelengths of light. Those cones that detect green colours respond to medium wavelengths of light
and those that detect red colours respond to longer wavelengths of light. While we commonly refer to the
cones as blue, green, and red, this can be confusing because different colours are perceived when different
wavelengths of light within the range of sensitivity for that cone are presented. Therefore it is better to
refer to cones by the wavelength of light they can detect: short, medium or long.

Key messages:
●​ Rod photoreceptors are mainly responsible for transduction during nighttime lighting and cone
photoreceptors are mainly responsible for transduction during daytime lighting
●​ Pigment (opsin) in the photoreceptor responds to light causing a reduction in sodium influx
●​ Photoreceptors need to pass information to ganglion cells whose axons form the optic nerve. To
achieve this, photoreceptors form connections with other retinal cells (e.g., bipolar cells) that
form connections with ganglion cells
●​ Ganglion cells are the only retinal cell to fire an action potential
●​ Cone photoreceptors have different pigments that are sensitive to different wavelengths of light
(i.e., long/red, short/blue, medium/green)

LO3: Describe the central pathways involved in the processing of visual information
The axons of retinal ganglion neurons form the optic nerve, which is responsible for communicating
visual information to areas of the nervous system that are responsible for processing and permitting sight.

The visual pathway is shown in the figure below.

This pathway can be summarised as follows: (1) axons of the retinal ganglion cells form the optic nerve
and enter the cranial cavity via the optic canal; (2) the two optic nerves, one from each eye, converge to
form the optic chiasm on the base of the brain; (3) at the chiasm the axons derived from the nasal half of
the two retina decussate and form the contralateral optic tract, while those from the temporal hemiretina
remain ipsilateral; (4) a small proportion of fibres in the optic tract terminate in the pretectal area and
superior colliculus, but the majority continue on and terminate in the lateral geniculate nucleus; (5) from
the lateral geniculate nucleus, nerve fibres project through the internal capsule and form the optic
radiation which terminates in the primary visual cortex, which is located predominantly on the medial
surface of the hemisphere in the region above and below the calcarine sulcus.
Objects in either half of the visual field project images on the nasal hemiretina of the contralateral eye and
the temporal hemiretina of the ipsilateral eye. This means that the left half of the visual field projects onto
the right side of each retina (the nasal side of the right eye and the temporal side of the left eye), and the
right half of the visual field projects onto the left side of each retina (the nasal side of the left eye and the
temporal side of the right eye). The optic nerve carries information from one eye, including both the nasal
and temporal hemiretinas of that eye. This information includes parts of both the left and right visual
fields due to the eye's field of view overlapping. At the optic chiasm, nerve fibers from the nasal
hemiretinas cross to the opposite side (decussate), while the fibers from the temporal hemiretinas continue
on the same side. This arrangement means that after the optic chiasm, all information from the left visual
field (from both eyes) is processed on the right side of the brain, and all information from the right visual
field is processed on the left side of the brain. Therefore, after the decussation at the optic chiasm, each
optic tract, lateral geniculate nucleus, and primary visual cortex receive information relating to the
contralateral half of the visual field only.

As fibres leave the lateral geniculate nucleus, they terminate in the visual cortex such that fibres
representing the lower part of the visual field terminate in the visual cortex above the calcarine sulcus.
Nerve fibres representing the upper part of the visual field pass through the temporal lobe via the Meyer’s
loop before terminating below the calcarine sulcus. The rest of the occipital lobe comprises the visual
association cortex, which is concerned with the interpretation of visual images. Lesions to the primary
visual cortex cause blindness in the corresponding part of the visual field. Damage to the visual
association cortex cases deficits in visual interpretation and recognition.

The visual pathway can be quite complicated to understand, especially given the crossing over of fibres
and the inversion of information coming from upper and lower visual fields. To help consolidate your
understanding of this pathway, now watch the video below.

Key messages:
●​ Ganglion cells give rise to the optic nerve
●​ Nerve fibres from the nasal hemiretina cross at the optic chiasm whereas nerve fibres from the
temporal hemiretina remain on the same side (ipsilateral)
●​ After crossing at the chiasm, nerve fibres form optic tract and terminate in the lateral geniculate
nucleus, which sends projections via the optic radiations to the primary visual cortex
●​ Optic tract, lateral geniculate nucleus and primary visual cortex receive information about the
contralateral visual field

LO4: Describe the peripheral structures involved in hearing

Hearing relates to the sensation of sound, which is simply audible variations in air pressure. When we
hear a sound, we are simply hearing audible variations in air pressure.

