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Chapter 10 - Sensory

General Principles
The afferent neurons of the peripheral nervous system (PNS) pick up information received by sensory
receptors.
Stimuli detected inside the body travel to the CNS by means of visceral afferents and are detected by
visceral receptors. We are not aware of these receptors.
Visceral receptors include:
Chemoreceptors that monitor O2 and H+ levels in major blood vessels.
Baroreceptors that monitor blood pressure.
Mechanoreceptors that monitor the degree of stretch or distension in hollow organs.
We are aware of stimuli from the external environment. These receptors fall into two broad systems:
1. Somatosensory system that include various receptors in the skin (somesthetic sensations), and
proprioceptors, or receptors that provide information about the position of the limbs and the body.
2. Special sensory system that include for vision, hearing, equilibrium and balance, taste and
smell.
Receptor Physiology
Receptors respond to stimuli which are different forms of energy including chemical gradients,
pressure, temperature, sound waves and photons. The specific form of energy of a stimulus is its
modality. According to the law of specific nerve energies sensory receptors are designed to respond best
to specific modalities. The modality to which the receptor responds to best is the adequate stimulus.
Sensory Transduction
Transduction (trans - across; duct - lead) is the conversion of one form of energy into another.
Sensory transduction converts the energy of the stimulus into a receptor or generator potential. The
sensory receptor may be a specialized structure at the end of a peripheral neuron or a separate cell that
communicates with an afferent neuron by means of a chemical synapse.

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Receptor Adaptation
Adaptation is a decrease in the size of the receptor potential with a constant stimulus. Slowly
adapting or tonic receptors show little adaptation in response to a prolonged stimulus. Rapidly
adapting or phasic receptors adapt quickly. These receptors detect changes in stimuli intensity.

Slowly adapting receptors are better at coding the intensity of a stimulus for its entire duration. Rapidly
adapting receptors code changes in stimulus intensity better but not the duration.
Sensory Pathways
The specific pathway that transmits information about a specific modality is a labeled line.
Stimulation of a labeled line only produces a sensation of its modality no matter what type of energy
produces the action potentials. Pathways for different modalities terminate on different places of the
cerebral cortex. A pathway begins with a sensory unit.
A sensory unit is a single afferent neuron and all receptors associated with it. All the receptors of an
afferent neuron can respond to an adequate stimulus in an area called a receptive field.

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In a generalized pathway, the afferent neuron of the sensory unit is the first in a chain of neurons that
relays information to the CNS (spinal cord and brain). The afferent neuron is the first order neuron. It
synapses with a second order neuron that synapses in turn with a third order neuron in the thalamus.
The third order neuron guides the impulse to the sensory cortex where it is perceived.

Sensory Coding
Stimulus type
Stimulus type is coded by the receptor type and the pathway activated when the stimulus is applied.
The perception of a stimulation often results from the simultaneous activation of more than one sensory
pathway. The final perception results from integration in the brain of information from various sensory
systems. Pinocchio illusion
Intensity and Duration
Intensity is coded by the frequency of action potentials (frequency coding) and the number of receptors
activated (population coding). Stronger stimuli produce a higher frequency of actions potentials.

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Intensity is interpreted by population coding because the more intense the stimulus, the greater the number of
receptors stimulated. This results from either a single sensory unit (sensory afferent neuron) sending more action
potentials to the CNS or more sensory units sending action potentials to the CNS.

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Location
The receptive fields of specific afferent neurons code for stimulus location. Tactile receptors in the skin
illustrate this. The precision of stimulus location is acuity. Acuity depends on:
1. Size and number of receptive fields.
The smaller the receptive field the greater the acuity. For example, tactile acuity can be measured by
two-point discrimination or the ability to perceive two fine points pressed against the skin as two points
and not as one. The smaller the receptive fields the smaller the distance between two points of stimulation
can be and still be discriminated. Areas with smaller receptive fields (lips, fingertips) have better
two-point discrimination than those with larger fields (back, shoulder).
2. Lateral inhibition
Lateral inhibition occurs when a strong stimulus applied to the receptive field of one neuron causes
that neuron to inhibit transmission of signals by neurons with neighboring receptive fields. Lateral
inhibition increases acuity because it increases the contrast of signals in the nervous system. In other
words, the difference between the strength of the signals coming from the central neuron in the affected
field and the neurons on the periphery is increased as the information is processed in the central nervous
system.

