Unit 4 Tutorials: Regulation, Integration,

and Control
INSIDE UNIT 4

The Nervous System and Nervous Tissue

Basic Structure and Function of the Nervous System

Nervous Tissue
Membrane Potential
Action Potential
Graded Potentials
Communication Between Neurons

The Anatomy of the Nervous System

Central Nervous System: Spinal Cord

Central Nervous System: Brainstem and Cerebellum
Central Nervous System: Cerebrum and Diencephalon
Meninges
Ventricular System of the Central Nervous System
Peripheral Nervous System: Spinal Nerve
Peripheral Nervous System: Reflexes
Peripheral Nervous System: Cranial Nerves

The Somatic Nervous System

Sensory Perception
General Senses
Special Senses: Taste and Smell
Special Senses: Hearing and Equilibrium
Special Senses: Vision
Central Processing: Spinal Cord and Brainstem
Central Processing: Sensory Cortex
Central Processing: Motor Responses

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 The Autonomic Nervous System

Divisions of the Autonomic Nervous System

Autonomic Reflexes and Homeostasis
Central Control

Basic Structure and Function of the Nervous

System
by Sophia

 WHAT'S COVERED

In this lesson, you will learn how the nervous system is categorized into anatomical and physiological
divisions. Specifically, this lesson will cover:
1. The Central and Peripheral Nervous Systems
2. Functional Divisions of the Nervous System
2a. Basic Functions
2b. Controlling the Body

1. The Central and Peripheral Nervous Systems

 THINK ABOUT IT

The picture you have in your mind of the nervous system probably includes the brain, the organ composed
primarily of nervous tissue contained within the cranium, and the spinal cord, the organ composed primarily
of nervous tissue within the vertebral column. That suggests it is made of two organs—and you may not
even think of the spinal cord as an organ—but the nervous system is a very complex structure. Within the
brain, many different and separate regions are responsible for many different and separate functions. It is
as if the nervous system is composed of many organs that all look similar and can only be differentiated
using tools such as the microscope or electrophysiology. In comparison, it is easy to see that the stomach is
different than the esophagus or the liver, so you can imagine the digestive system as a collection of specific
organs.
The nervous system can be divided into two major regions: the central and peripheral nervous systems. The
central nervous system (CNS) is the brain and spinal cord (the central axis), and the peripheral nervous system
(PNS) is the structure outside of the central axis, nerves, and related structures. The brain is contained within

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 the cranial cavity of the skull, and the spinal cord is contained within the vertebral cavity of the vertebral
column. It is a bit of an oversimplification to say that the CNS is what is inside these two cavities and the
peripheral nervous system is outside of them, but that is one way to start to think about it. In actuality, there are
some elements of the peripheral nervous system that are within the cranial or vertebral cavities and some
elements of the central nervous system that are outside of them. Depending on different aspects of the nervous
system, the dividing line between central and peripheral is not necessarily universal.

Central and Peripheral Nervous System - The structures of the PNS are referred to as ganglia and nerves, which can

be seen as distinct structures. The equivalent structures in the CNS are not obvious from this overall perspective and

are best examined in prepared tissue under the microscope.

Nervous tissue is present in both the CNS and PNS. Recall that nervous tissue contains two basic types of cells:
neurons and neuroglial cells, sometimes referred to as glial cells or glia (glia, glue). Remember that neurons

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 function to generate and propagate electrochemical signals throughout the body while glial cells provide
support for neurons.

In order to understand the organization patterns of the nervous system, it is important to review the structure of
a neuron. Recall that a neuron is composed of a cell body, or soma, with two types of processes that extend
outwards. Dendrites branch off of the cell body and monitor the electrochemical activity in their surrounding
area, bringing electrical signals into the soma. The axon extends away from the cell body and propagates an
electrical signal onto the next cell. Some neurons will have a wrapping called myelin around portions of the
axon.

At the end of the axon are axon terminals which form connections with and transfer electrochemical signals to
other cells.

