The Sensory Systems of the Human Body

The sensory organs enable us to interact with our surroundings and perceive
things outside our bodies. Their functions are fascinating, but the topic seems
vast and complex during medical studies. However, all sensory organs follow a
few fundamental principles in their structure and function.

The Fundamentals of Sensory Perception

An adequate stimulus (i.e. a threshold energy input that causes a reaction of the sensory
organ) triggers the stimulation of the organ, such as light in the eye. The stimulation is
relayed via peripheral pathways (in this example, the eye and optic nerve) and processed
in the respective brain areas optic nerve and corpus geniculatum laterale). This means
that there are 2 central processes:

Transduction: the conversion of a stimulus into a receptor potential

Transformation: the conversion of receptor potentials into action potentials
for forwarding to the processing brain areas

Here, 4 traditional senses—sight, hearing, smell, and taste—, as well as the sense of
balance, will be discussed.
Sensory adaptation
Sensory adaptation is defined as the change over time in the responsiveness of the
sensory system to a constant stimulus. This allows the brain to tune out unimportant
information. The system is designed to respond to changing, not constant, information.
Nociceptors (pain receptors) do not adapt under any circumstances.

Sensory Pathways
Sensory receptors are sensory nerves that are either cells or nerve endings. They can
detect both internal (interoceptor) and external (exteroceptor) stimuli. Activation initiates
signal transduction by creating graded or action potentials. Sensory pathways begin with
the receptor and continue through to ganglion cells and, finally, to the spinal cord.
Types of Sensory Receptors
Sensory receptors are specific to only 1 sensory modality.

Type of sensory receptor Description

Auditory hair cells detect mechanical movement of cells
Mechanoreceptors from fluid in the ear.
Olfactory receptors detect airborne chemicals, allowing
smell
Chemoreceptors Taste (gustatory receptors) detect chemicals on the
tongue.
Nociceptors Receptors that respond to tissue injury, leading to pain
Thermoreceptors Receptors that respond to temperature
Photoreceptors Activation of rod and cone cells in the eye
The Sense of Sight

The optical apparatus

Image: Structures of the Eye, by Phil Schatz.

An object has to be projected onto the ocular fundus for us to see it. For this purpose,
light is refracted at the air–ocular border of the globe, and the image of the object is
projected on the retina upside down and scaled down. The total refractive power of the
eye is 58.8 diopters (D; distance-accommodating power of cornea, anterior chamber, and
lens).

A higher refraction can be achieved with greater curvature of the lens, and thus, the
refractive power can be adjusted according to the distance from the object. The distance
between near and far-off objects (i.e. between the minimum and maximum distances that
allow for a sharp image) is also referred to as the range of accommodation.

The difference between the respective diopter values forms the amplitude of
accommodation. This value cannot exceed a maximum of 14 D. In addition, the pupil
contributes to the intensification of the projected object by blanking out marginal rays.

The retina
The retina lines the ocular bulb. The pars optica, which consists of the stratum
pigmentosum and the stratum nervosum, is located in the rear area of the bulb. The pars
caeca is located in the anterior area and consists of stratum pigmentosum only. From
outside to inside, the retina can be divided into the following 10 layers:

Stratum pigmentosum

Pigment epithelium: supply of retinal cells

Stratum nervosum

Photoreceptors: transduction of light signals

External glia limitans
 External granular layer: cell nuclei of photoreceptors (first neuron)
External plexiform layer: first synapse
Internal granular layer: cell nuclei of bipolar cells (second neuron)
Internal plexiform layer: second synapse
Ganglion cell layer: cell nuclei of ganglia (third neuron)
Nerve fiber layer
Internal glia limitans

The internal granular layer contains additional horizontal cells, amacrine cells, and Müller
glia cells, which have a modulating effect on the processing of signals and can amplify
the contrast.

Photoreceptors – rods and cones

There are 2 types of photoreceptors. The rods are responsible for providing light/dark
vision, also referred to as scotopic vision, and the cones are responsible for color vision,
also called photopic vision. The cones can be divided into 3 subtypes according to the 3
spectral colors. The human eye has approximately 120 million rods and approximately 6
million cones.

Phototransduction of the eye

The sensory cells contain the visual pigment 11-cis retinal, which absorbs light and
changes its conformation to all-trans-retinal. As a result, the sodium channels of the cell
membrane close, leading to a hyperpolarized signal at the glutamate photoreceptor
synapses.
 Image: Retinal Isomers, by Phil Schatz. License: CC BY 4.0

The signals are processed in a network of bipolar cells, amacrine cells, and horizontal
cells and are interconnected with ganglion cells. Therefore, rods and cones are secondary
sensory cells.