A low frequency sound travels at the same speed as a high frequency sound but has fewer low frequency
oscillations in air pressure compared with a high frequency sound. Our auditory system responds to
sounds over a wide range of frequencies ranging from 20 to 20000 Hz. Another aspect of sound that
relates to frequency is pitch. Sounds that have a low tone, or pitch, create changes in air pressure that
oscillate at a low frequency. For example, a low key on a piano. Whereas sound that have a high tone, or
pitch, create changes in air pressure that oscillate at a high frequency - for example, a child screaming.
The last property of sound waves that is important to know about is intensity. This property relates to the
amplitude of the changes in air pressure associated with a particular sound. Low intensity sounds produce
changes in air pressure that have a small amplitude whereas high intensity sounds produce changes in air
pressure that have a large amplitude.

While we have different components of sound i.e. frequency and intensity, we rarely hear sounds that are
characterised by one particular frequency or intensity. In reality, we hear complex sounds that are made up
of different frequency waves that have different intensities. This gives the unique tonal qualities that we
associate with different sounds e.g. a musical instrument or the human voice. Watch this short video to
consolidate your understanding of the properties of sound.

To understand how these variations in air pressure are translated into nerve signals that reach the brain,
you need to know the structure of the ear. There are three overall divisions of the ear: the outer ear, middle
ear and inner ear. The outer ear is made up of all structures from the pinna through to the tympanic
membrane. The middle ear is a small air-filled chamber that is made up of the tympanic membrane and
the ossicles. The oval window marks the beginning of the inner ear.

Interact with the image below to learn more about the structure of the ear.

●​ Pinna: This is the visible convoluted portion of ear that is made up primarily of cartilage covered
by skin. This convoluted structure acts as a funnel to help collect sounds from a wide area and
 assists in localising sounds. The shape of the pinna facilitates the collection of sounds coming
from in front of the ear rather than from behind. Because of this, we are able to localise sounds
that are in front of us more easily. This is particularly the case for humans as we have a pinna that
are more or less in a fixed position, which is why we find it easier to hear someone speak when
they are looking at us. This contrasts dogs, for example, who have muscular control of their pinna
and can change the orientation of their pinna to towards the source or a sound.
●​ The tympanic membrane is also known as the eardrum. This structure separates the outer ear from
the middle ear and vibrates when sound waves hit it. The tympanic membrane is responsible for
transferring these vibrations to structures in the middle ear.
●​ The auditory canal serves as the entrance to the inner ear. The auditory canal extends for
approximately two and a half centimetres inside the skull and ends at the tympanic membrane.
●​ Connected to the medial surface of the tympanic membrane is a series of bones called ossicles.
The ossicles are a series of three very small bones called the malleus, incus and stapes. The
ossicles are connected at one end to the tympanic membrane and at the other end to a second
membrane that covers a hole in the bone of the skull called the oval window. The ossicles act as a
conduit between the tympanic membrane and the oval window.
●​ The tympanic membrane is almost conical in shape, with the point of the cone extending into the
middle ear and attaching to the malleus. The malleus forms a rigid connection with the incus
which itself has a flexible connection with the stapes. The flat bottom, or footplate, of the stapes
moves in and out against the oval window like a piston. Therefore when the tympanic membrane
vibrates in response to sound waves hitting it this causes the malleus to move. This causes the
incus to move and thereby the stapes to move against the oval window. Therefore these bones
transfers movement that has been produced in the tympanic membrane by the sound wave to the
oval window.
●​ The oval window marks the beginning of the inner ear.
●​ The cochlea is a fluid filled structure that is responsible for converting changes in movement in
the oval window into a change in nerve activity within the cochlear branch of the
vestibulocochlear nerve.