The Neuron - The cell body of a neuron, also called the soma, contains the nucleus and mitochondria. The dendrites

transfer the electrochemical signal to the soma. The axon carries the electrochemical signal away to another excitable

cell. The axon terminals for the synapse (connection) to the next excitable cell.

Looking at nervous tissue, there are regions that predominantly contain cell bodies and regions that are largely
composed of just axons. These two regions within nervous system structures are often referred to as gray
matter (the regions with many cell bodies and dendrites) or white matter (the regions with many axons).

 KEY CONCEPT

The image below demonstrates the appearance of these regions in the brain and spinal cord. The colors
ascribed to these regions are what would be seen in “fresh,” or unstained, nervous tissue. Gray matter is
not necessarily gray. It can be pinkish because of blood content, or even slightly tan, depending on how
long the tissue has been preserved. But the white matter is white because axons are insulated by myelin
which is lipid-rich. Lipids can appear as white (“fatty”) material, much like the fat on a raw piece of chicken
or beef. Actually, the gray matter may have that color ascribed to it because next to the white matter, it is
just darker—hence, gray.

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 The distinction between gray matter and white matter is most often applied to central nervous tissue, which has
large regions that can be seen with the unaided eye. When looking at peripheral structures, a microscope is
often used and the tissue is stained with artificial colors. That is not to say that central nervous tissue cannot be
stained and viewed under a microscope but unstained tissue is most likely from the CNS—for example, a frontal
section of the brain or a cross-section of the spinal cord.

Gray Matter and White Matter - A brain removed during an autopsy, with a partial section removed, shows white
matter surrounded by gray matter. Gray matter makes up the outer cortex of the brain.

Regardless of the appearance of stained or unstained tissue, the cell bodies of neurons or axons can be located
in discrete anatomical structures. Those structures are named based on whether they are located in the central
or peripheral nervous system. A localized collection of neuron cell bodies in the CNS is referred to as a nucleus
(CNS).

In the PNS, a cluster of neuron cell bodies is referred to as a ganglion. Admittedly, the term ‘nucleus’ has
multiple meanings in this anatomy and physiology course alone. The table below outlines those definitions to
help you better understand its uses.

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 What Is a Nucleus? - (a) The nucleus of an atom contains its protons and neutrons. (b) The nucleus of a cell is the

organelle that contains DNA. (c) A nucleus in the CNS is a localized center of function with the cell bodies of several

neurons, shown here circled in red.

The terminology applied to bundles of axons also differs depending on location. A bundle of axons, or fibers,
found in the CNS is called a tract whereas the same thing in the PNS would be called a nerve. It is important to
note that the same bundle of axons can be referred to as both a tract and nerve if it spans both central and
peripheral regions.

EXAMPLE One example of this is the axons that project from the retina in your eye to the brain. Those
axons are called the optic nerve as they leave the eye, but when they are inside the cranium, they are
referred to as the optic tract. This naming scheme can be applied to roads as well. A single road from point
A to point B in different towns may have different names on either side of the town, county, or state line.

Table: Structures of the CNS and PNS

CNS PNS

Group of Neuron Cell Bodies (i.e., gray matter) Nucleus Ganglion

Bundle of Axons (i.e., white matter) Tract Nerve

 TERMS TO KNOW

Brain
A nervous system organ located in the cranial cavity and primarily composed of nervous tissue.

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 Spinal Cord
A nervous system organ located in the vertebral cavity and primarily composed of nervous tissue.

Central Nervous System (CNS)

The anatomical division of the nervous system that consists of the brain and spinal cord.

Peripheral Nervous System (PNS)

The anatomical division of the nervous system that consists of structures outside of the brain and spinal
cord.

Gray Matter
Regions of nervous tissue with many cell bodies and dendrites.

White Matter
Regions of nervous tissue with many axons.

Nucleus (CNS)
A group of neuron cell bodies in the central nervous system.

Ganglion
A group of neuron cell bodies in the peripheral nervous system.

Tract
A bundle of axons in the central nervous system.