The macula lutea is a yellow oval spot at the center of the retina. It is the part of the
retina that is responsible for sharp, detailed central vision (also called visual acuity), and
it contains a very high concentration of cones. The fovea centralis, the center of the
macula lutea, consists only of cones. This is the site of clearest vision.

The papilla nervi optici is located medial to the macula lutea and marks the exit of the
optic nerve. Because of missing photoreceptors in this area, vision is not possible here.
Therefore, this area is also referred to as the ‘blind spot.’ This spot is located laterally in
the field of vision.

The visual pathway

The efferent fibers of the ganglion cells form the optic nerve. This nerve exits the eye at
the papilla nervi optici, passes through the orbital foramen, and enters the cranial
cavity. The optic nerves of both eyes join in the chiasma opticum.

Here, the medial parts of the retina cross to the opposite side. The temporal parts remain
ipsilateral. Therefore, the ipsilateral temporal fibers and the contralateral medial fibers
run together starting from the chiasma opticum.
The tractus opticus projects onto the corpus geniculatum laterale inside the
thalamus.

From here, the visual pathway then proceeds to the visual cortex via the broad radiatio
optica. The visual cortex is divided into primary and secondary visual cortex. The
primary visual cortex is responsible for making us conscious of visual impulses, which are
then analyzed in the secondary visual cortex.

Paresis of the visual pathway

Symptoms can often lead to conclusions about the location of lesions of the visual
pathway. The following examples illustrate this:

Blindness in 1 eye: The lesion is probably located in the nervus opticus

because this nerve houses all fibers of 1 eye.
Homonymous hemianopia: The same side of the visual field is affected in
both eyes; the lesion is located behind the chiasma opticum.
Heteronymous hemianopia: The opposite sides of both eyes are affected,
i.e. the patient has a vision as if he had blinkers on, meaning that both
temporal visual fields are defected; the lesions are located inside the chiasma
opticum.
Hemianopia with intact optical reflexes: A lesion of the corpus

geniculatum laterale. The collaterals of the reflex pathways in the

mesencephalon exit before the corpus; thus, the reflexes remain intact despite
the loss of vision.
Minor scotomas: The lesion is probably located in the optic radiation because

only particular areas are damaged due to the broad fragmentation.

The Auditory System

The auditory sense converts acoustic waves—meaning fluctuations in pressure in our
surroundings—into electrical signals and, consequently, perceives tones, sounds, and
noises.

Sound pressure level and volume level

The most important measurement unit in this context is the sound pressure level,
measured in decibels (dB). This describes the acoustic pressure in relation to the auditory
threshold.

The auditory threshold is the minimal acoustic pressure at which a tone of a specific
frequency can be heard. The relationship of the sound pressure level to a specific value
(auditory threshold) makes it a measurable value that can be calculated as follows:

sound pressure level = 20 × lg (p1 / p),

where p is the reference value meaning the auditory threshold.

The volume level is not the same as the sound pressure level. The volume level
describes the subjective perception of the sound volume and is based on the phon scale.
The phon value equals the sound pressure level at a frequency of 1 kHz. This means that,
for instance, values of lower frequency need a higher sound pressure level than 1 kHz to
be perceived as equally loud. These differences are shown with the help of isophones.
Structure of the ear

Image: Structures of the Ear, by Phil Schatz. License: CC BY 4.0

The ear is divided into the outer ear, middle ear, and inner ear. Sound passes through the
outer ear to the tympanic membrane. There, the auditory ossicles—malleus, incus, and
stapes relay the sound to the oval window of the inner ear.

The middle ear is responsible for impedance adjustment, which allows a sound
transmission of 60%. Without this adjustment, 98% of the sound would be reflected. The
adjustment occurs via 3 mechanisms:

1. The leverage effect of the ossicles amplifies the sound force onto the oval window,
contrary to the force on the tympanic membrane.
2. The smaller area of the stapes compared with the tympanic membrane leads to an
increase in pressure.
3. Compared with the oscillations of the tympanic membrane, the stapes moves more
slowly. Because impedance = pressure/velocity, a decrease of velocity results in
an increased impedance.
 Image:

The inner ear – Structure of

the cochlea
The cochlea is the part of the inner ear responsible for hearing. The cochlea consists of
several tubes (scala tympani, scala media, and scala vestibuli) and the organ of Corti,
which are coiled into the cochlea.

The scala tympani and scala vestibuli are filled with sodium-rich perilymph. The scala
media contains potassium-rich endolymph, and it is bordered by the Reissner membrane
at the scala vestibuli and by the basilar membrane at the scala tympani, as well as by the
stria vascularis.

The organ of Corti contains the actual sensory cells, the outer and inner hair cells, and is
covered by the tectorial membrane. There are 3 rows of outer hair cells that touch the
tectorial membrane with their longest stereocilia, as well as 1 row of inner hair cells
whose stereocilia do not touch the membrane but, rather, are deflected by the
displacement of the endolymph (hydrodynamic coupling) and are thus stimulated.