Somatosensory System

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Body sensations such as pressure, temperature, pain and body position. Somatosensory receptors
include:
Body-sense sensations
Proprioceptors are receptors that give information about body position. These receptors are located
in muscles, tendons, ligaments, joints and skin.
Somesthetic sensations (senses associated with the surface of the body).
Mechanoreceptors detect pressure, force and vibration. These include:

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Merkel's disks and Meissner's corpuscles in the superficial layer of the skin and,
hair follicle receptors, Pacinian corpuscles and Ruffini's endings in deeper layers.
Thermoreceptors respond to temperature of receptor endings themselves.
Warm receptors respond to temperature between 30o C and 45o C with action potentials
increasing as temperature increases.
Cold receptors respond to temperatures between 35o C and 20o C with action potentials
increasing as the temperature falls.
Both warm and cold receptors respond rapidly to temperature changes and show rapid
adaptation. The brain uses the relative changes in the responses of hot and cold receptors to interpret
the temperature of the environment.
Nociceptors transduce harmful stimuli that we perceive as pain. These consist of free nerve endings.
There are three types of nociceptors:
Mechanical - respond to intense mechanical stimuli.
Thermal - respond to intense heat.
Polymodal - respond to a variety of stimuli including mechanical, intense heat and chemicals
released from damaged tissue.
Somatosensory Cortex
Somatosensory sensations from all parts of the body go to the somatosensory cortex. All sensory
information (except olfaction) goes to that thalamus and is relayed from there to the cortex. Also, all
sensations from one side of the body goes to the thalamus and cerebral cortex on the opposite side. There
are two major somatosensory pathways:

1. Dorsal Column-Medial Lemniscal Pathway

This pathway transmits information from mechanoreceptors and proprioceptors. First order
neurons from the periphery enter the spinal cord through the dorsal root. The main axons ascend the
spinal cord in the ipsilateral dorsal column and end in the dorsal column nuclei in the medulla
oblongata where they synapse with second-order neurons. Second order neurons cross over to the

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contralateral side of the medulla in the medial lemniscus and ascend to the thalamus. Third order
neurons transmit information from the thalamus to the somatosensory cortex.

2. Spinothalamic Tract
This pathway transmits information from thermoreceptors and nociceptors. First order neurons
from the periphery enter the spinal cord through the dorsal root and may ascend or descend (a few
spinal segments) along Lissauer's tract before synapsing with second order neurons in the dorsal
horn. Second order neurons cross to the contralateral side of spinal cord and ascend in the
anterolateral quadrant of the spinal cord through the brainstem to the thalamus. Third order
neurons in the thalamus ascend to the somatosensory cortex.
Pain Perception Article about "Feeling No Pain"
Pain is important because it helps us to avoid damaging stimuli and, when tissue damage occurs, helps
us prevent further damage.
Pain Response
Stimulation of nociceptors causes pain perception but also causes
1. Autonomic responses
2. Fear and anxiety
3. Reflexive withdrawal
There are two types of pain:
1. Fast pain is perceived as an easily localized, sharp, pricking sensation and is transmitted by Ad
fibers (12-30 m/sec).
2. Slow pain perceived as a poorly localized, dull, aching sensation and is transmitted by C fibers
(0.2 - 1.3 m/sec).
Communication between first and second order neuron involves a neurotransmitter called substance P.

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The information that travels along the spinothalamic tract provides information about the location and
type of pain. Pain afferents also ascend along pathways that influence the behavioral and emotional
aspects of pain.
Visceral Pain
Generally, stimulation of nociceptors in the viscera produces a pain called referred pain. It is called
referred pain because it is referred to a body surface. Referred pain is due to second order neurons
receiving input from both somatic and visceral afferents. The impulses coming from the second-order
neurons is interpreted by the somatosensory cortex as coming from the somatic afferents.

Modulation of Pain Signals

Pain information ascending to higher centers can be either facilitated or attenuated.
Gate-Control Theory

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Describes how pain is modulated at the spinal level. Interneurons inhibit second order neurons that
transmit pain. C fibers transmitting pain signals from the periphery inhibit these interneurons. However,
A fibers transmitting touch, pressure or vibration stimulate these inhibitory neurons. Hence, non-painful
mechanical stimuli decrease pain transmission.
The brain can also influence the perception of pain through descending pathways that are part of the
endogenous analgesia (pain-blocking) systems.