Nerve
A bundle of axons in the peripheral nervous system.

2. Functional Divisions of the Nervous System

In a separate method, the nervous system can be divided on the basis of its functions. The CNS and the PNS
both contribute to the same functions, but those functions can be attributed to different regions of the brain
(such as the cerebral cortex or the hypothalamus) or to different structures in the periphery. Furthermore, the
same structure can sometimes be part of several functions.

There are two ways to consider how the nervous system is divided functionally. First, the basic functions of the
nervous system are sensation, integration, and response. In other words, the system must detect a change,
process that information, and respond accordingly. Secondly, control of the body can be conscious or
subconscious, a division that is largely defined by the structures that are involved in the response.

2a. Basic Functions

Sensation: The first major function of the nervous system is sensation—the detection of information about the
environment to gain input about what is happening outside the body (or, sometimes, within the body) and the
translation of that into electrochemical signals. The sensory functions of the nervous system register the
presence of a change from homeostasis or a particular event in the external or internal environment, known as
a stimulus.

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  REFLECT

The senses we think of most are the “big five”: taste, smell, touch, sight, and hearing. The stimuli for taste
and smell are both chemical substances (molecules, compounds, ions, etc.), touch is physical or mechanical
stimuli that interact with the skin, sight is light stimuli, and hearing is the perception of sound, which is a
physical stimulus similar to some aspects of touch. There are actually more senses than just those, but that
list represents the major senses. Those five are all senses that receive stimuli from the outside world and of
which there is conscious perception. Additional sensory stimuli might be from the internal environment
(inside the body), such as the stretch of an organ wall or the concentration of certain ions in the blood.
Integration: Stimuli that are received by sensory structures are communicated to the nervous system where that
information is processed. This is called integration. Stimuli are compared with, or integrated with, other stimuli,
memories of previous stimuli, or the state of a person at a particular time. This leads to the specific response
that will be generated.

EXAMPLE Seeing a baseball pitched to a batter will not automatically cause the batter to swing. The
trajectory of the ball and its speed will need to be considered. Maybe the count is three balls and one
strike, and the batter wants to let this pitch go by in the hope of getting a walk to first base. Or, maybe the
batter’s team is so far ahead that it would be fun to just swing away.
Response: The nervous system produces a response, or internal event, on the basis of the stimuli perceived by
sensory structures. An obvious response would be the movement of muscles, such as withdrawing a hand from
a hot stove, but there are broader uses of the term. The nervous system can cause the contraction of all three
types of muscle tissue.

EXAMPLE Skeletal muscle contracts to move the skeleton, cardiac muscle is influenced as heart rate
increases during exercise, and smooth muscle contracts as the digestive system moves food along the
digestive tract. Responses also include the neural control of glands in the body as well, such as the
production and secretion of sweat by the eccrine and merocrine sweat glands found in the skin to lower
body temperature.
Responses can be divided into those that are voluntary or conscious (contraction of skeletal muscle) and those
that are involuntary (contraction of smooth muscles, regulation of cardiac muscle, and activation of glands).

2b. Controlling the Body

The nervous system can be divided into two parts mostly on the basis of a functional difference in responses.

1. The somatic nervous system (SNS) is a functional division of the nervous system which is responsible for
conscious perception and voluntary motor responses. Voluntary motor response means the contraction of
skeletal muscle. At times, these motor responses are intentional such as reaching for a cup and filling it with
water when you realize you are thirsty. At other times, these motor responses are performed unintentionally
such as screaming or jumping back when a friend jumps out from behind a corner and yells “Boo!” In both
cases, it is skeletal muscle that is activated in response to the received stimuli. The stimuli that elicit these
types of responses are also referred to as somatic stimuli.