Transduction of the cochlea

The traveling wave inside the scala vestibuli caused by the stapes leads to a frequency-
specific maximum deflection of the basilar membrane at the corresponding location of
the cochlea. This results in a shearing motion of the tectorial membrane against the
organ of Corti and the hair cells. High-pitched tones are reflected near the oval window,
low-pitched tones in the direction of the helicotrema.

The deflection of stereocilia creates a receptor potential and leads to mechanoelectrical

transduction. The endocochlear potential, which is created through the higher potassium
concentration of the endolymph compared with the perilymph, is essential for the
creation of potential. This potential is built by the stria vascularis, which actively
transports potassium ions into the endolymph.

The deflection of the stereocilia opens potassium channels. Potassium from the
endolymph flows into the cell and depolarizes it. Thereafter, calcium channels open, and
the calcium influx results in a glutamatergic synaptic transmission of the potential to the
first neuron of the sound conduction system.
The auditory pathway
The perikaryon of the first neuron is located inside the ganglion spiral. These bipolar cells
transmit the potentials to the cochlear nuclei of the medulla oblongata. Their extensions
form an important part of the vestibulocochlear nerve. The tonotopic organization of the
cochlear coil is preserved.

Part of the fibers cross inside the trapezoid body, while another part continues on
uncrossed. The crossed part is interconnected in the nuclei olivares superiores, which is
essential for directional hearing. Contralaterally, both strands continue together to the
inferior colliculi as the lemniscus lateralis. After dispensing with some of the smaller
branches and back-crossings, the main part of the fibers continues to the corpus
geniculatum mediale and, from there, to the primary auditory cortex as acoustic waves.

Therefore, the auditory cortex receives information from both cochleae. This has a
positive effect in the case of unilateral injury of the auditory pathway and on directional
hearing. The primary auditory cortex is responsible for making us conscious of sounds.
The meaningful connection to words or melodies takes place in the secondary auditory
cortex.

The Vestibular System

The vestibular apparatus, which is responsible for the sense of balance or sense of
equilibrium, is located in the inner ear along with the cochlea. It consists of 3 semicircular
ducts and 2 otolith organs, which together facilitate spatial orientation and registration of
movements.

Structure of the Vestibular Apparatus

The structure of the vestibular apparatus is similar to that of the cochlea. The ducts of the
vestibular apparatus are also filled with endolymph, and the sensory cells are also hair
cells. However, unlike the hair cells of the cochlear, these develop cilia and several
stereocilia that are connected via tip links.

They are covered by a gelatinous mass. Inside the semicircular ducts, this mass, which
contains mucopolysaccharide, is called the cupula. In addition, this mass contains small
calcium carbonate crystals inside the otolith organ and is, therefore, called the otolithic
membrane.

Transduction of the vestibular apparatus

The tough cupula/otolithic membrane is shifted against the sensory cells through
acceleration, deceleration, or rotating of the head. Just like inside the cochlea, the shifting
leads to shear movement and a deflection of cilia and stereocilia and causes a receptor
potential.

The transduction process is the same in the semicircular ducts and the otolith organs.
However, because of their anatomic differences, the 2organs measure different
movements.

Translational motion: Otolith organs measure acceleration and deceleration. Macula

sacculi measure vertical translational motions, and macula utriculi measure horizontal
motions.
 Image: Linear Acceleration Coding, by Phil Schatz. License: CC BY 4.0

Rotational motion: The endolymph in the semicircular ducts is usually arranged

circularly. Because of inertia, the fluid is shifted against the sensory epithelium during
rotations, and thus the cilia of the cells are deflected. Cilia are built in a way that, if
deflected medially toward the utricle, they will cause a potential. This means that when
the head is rotated to the left side, the fluid in the horizontal semicircular ducts shifts to
the right, which leads to activity in the left semicircular ducts and afferent nerves.

Image: Rational Coding by Semicircular Canals, by Phil Schatz. License: CC BY 4.0

Vestibular pathway
The generated potentials are transferred from the first neuron as part of the
vestibulocochlear nerve to the vestibular nuclei inside the rhombencephalon and to
the second neuron. From this point on, the crossed and uncrossed pathways continue on
to the nucleus ventralis posterior of the thalamus. The impulses are then transmitted
to the vestibular areas of the cerebrum.
The central vestibular system
The information from the vestibular apparatus is continuously offset by somatosensory
information from the brain and neck area, as well as from other joints, for the central
nervous system to acquire information about the posture of the entire body.

The 4 vestibular nuclei involved are the nucleus superior of Bechterew, nucleus inferior of
Roller, nucleus medalis of Schwalbe, and nucleus laterals of Deiters. This is also true for
muscular reflexes activated to maintain body balance.