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One example is shown in figure above. Neurons in the periaqueductal gray matter of the midbrain
communicate with the nucleus raphe magnus in the medulla and lateral reticular formation. Neurons
from these areas descend the spinal cord and synapse with inhibitory interneurons that release the
endogenous opiate neurotransmitter enkephalin. These in turn synapse with the axon terminals of
afferent neurons to decrease the release of substance P and the cell bodies and dendrites of second order
neurons to induce inhibitory post synaptic potentials.

Vision Web site devoted to vision

Anatomy of the Eye
Review the anatomy of the eye.
Nature and Behavior of Light Waves

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Visible light is a part of the electromagnetic spectrum with

wavelengths between 350 nm and 750 nm

Properties of light: Reflection and Refraction; Refraction through a Prism

Reflection
Light waves strike and bounce off surfaces that we see. Although we receive some light directly
(sun & light bulb) most light is reflected off the objects with non-perceived wavelengths absorbed.
Refraction
Light waves bend as they pass through transparent materials of different densities.

When parallel light waves strike a concave lens the waves striking the lens surface at a right angle goes
straight through but light waves striking the surface at other angles diverge. In contrast, light waves
striking a convex lens converge at a single point called a focal point. The distance from the long axis of
the lens to the focal point is the focal length.

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Both the cornea and the lens of the eye have convex surfaces and help to focus light rays onto the
retina. The cornea provides for most of the refraction but the curvature of the lens can be adjusted to
adjust for near and far vision.
Accommodation Accommodation applet

Light rays that enter the eye from distant objects are nearly parallel and require little bending or refraction to
focus on the retina. Light rays from close objects are diverging as they enter the eye and require more
refraction. The eye adjusts its refractive power by controlling the shape of the lens. The lens is suspended by
zonular fibers that exert a tension on the lens that causes it to flatten and have a lesser curvature for distant
objects. When viewing objects that are near the ciliary muscles contract releasing the tension on the lens.
Because of the elasticity of the fibers in the lens, the lens rounds up and the curvature and refractive power
increases.
Clinical Defects in Vision The Human Eye
Emmetropia

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This is normal vision. The eye can see distant objects clearly without accommodation and nearby objects
clearly with accommodation. In myopia and hyperopia this is not the case because there is a mismatch
between the refractive power of the lens and cornea and eyeball length.
Myopia

Near objects are seen clearly but not distant objects because the refractive power of the lens or cornea is too
strong for the length of the eyeball. Corrected by the use of concave lens that cause light waves to diverge.
Eye accommodates for close vision.
Hyperopia

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The lens or cornea is too weak for the length of the eyeball. Distant objects can be focused only with
accommodation and lens cannot accommodate enough for close objects. Corrected by the use of a convex
lens.
Astigmatism
Irregularities on the surface of the cornea or lens cause uneven refraction and results in distortion.
Presbyopia
Loss in elasticity of lens with age results in loss of accommodation (reading glasses are needed!).
Cataract
Age related loss in transparency of lens.
Glaucoma
Increase in the volume of aqueous humor raises pressure in anterior cavity of eyeball. Can lead to blindness
if unchecked.
Regulating Light Entry (extreme close up slow motion human eye with pupil dilating + expanding

The eyes can regulate the amount of light entering the eye by adjusting the size of the pupil. The pupil is
constricted in bright light and dilated in dim light. The size of the pupil is controlled by the iris. The iris has a
circular muscle layer that constricts the pupil and a radial muscle layer that dilates the pupil.
Parasympathetic neurons control the circular muscle layer while sympathetic neurons control the radial
muscle layer.

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Retina
Retina is the location of photoreceptors:
Rods give black and white vision under low light conditions.
Cones give color vision in bright light.

Retina has three distinct layers

Inner layer (closest to middle of eyeball) contains ganglion cells.
Middle layer contains bipolar cells.
Outer layer contains rods and cones.
Amacrine and horizontal cells are also present and modulate communication by lateral inhibition.

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To increase visual acuity the retina has an area where cells of the inner and middle layers are laterally
displaced. This area is called the fovea (depression). The fovea is the center of a spot that contains a high
concentration of cones called the macula lutea (yellow spot). The concentration of the cones is highest here
and decreases towards the periphery where there are more rods.
Phototransduction
Light energy is converted to electrical signals by rods and cones. These photoreceptors can be divided into
outer and inner segments. The outer segment contains invaginations of the cell membrane or membranous
discs. The membranes contain molecules that absorb light. The inner segment contains the cell nucleus,
organelles and the synaptic terminal.