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 2. The autonomic nervous system (ANS) is a functional division of the nervous system which is responsible
for involuntary perception and control of the body, usually for the sake of homeostasis (regulation of the
internal environment). Sensory input for autonomic functions can be from sensory structures tuned to
external or internal environmental stimuli. The motor output extends to smooth and cardiac muscle as well
as glandular tissue. The role of the autonomic system is to regulate the organ systems of the body, which
usually means to control homeostasis. Sweat glands, for example, are controlled by the autonomic system.
When you are hot, sweating helps cool your body down. That is a homeostatic mechanism. But when you
are nervous, you might start sweating also. That is not homeostatic; it is the physiological response to an
emotional state.

There is another division of the nervous system that describes functional responses. The enteric nervous
system (ENS) is responsible for controlling the smooth muscle and glandular tissue in your digestive system. It
is a large part of the PNS and is not dependent on the CNS. It is sometimes valid, however, to consider the
enteric system to be a part of the autonomic system because the neural structures that make up the enteric
system are a component of the autonomic output that regulates digestion. There are some differences between
the two, but for our purposes here, there will be a good bit of overlap.

Somatic, Autonomic, and Enteric Structures of the Nervous System - Somatic structures include the spinal nerves,

both motor and sensory fibers, as well as the sensory ganglia (posterior root ganglia and cranial nerve ganglia).
Autonomic structures are found in the nerves also but include the sympathetic and parasympathetic ganglia. The

enteric nervous system includes the nervous tissue within the organs of the digestive tract.

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 IN CONTEXT
Everyday Connection - How Much of Your Brain Do You Use?

Have you ever heard the claim that humans only use 10 percent of their brains? Maybe you have seen
an advertisement on a website saying that there is a secret to unlocking the full potential of your mind
—as if there were 90 percent of your brain sitting idle, just waiting for you to use it. If you see an ad
like that, don’t click. It isn’t true.

An easy way to see how much of the brain a person uses is to take measurements of brain activity
while performing a task. An example of this kind of measurement is functional magnetic resonance
imaging (fMRI), which generates a map of the most active areas and can be generated and presented
in three dimensions. This procedure is different from the standard MRI technique because it is
measuring changes in the tissue in time with an experimental condition or event.

fMRI - This fMRI shows activation of the visual cortex in response to visual stimuli.

The underlying assumption is that active nervous tissue will have greater blood flow. By having the

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 subject perform a visual task, activity all over the brain can be measured. Consider this possible
experiment: the subject is told to look at a screen with a black dot in the middle (a fixation point). A
photograph of a face is projected on the screen away from the center. The subject has to look at the
photograph and decipher what it is. The subject has been instructed to push a button if the
photograph is of someone they recognize. The photograph might be of a celebrity, so the subject
would press the button, or it might be of a random person unknown to the subject, so the subject
would not press the button.

In this task, visual sensory areas would be active, integrating areas would be active, motor areas
responsible for moving the eyes would be active, and motor areas for pressing the button with a finger
would be active. Those areas are distributed all around the brain and the fMRI images would show
activity in more than just 10 percent of the brain (some evidence suggests that about 80 percent of the
brain is using energy—based on blood flow to the tissue—during well-defined tasks similar to the one
suggested above). This task does not even include all of the functions the brain performs. There is no
language response, the body is mostly lying still in the MRI machine, and it does not consider the
autonomic functions that would be ongoing in the background.

 TERMS TO KNOW

Sensation
A nervous system function in which sensory information about the environment is detected translated
into electrochemical signals.

Integration
A nervous system function in which various stimuli are compared and/or integrated to produce a
response.

Response
A nervous system function in which an event is caused to occur as a consequence of a specific stimulus
or stimuli.

Somatic Nervous System (SNS)

A functional division of the nervous system which is responsible for conscious perception and voluntary
motor responses.

Autonomic Nervous System (ANS)

A functional division of the nervous system which is responsible for involuntary perception and control
of the body.

Enteric Nervous System (ENS)

A division of the autonomic nervous system which is responsible for controlling the smooth muscle and
glandular tissue in your digestive system.