Particularly interesting are the vestibulo-ocular reflexes, which connect the vestibular
apparatus with the eye muscles. This is, for instance, important for rotational
movements. Vestibular nystagmus is a slow, vestibular-induced eye movement followed
by a fast return movement.

Example: If a person sitting in a chair turns to the right, the sensory cells in the right
semicircular duct are activated. They project via vestibular nuclei to the nuclei of the eye
muscle and cause an eye movement to the left. Vision stabilization follows. The fast
return movement is mediated centrally and follows the turning movement.

The Olfactory Sense

Structure and function of the olfactory mucosa

This area of olfactory perception (regio olfactoria) is strongly underdeveloped in
humans and covers only the upper nasal concha and nasal septum. The stratified
olfactory epithelium consists of 3 cell populations.

Supporting cells, basal cells and olfactory cells

Resting on the basal lamina of the olfactory epithelium, basal cells are stem cells capable
of division and differentiation into either supporting or olfactory cells. The constant
divisions of the basal cells lead to the olfactory epithelium being replaced every 2–4
weeks.

The olfactory cells are the primary bipolar sensory cells of the olfactory organ. They form
long cilia (olfactory cilia) that bind molecules of breathable air with the help of
chemoreceptors and thereby stimulate the sensory cells.

Each sensory cell can perceive only 1 olfactory quality, such that 1 olfactory quality can
be perceived by tens of thousands of sensory cells. Humans have approximately 350
different receptors and can differentiate between 7 typical kinds of smells.

Primary sensory cells transmit the stimulation directly to the central nervous system via
their axons and without any interconnection. The axons, or fila olfactoria, are bundled
in the olfactory nerve and pass directly into the bulbus olfactorius through the lamina
cribrosa of the ethmoid bone plate. The bulbus olfactorius is considered a part of the
central nervous system located in front of it. The first interconnections take place here.

Bulbus olfactorius
The endings of the fila olfactoria, together with the dendrites of the mitral cells, form
glomeruli inside the bulbus olfactorius, which are the smallest functional units of the
olfactory organ. The first synapse of the olfactory system is located there. During this
process, convergence occurs: More than 1000 axons of sensory cells project onto the
dendrites of 1 mitral cell.

Periglomerular cells and granular cells of the glomeruli can modulate the signal. A lateral
inhibition facilitates periglomerular cells to amplify the signal of the stimulated
glomerulus and to define it better against weaker neighboring signals.

The olfactory pathway – tractus olfactorius

Approximately 30,000 axons of mitral cells exit the bulbus as the so-called tractus
olfactorius, which splits into a main branch and a side branch. The main branch crosses
at the anterior commissure to the bulbus of the opposite side of the brain, whereas the
side branch projects onto the olfactory bulb.

The olfactory bulb is located in the paleocortex and consists of many olfactory projection
fields. Information is sent from here to the neocortex and the cortex praepiriformis, as
well as to the limbic system. From the limbic system, it is sent to the nuclei areas of the
hypothalamus and the formatio reticularis.

Image: The Olfactory System, by Phil Schatz. License: CC BY 4.0

The Sense of Taste

Structure of the organ of taste

Taste is perceived via the tongue, on which the appropriate structures are located. There
are 3 different types of taste papillae:

Papillae fungiformes: 200–400, distributed on the entire surface

Papillae foliatae: 15–20, located on the posterior margin in consecutive rows
Papillae vallatae: 7–12, located at the border to the tongue base

The taste papillae contain the taste buds. There are 2,000–4,000 buds, each having
approximately 10–50 sensory cells. The taste buds develop a cavity filled with fluid. The
sensory cells and their microvilli extend into this cavity. The microvilli contain the actual
taste receptors.

Transduction of taste
Here, a chemical stimulus is converted into an electrical signal. The chemical
substances/relationships always cause a depolarization of the sensory cell through
different receptors or channels, which leads to the release of transmitters and the
activation of innervating nerves. Therefore, sensory cells of taste are secondary sensory
cells.

We can differentiate between 4 different qualities of taste: sweet, sour, salty, bitter.

A sensory cell can perceive just 1 quality of taste or all 4 of them but with a predefined
ranking order of the 4 qualities.

Gustatory pathway
The papillae are innervated by the nervus glossopharyngeus and the nervus vagus.
These nerves proceed to the nucleus tractus solitarii of the brain stem. The
information is switched over to the second neuron and transferred ipsilaterally to the
third neuron in the nucleus parabrachialis of the formatio reticularis.

This neuron projects into the contralateral nucleus ventralis posterior of the thalamus.
The thalamus transfers the information to different areas of the brain. There, we become
conscious of the taste and of links to other perceptions, i.e. the sense of smell.