We can summarise the conversion of a sound wave, or change in air pressure, into a nerve signal in five
simple steps:

Step 1. Sound waves travel down the auditory canal and produce movement in the tympanic membrane

Step 2. Movement in the tympanic membrane causes the ossicles to move

Step 3. Movement in the ossicles results in movement within the membrane of the oval window

Step 4. The motion of the oval window causes the fluid in the cochlea to move

Step 5. Movement of fluid in the cochlea causes a change in activity in the sensory neurons of the
vestibulocochlear nerve.
Key messages:
●​ We 'hear' variations in air pressure
●​ Sound waves are funnelled by the outer ear where they cause movement in the tympanic
membrane
●​ Movement in the tympanic membrane causes the ossicles to move (piston action)
●​ Movement of the ossicles amplifies the sound waves and causes movement of the oval window
●​ Movement of the oval window causes fluid movement surrounding the cochlea (located in inner
ear) that activates sensory cells

LO5: Describe how auditory information is transduced

The inner ear contains two major structures: the cochlea and the semicircular canals.

The cochlea is the only component of the inner ear that is relevant to hearing and is spiral in shape and
wraps around a bony core. The walls of the cochlear are made of bone and the cochlea is hollow on the
inside. At the base of the cochlea are two membrane-covered holes: the oval window and the round
window.
If you were to take a cross section of the cochlea you would see the spiral shape of the cochlear creates
three fluid-filled chambers. These are the scala vestibuli, scala media and scala tympani:

The scala vestibuli is separated from the scala media by the Reissner's membrane and the scala tympani
separated from the scala media by the basilar membrane. The organ of Corti sits on top of the basilar
membrane and contains the auditory receptor neurons. On top of the organ of Corti is the tectorial
membrane.

The mechanical transformation of sound energy that occurs in the middle and inner ear need to be
transferred into a neural signal. This occurs within the organ of Corti which contains the auditory receptor
cells (also referred to as hair cells). These cells are not neurons because they do not have an axon, and are
incapable of generating an action potential. Instead, they are a specialised epithelial cell. At the tip of each
of hair cell are a series of stereocilia, with each hair cell possessing between 10 and 300 stereocilia. The
movement or bending of these stereocilia represents the critical event in the transduction of sound into a
neural signal.

The hair cells of the organ of Corti are squashed between the basilar membrane and a thin sheet of tissue
called the reticular lamina. Within the organ of Corti are another group of cells called the rods of Corti.
These rod cells provide structural support and act to define the boundary of two different categories of
hair cells. Hair cells that are located between the modiolus and the rods of Corti are called inner hair cells.
There are about four and a half thousand inner hair cells that form a single row. Hair cells further away
from the rods of Corti are called outer hair cells. There are three rows of these outer hair cells which total
between 12000 and 20000 in humans.

The stereocilia of both inner and outer hair cells extend above the reticular lamina into the endolymph of
the scala media. The tips of the stereocilia of outer hair cells end in the tectorial membrane whereas for
inner hair cells they stop just below the tectorial membrane. Both types of hair cells form synapses on
neurons whose cell bodies are located in the spiral ganglion within the modiolus. These ganglion cells are
bipolar cells and have neurites that extend to the base and sides of the hair cells where they receive
synaptic input from the hair cell. The axon from the spiral ganglion cells enter the cochlear branch of the
vestibulocochlear nerve and project to the cochlear nuclei within the medulla.
When the stapes hits the oval window, this will cause a movement in the basilar membrane and therefore
the hair cells of the organ of Corti. When the basilar membrane move up so does the reticular lamina,
positioning it closer to the modiolus whereas when the basilar membrane moves down, the reticular
lamina also moves down pulling it away from the modiolus. This creates a movement of the reticular
lamina relative to the tectorial membrane. Because the tectorial membrane holds the tips of the stereocilia
of outer hair cells, lateral motion of the reticular lamina relative to the tectorial membrane will cause the
tips of the stereocilia to move laterally away from the direction of movement of the reticular lamina. For
example if the reticular lamina moved to the right, the tips of the stereocilia of the outer hair cells would
be bent to the left relative to the body of the hair cell. The tips of the stereocilia of inner cells also bend,
but because their stereocilia are not held in place by the tectorial membrane, this movement is likely
produced by the movement of endolymph within the scala media.