 SUMMARY

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 In this lesson, you learned about the ways in which the nervous system is divided. The nervous system
can be divided anatomically into the central and peripheral nervous systems. Physiologically, it also
forms functional divisions of the nervous system which are categorized as the basic functions of
sensation, integration, and response or those that deal with controlling the body voluntarily or
involuntarily.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Autonomic Nervous System (ANS)

A functional division of the nervous system which is responsible for involuntary perception and control of
the body.

Brain
A nervous system organ located in the cranial cavity and primarily composed of nervous tissue.

Central Nervous System (CNS)

The anatomical division of the central nervous system that consists of the brain and spinal cord.

Enteric Nervous System (ENS)

A division of the autonomic nervous system which is responsible for controlling the smooth muscle and
glandular tissue in your digestive system.

Ganglion
A group of neuron cell bodies in the peripheral nervous system.

Gray Matter
Regions of nervous tissue with many cell bodies and dendrites.

Integration
A nervous system function in which various stimuli are compared and/or integrated to produce a
response.

Nerve
A bundle of axons in the peripheral nervous system.

Nucleus (CNS)
A group of neuron cell bodies in the central nervous system.

Peripheral Nervous System (PNS)

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 The anatomical division of the central nervous system that consists of structures outside of the brain and
spinal cord.

Response
A nervous system function in which an event is caused to occur as a consequence of a specific stimulus
or stimuli.

Sensation
A nervous system function in which sensory information about the environment is detected translated into
electrochemical signals.

Somatic Nervous System (SNS)

A functional division of the nervous system which is responsible for conscious perception and voluntary
motor responses.

Spinal Cord
A nervous system organ located in the vertebral cavity and primarily composed of nervous tissue.

Tract
A bundle of axons in the central nervous system.

White Matter
Regions of nervous tissue with many axons.

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 Nervous Tissue
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the cells and structures of nervous tissue. Specifically, this lesson will
cover:
1. Neurons
1a. Parts of a Neuron
1b. Types of Neurons
2. Glial Cells
2a. Glial Cells of the CNS
2b. Glial Cells of the PNS
2c. Myelin

 BEFORE YOU START

As you know, nervous tissue is composed of two types of cells, neurons and neuroglia (commonly referred
to as glial cells). Neurons are the primary type of cell that most anyone associates with the nervous system.
They are responsible for the computation and communication that the nervous system provides. They are
electrically active and release chemical signals to target cells. Neuroglia, henceforward referred to as glial
cells, or glia, are known to play a supporting role for nervous tissue. Ongoing research pursues an
expanded role that glial cells might play in signaling, but neurons are still considered the basis of this
function. Neurons are important, but without glial support, they would not be able to perform their function.

1. Neurons
Neurons are the cells considered to be the basis of nervous tissue. They are responsible for the electrical
signals that communicate information about sensations and that produce movements in response to those
stimuli, along with inducing thought processes within the brain. An important part of the function of neurons is in
their structure, or shape. The three-dimensional shape of these cells makes the immense number of
connections within the nervous system possible.

1a. Parts of a Neuron

Recall that the main part of a neuron is the cell body, which is also known as the soma (soma, body). The cell
body contains the nucleus and most of the major organelles. But, what makes neurons special is that they have
many extensions of their cell membranes, which are generally referred to as processes. Neurons are usually

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 described as having one, and only one, axon—a fiber that emerges from the cell body and projects to target
cells. That single axon can branch repeatedly to communicate with many target cells. It is the axon that
propagates the nerve impulse, which is communicated to one or more cells. The other processes of the neuron
are dendrites, which receive information from other neurons at specialized areas of contact called synapses.
The dendrites are usually highly branched processes, providing locations for other neurons to communicate
with the cell body. Information flows through a neuron from the dendrites, across the cell body, and down the
axon. This gives the neuron a polarity—meaning that information flows in this one direction. The image below
shows the relationship between these parts to one another.

Parts of a Neuron - The major parts of the neuron are labeled on a multipolar neuron from the CNS.