There are four different kinds of photoreceptors each containing a different photopigment that absorbs
light. One type of photopigment is found in rods and three different photopigments are found in the three kinds
of cones.
Each photopigment contains a light absorbing portion retinal and a protein called opsin. The kind of opsin
determines which wavelengths of light are optimally absorbed by the retinal.

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The photopigment of the rods is rhodopsin. It is associated with a G protein called transducin and the
enzyme phosphodiesterase which degrades cyclic GMP (cGMP).
The Process of Phototransduction Photoisomerization of rhodopsin
A. Photoreceptor in the Dark Transduction Animation
In the dark levels of cGMP are high.
1. cGMP in cytosol opens sodium channels
in the membrane of the outer segment.
2. Na+ moves in and depolarizes the
membrane.
3. Depolarization spreads to the inner
segment.
4. Depolarization opens Ca++ channels.
5. Ca++ enters the cell and triggers release of
transmitter by exocytosis.
6. Transmitter causes graded potential in
the bipolar cell.
B. Photoreceptor in the Light

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1. Light is absorbed by rhodopsin.

Retinal changes shape and is released
by opsin.
2. This creates bleached opsin that
activates transducin.
3. Activated transducin activates
phosphodiesterase.
4. Phosphodiesterase breaks down
cGMP.
5. Decreased cGMP causes Na+
channels to close.
6. K+ continues to leak out
hyperpolarizing the membrane.
7. Hyperpolarization spreads to the
inner segment and closes Ca++
channels.
8. Less Ca++ enters the cell.
9. Less transmitter is released.
10. Graded potential in bipolar cells
decreases.
Rods are sensitive to light in a wide spectrum but is most sensitive to the blue-green range. Rods are
highly sensitive and can respond to a single photon.

Cones are not as sensitive as rods and respond best to narrower ranges of wavelengths:
S (blue) cones - most sensitive at 430 nm
M (green) cones - most sensitive at 530 nm

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L (red) cones - most sensitive at 560 nm (see Table 10.4)

Bleaching of Photoreceptors in Light Illusions Blind Spot
When exposed to bright light the rhodopsin becomes bleached and opsin is in its active form. No
more light can be absorbed. When a dark environment (movie theater) is entered bleached rods are not
sensitive to light. Retinal and opsin reassociate in dim light and rhodopsin becomes sensitive to light
again.
Neural Processing in the Retina
There are varying degrees of photoreceptors converging onto bipolar cells. The lesser the
convergence the greater the visual acuity. For example, at the macula lutea where there is maximum
visual acuity very few photoreceptors converge on a bipolar cell. In the periphery where there is less
visual acuity but more sensitivity many photoreceptors converge on each bipolar cell.
Both the photoreceptor and bipolar cells are incapable of generating action potentials and the graded
potentials caused by stimuli is modulated by the horizontal and amacrine cells.

The ganglion cells transmit signals to the brain by action potentials. Each ganglion cell has a
receptive field that can be complex. For example, there are on-center, off-surround ganglion cells which
fire most strongly when there is light in the center of the field but no light in the surround. There are also
off-center, on-surround ganglion cells that fire most strongly when the opposite is true. The axons of the
ganglion cells form the optic nerve.
Neural Pathways for Vision
The optic nerve exits each eye and combines in front of the brainstem to form the optic chiasm. The
light from the left visual field strikes the nasal retina of the left eye and the temporal retina of the right.

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The ganglion axons from the left nasal retina cross over to the right brain at the optic chiasm while those
from the right temporal retina stay on the same side. The same is true for information from the right visual
field.

The result is that, after information arrives at the optic chiasm, input from the left visual field goes to
the right side of the brain and vice versa. The fibers that continue from the optic chiasm to the lateral
geniculate body of the thalamus due so in the optic tract. At the lateral geniculate body synapses are
formed with neurons that go to the visual cortex as part of the optic radiation.
The visual field is mapped onto the cortex in a topographic organization.
Parallel Processing
For each point in the visual field information about different qualities of the stimuli such as color,
shape and movement are transmitted by parallel pathways to the primary visual cortex where they are
interpreted.
Depth Perception
Most areas of the right and left visual fields are detected by both eyes. This is necessary for depth
perception because the right and left eyes see objects from different angles. The cortex converts these
differences into three dimensional images.

Ear
Anatomy

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Review the anatomy on your own.