The bending of the stereocilia of the hair cells causes the opening or closing of mechanically gated ion
channels. When the stereocilia move in a direction that opens these ion channels, inward flow of ions
occurs generating a potential within the hair cell. The exact processes involved are still unclear but the
working theory is that when the stereocilia move in toward the kinocilium, which is the tallest of the
stereocilia of each hair cell and is usually the stereocilia located farthest to the right or left, this causes
tension on mechanically gated ion channels causing them to open and potassium to enter the hair cell.
Movement in the opposite direction away from the kinocilium relieves the tension on these ion channels,
allowing them to spend more time closed which reduces potassium influx. The influx of potassium into
the hair cell causes it to depolarise, which in turn opens voltage-gated calcium channels. This allows the
entry of calcium into the hair cell, which triggers the release of glutamate contained within vesicles at the
base of the hair cell. The glutamate that is released then activates spiral ganglion fibres that form synapses
with the hair cells, which causes an action potential within the spiral ganglion neuron.

But hang on – the opening of potassium channels causes depolarisation of the hair cell. This is not what
happens in a neuron. Why do you think that is?
The reason that potassium causes depolarisation of hair cells is because the concentration of endolymph is
particularly high in potassium and therefore the equilibrium potential for potassium in hair cells is 0 mV,
while -80 mV in neurons. Therefore entry of potassium into the hair cell causes the membrane potential to
increase and the hair cell to depolarise. The other reason is the presence of endocochlear potential which
creates a large gradient across the stereocilia membrane.

If you have found the information above conceptually difficult to understand, watch the video below
which contains an animation describing how sound waves can be converted into neural signals. This
video is at a level below what you need to understand this content at, but may be helpful for you to watch
before then revisiting the content above.

Key messages:
●​ The cochlea is the only component of the inner ear important for hearing
●​ Hair cells within the Organ of Corti are the sensory receptor cells responsible for hearing
●​ Bending of the cilia on the hair cells causes potassium influx and depolarisation of the hair cell
●​ Hair cells are not neurons but in response to depolarisation release glutamate that then activates
spiral ganglion neurons

LO6: Describe the central pathways involved in the processing of auditory information
In order for us to perceive and interpret sound, auditory information must reach the nervous system. This
occurs via a pathway from starts within the cochlea and ultimately terminates within the auditory cortex.
This pathway is presented in the following image.

This pathway can be summarised as follows:

 1.​ The cochlear nerve fibres make dendritic contact with hair cells of the organ of Corti within the
inner ear. The cell bodies of these fibres lie within the cochlea and are collectively called the
spiral ganglion. The cochlear nerve leaves the inner ear and joins the brainstem at the level of the
rostral medulla, and then its fibres bifurcate and synapse in the dorsal and ventral cochlear nuclei.
2.​ Once within the brainstem, a pathway that is fine-tuned to auditory information becomes
activated. This pathway is slightly complicated, but regardless has one goal. To get to the
thalamus and from there to the cerebral cortex. The first stop once the auditory information has
been transmitted to the cochlear nucleus is the pons. From the cochlear nuclei, nerve fibres ascend
into the pons. Some of these nerve fibres cross to the other side at the level of the trapezoid body
and some remain on the same side. Within the pons some of these nerve fibres terminate within
the superior olivary nucleus.
3.​ After the superior olivary nuclei, ascending auditory information joins the lateral lemniscus and
ends up in the inferior colliculus which is located in the midbrain. Those fibres that left the
cochlear nucleus but did not terminate within the superior olivary nucleus also terminate within
the inferior colliculus.
4.​ From the inferior colliculus the next stop is the thalamus and axons from the inferior colliculus
terminate in the medial geniculate nucleus of the thalamus.
5.​ The final step in the ascending auditory pathway consists of axons that originate in the medial
geniculate nucleus and pass through the internal capsule to the primary auditory cortex of the
temporal lobe. This is located predominantly in Heschl’s gyri, which are situated on the dorsal
surface of the superior temporal gyrus and, therefore, are largely hidden within the lateral fissure.

Key messages:
●​ The cochlea nerve terminates in the cochlea nucleus
●​ Information is relayed from the cochlea nucleus to the primary auditory cortex via the superior
olivary nucleus (not all fibres terminate here), inferior colliculus and medial geniculate nucleus
●​ Some nerve fibres decussate at the level of the pons and some nerve fibres remain ipsilateral (i.e.,
the auditory cortex receives input from both ears)