Where the axon emerges from the cell body, there is a special region referred to as the axon hillock. This is a
tapering of the cell body toward the axon. Within the axon hillock, the cytoplasm changes to a solution of limited
components called axoplasm. Superficially, the proteins in the cell membrane change. The cell membrane
around the axon is referred to as the axolemma. The proximal portion of the axon, distal and adjacent to the
axon hillock, is known as the initial segment and plays a large role in the ability of the neuron to produce and
transfer electrical signals. You’ll learn more about this in a future lesson.

Many axons are wrapped by an insulating substance called myelin, which is actually made from glial cells. Recall
that myelin acts as insulation much like the plastic or rubber that is used to insulate electrical wires. A key
difference between myelin and the insulation on a wire is that there are gaps in the myelin along an axon where

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 the axolemma is exposed. Each gap is called a Node of Ranvier and is important to the way that electrical
signals travel down the axon. At the end of the axon is the axon terminal, where there are usually several
branches extending toward the target cell, each of which ends in an enlargement called a synaptic end bulb.
These bulbs are what make the connection with the target cell at the synapse.

1b. Types of Neurons

There are many neurons in the nervous system—a number in the trillions. And there are many different types of
neurons. They can be classified by many different criteria. The first way to classify them is by the number of
processes attached to the cell body. Using the standard model of neurons, one of these processes is the axon,
and the rest are dendrites. Because information flows through the neuron from dendrites or cell bodies toward
the axon, these names are based on the neuron's polarity.

Neuron Classification by Shape - Unipolar cells have one process that includes both the axon and dendrite. Bipolar
cells have two processes, the axon and a dendrite. Multipolar cells have more than two processes, the axon and two

or more dendrites.

Unipolar neurons have only one process emerging from the cell. True unipolar cells are only found in
invertebrate animals, so the unipolar cells in humans are more appropriately called “pseudo-unipolar” cells.
Invertebrate unipolar cells do not have dendrites. Human unipolar cells have an axon that emerges from the cell
body, but it splits so that the axon can extend along a very long distance. At one end of the axon are dendrites,

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 and at the other end, the axon forms synaptic connections with a target. Unipolar cells are exclusively sensory
neurons and have two unique characteristics.

First, their dendrites are receiving sensory information, sometimes directly from the stimulus itself.
Secondly, the cell bodies of unipolar neurons are always found in ganglia (groups of neuron cell bodies
located in the peripheral nervous system). Sensory reception is a peripheral function. The axon projects
from the dendrite endings, past the cell body in a ganglion, and into the central nervous system.

Bipolar neurons have two processes, which extend from each end of the cell body, opposite to each other. One
is the axon, and one is the dendrite. Bipolar cells are not very common. They are found mainly in the olfactory
epithelium (where smell stimuli are sensed) and as part of the retina.

Multipolar neurons are all of the neurons that are not unipolar or bipolar. They have one axon and two or more
dendrites (usually many more). The majority of neurons in the human body are multipolar. Some cutting-edge
research suggests that certain neurons in the CNS do not conform to the standard model of “one, and only one”
axon. Some sources describe a fourth type of neuron, called an anaxonic neuron. The name suggests that it has
no axon (an, without), but this is not accurate. Anaxonic neurons are very small, and if you look through a
microscope at the standard resolution used in histology (approximately 400X to 1000X total magnification), you
will not be able to distinguish any process specifically as an axon or a dendrite. Any of those processes can
function as an axon depending on the conditions at any given time. Nevertheless, even if they cannot be easily
seen and one specific process is definitively the axon, these neurons have multiple processes and are therefore
multipolar.

Neurons can also be classified on the basis of where they are found, who found them, what they do, or even
what chemicals they use to communicate with each other. Some neurons referred to in this Unit on the nervous
system are named on the basis of those sorts of classifications. For example, a multipolar neuron that has a very
important role to play in a part of the brain called the cerebellum is known as a Purkinje (per-KIN-gee) cell. It is
named after the anatomist who discovered it (Jan Evangelista Purkinje, 1787–1869).