Nature of Sound Waves

Sound waves are mechanical waves caused by air molecules in motion. Sound has properties of
loudness and pitch. Loudness (amplitude) is due to differences in the densities of compressed and
rarified areas. The loudness is expressed in decibels (db) on a logarithmic scale. Pitch is determined
by the frequency of sound waves. Frequency is measured by the number of waves per second or Hertz
(Hz). The average range of hearing is 20-20,000 Hz with the greatest sensitivity in 1000-4000 Hz.
Sound Amplification in the Middle Ear Tutorial 45.1 Sound Transduction in the Human Ear
It takes greater pressure to produce waves in the fluid of the cochlear than in the air of the outer ear.
The ear amplifies the pressure wave in the air by two means:
1. The three ear ossicles are arranged to function as a series of levers. The movement of the maleus
causes a lesser movement of greater force of the incus which in turn causes a lesser movement of
greater force of the stapes. The oval plate of the stapes then presses on the fluid of the cochlear duct
with greater force in producing a pressure wave.
2. The diameter of the tympanic membrane is much larger than the membrane on the oval
window. The pressure on the larger surface translates to a larger pressure on the smaller surface.
Anatomy of the Cochlea

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Sound transduction occurs in the cochlear. Review the anatomy of the cochlea.
Anatomy of the Organ of Corti
The organ of Corti is on top of the basilar membrane and contains hair cells, supporting cells and
an overlying tectorial membrane. The hair cells have stereocilia the tips of which are embedded in
the tectorial membrane.
Sound Transduction by Hair Cells Sound Transduction

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Sound waves causes pressure waves in the endolymph of the cochlear duct. The basilar membrane
moves relative to the tectorial membrane in which the stereocilia of hair cells is embedded. This causes
the stereocilia to bend. The distortion of stereocilia causes potassium channels to either open or close.
Because the concentration of K+ in endolymph is higher than that in the cell, the opening of K+ channels
causes the cell to depolarize as K+ rushes in. The closing of these channels cause hyperpolarization.

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The stereocilia are arranged in decreasing sizes and are connected by elastic protein filaments. When
the hair cell is at rest potassium channels are partially open and partial depolarization causes Ca++
channels to open. This results in the release of neurotransmitter by the hair cell which causes neurons of
the cochlear nerve to fire action potentials.
When stereocilia bend in the direction of the tallest stereocilia the increased tension on stereocilia
open the potassium channels more increasing depolarization. This increases Ca++ entry and
neurotransmitter release and the frequency of action potentials in afferent neurons. When stereocilia
bend away from the tallest stereocilia potassium channels close and the results are opposite.
Coding Sound Intensity and Pitch
The greater the intensity (loudness) of sound the greater the bending of the stereocilia. The larger the
variations in transmitter release produces greater variations in action potentials in the afferent neurons.

The basilar membrane varies in structure over its length with the membrane being narrow and stiff
near the oval and round windows and wider and more flexible near the helicotrema. High frequency
sounds cause greater deflection of the basilar membrane where it is narrow and stiff and lower frequency
sounds produce greater deflection where the basilar membrane is loose and flexible. The frequency of
sound (pitch) is coded by where along the basilar membrane there is the greatest deflection.
Neural Pathways for Sound
The afferent neurons travel in the cochlear nerve (VIII) with frequency of action potential coding
intensity of sound. In the brainstem afferent neurons synapse with second-order neurons in the cochlear
nuclei that travel to the medial geniculate body of the thalamus. Third-order neurons travel to the
auditory cortex of the temporal lobe. The frequency of sound is mapped out in the auditory cortex in a
tonotopic manner.

Ear and Equilibrium

The ear in addition to transducing sound also detects the acceleration of the body and the position of the

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head in relation to the rest of the body. Please note, that acceleration is a change in the motion of a body. It
can be a change of speed (linear) or a change in direction (rotation).
Anatomy of Vestibular Apparatus

Review anatomy
The semicircular canals detect rotational acceleration. There are three:
1. Anterior
2. Posterior
3. Horizontal
Utricle detects linear acceleration forward and backward.
Saccule detects linear acceleration up and down.
Transduction of Rotational (Angular) Acceleration

Hair cells are found in the ampulla of each semicircular canal. The stereocilia of these receptor cells are
similar to those of the organ of Corti with the largest ones next to a cilium called a kinocilium. The

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stereocilia and kinocilium are embedded in a gelatinous mass called the cupula.

When the head is at rest the hair cells are partially depolarized and cause action potentials in the
afferent neurons at a low frequency. When the head rotates the bony labyrinth rotates but the endolymph
within the membranous labyrinth lags behind because of inertia. Depending on the angle of rotation, the
stereocilia within the cupula either bend toward or away from the kinocilium. If the stereocilia bend away
from the kinocilium the hair cell is hyperpolarized and action potentials in the afferent neuron decrease.
If the stereocilia bend toward the kinocilium the hair cell is depolarized and action potentials in the
afferent neuron increase.
Transduction of Linear Acceleration

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Within the utricle and saccule are areas containing hair cells with stereocilia embedded in a gelatinous
mass. The surface of the gelatinous mass has calcium carbonate crystals called otoliths. During linear
acceleration hair cells within the utricle and saccule are subjected to the same kinds of mechanical
distortion as described in the cochlea and semicircular canals. These cells not only detect linear
acceleration but provide information about the relative position of the head in space.
Neural Pathways for Equilibrium

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The afferent neurons from the semicircular canals and utricle and saccule travel to the brain in the
vestibular nerve. The nerve goes to the vestibular nuclei and the cerebellum. The information is
compared to that received from the eyes, proprioceptors and somesthetic receptors to enable balance and
equilibrium and to control eye movements.

Taste
Anatomy of Taste Buds

The taste buds contain 50-150 receptor cells and numerous support cells. Taste receptor cells have
microvilli that extend through a pore exposing the microvilli to the saliva on the surface with its
dissolved food molecules.
Signal Transduction in Taste
There are four primary tastes
A. Sour - Caused by the presence of H+ in food.
1. Hydrogen ions bind to potassium channels.
2. Potassium channels close.
3. Taste receptor depolarizes and calcium channels open up.
4. Calcium triggers release of transmitter by exocytosis.
B. Salty - Caused by the presence of sodium in food.
1. Increased concentration of sodium increases flow of sodium into cell.
2. Voltage-gated calcium channels open as cell depolarizes.
3. Calcium triggers transmitter release.

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C. Sweet - Presence of organic molecules with structure similar to sucrose.

1. Organic molecule binds with membrane receptor.
2. G protein called gustducin activated.
3. Stimulates production of cAMP.
4. cAMP activates protein kinase.
5. Kinase catalyzes phosphorylation of K+ channels and they close.
6. The cell depolarizes and voltage gated Ca++ channels open.
7. Ca++ triggers release of transmitter.
D. Bitter - Associated with a variety of nitrogen-containing compounds. Two mechanisms
involved:
1. "Bitter" molecules block K+ channels.
2. K+ leakage decreases.
3. Cell depolarizes and calcium channel opens.
4. Calcium enters the cell and transmitter is released.
Other bitter molecules:
1. Ligands bind to receptors on cell membrane.
2. Receptor inactivates gustducin.
3. Decreases activity of adenylate cyclase
4. Decrease levels of cAMP
5. Calcium channels close.
A fifth taste has been discovered that is activated by amino acids such as glutamate
(monosodium glutamate MSG) and are flavor enhancers. This taste has been called "umami"
(delicious).
Receptors cells have all four transduction mechanisms. However, receptor cells respond stronger
to one primary taste than the other.
Several receptor cells converge on a single afferent neuron and afferent neurons respond in a
complex fashion to different tastes.
Neural Pathway for Taste
Taste afferent neurons travel in VII, IX and X. Afferent neurons synapse in gustatory nuclei of the
medulla with second order neurons that travel to the contralateral thalamus. Third order neurons go
from thalamus to the gustatory cortex.

Olfaction
Odorants (specific molecules) dissolve in mucus and bind to chemoreceptors.
Anatomy of Olfactory System

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Olfactory epithelium consists of supporting cells, basal cells and olfactory receptor cells. Basal
cells replace olfactory receptor cells.
Olfactory Signal Transduction
1. Odorant molecules bind to membrane receptors and activates a G protein.
2. G protein activates adenylate cyclase.
3. cAMP is formed.
4. cAMP binds and opens Na+ and Ca++ channels causing depolarization.
5. Ca++ also causes chloride channels to open and Cl- to exit from the cell, increasing the
depolarization.
6. If the depolarization is great enough, action potentials are triggered.
Neural Pathway for Olfaction

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The olfactory neurons form the olfactory nerve (cranial nerve I). The axons form synapses with
second-order neurons in the olfactory bulb in structures called glomeruli (sing. glomerulus). The
second-order neurons (mitral cells) form the olfactory tract which goes to the olfactory tubercle.
From the olfactory tubercle olfactory information goes either to the olfactory cortex concerned with
the discrimination of smells, or to the limbic system for triggering olfactory-driven behaviors, such as
sexual behaviors.

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