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 Other Neuron Classifications - Three examples of neurons that are classified on the basis of other criteria. (a) The
pyramidal cell is a multipolar cell with a cell body that is shaped something like a pyramid. (b) The Purkinje cell in the

cerebellum was named after the scientist who originally described it. (c) Olfactory neurons are named for the

functional group with which they belong.

 TERMS TO KNOW

Synapse
A cell-to-cell connection by which information is transferred and received.

Axon Hillock
A tapering of the neuron cell body towards the axon.

Axoplasm
The cytoplasm within the axon of a neuron.

Axolemma
The cell membrane of the axon of a neuron.

Initial Segment
The first portion of the axon of a neuron.

Node of Ranvier
A gap in the myelin surrounding a neuron where the axon is exposed.

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 Synaptic End Bulb
The expansion at the end of the axon where a synapse is formed.

Unipolar Neuron
A neuron with a single process.

Bipolar Neuron
A neuron with two processes.

Multipolar Neuron
A neuron with more than two processes.

2. Glial Cells
 BEFORE YOU START

You first learned about neuroglia or glial cells back in Unit 1 (towards the end of that unit). While you learned
the basics about the types of glial cells, here, they will be broken out in more detail. Remember you can
always go back and review previous lessons if desired.
Glial cells, or neuroglia or simply glia, are the other type of cell found in nervous tissue. They are considered to
be supporting cells, and many functions are directed at helping neurons complete their function for
communication. The name glia comes from the Greek word that means “glue” and was coined by the German
pathologist Rudolph Virchow. Today, research into nervous tissue has shown that there are many deeper roles
that these cells play, and research may find much more about them in the future.

There are six types of glial cells. Four of them are found in the CNS and two are found in the PNS. The table
below outlines some common characteristics and functions.

Table: Glial Cell Types by Location and Basic Function

CNS glia PNS glia Basic function

Astrocyte Satellite cell Support

Schwann
Oligodendrocyte Myelination (the wrapping of neuronal axons with myelin)
cell

Immune surveillance and phagocytosis (the endocytosis of large particles,

Microglia -
aka “cell eating”)

Ependymal cell - Creating cerebrospinal fluid (CSF)

2a. Glial Cells of the CNS

Recall that there are four glial cells that are located in the central nervous system (CNS). Below is a review of
each of these four cell types—astrocyte, oligodendrocyte, microglia, and ependymal cell.

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 1. The astrocyte is named because it has many processes extending in all directions which makes it appear
star-shaped (astro, star). These processes extend to interact with neurons, blood vessels, or a connective
tissue covering the CNS. Astrocytes support neurons by maintaining the concentration of chemicals in the
extracellular space, removing excess signaling molecules, reacting to tissue damage, and contributing to
the blood-brain barrier (BBB). The blood-brain barrier is a layer of astrocyte membrane around the blood
vessels in the brain which serves to restrict what can cross from circulating blood into the CNS. Nutrient
molecules, such as glucose or amino acids, can pass through the BBB but other items cannot. This protects
the CNS from toxins and pathogens but also restricts items such as cells of the immune system from being
able to enter to protect the body. Pharmaceutical companies are challenged to design drugs that can cross
the BBB as well as have an effect on the nervous system.

Glial Cells of the CNS - The CNS has astrocytes, oligodendrocytes, microglia, and ependymal cells that support the
neurons of the CNS in several ways.

2. Also found in CNS tissue is the oligodendrocyte, sometimes called just “oligo,” which is the glial cell type
that insulates axons in the CNS. The name means “cell of a few branches” (oligo, few; dendro, branches;
cyte, cell). There are a few processes that extend from the cell body. Each one reaches out and surrounds

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 an axon to insulate it in myelin. One oligodendrocyte will provide the myelin for multiple axon segments,
either for the same axon or for separate axons. The function of myelin will be discussed later.

3. Microglia (micro, small) are, as the name implies, smaller than most of the other glial cells. While their
origin is not conclusively determined, their function is to ingest and digest diseased or damaged cells or the
pathogens that cause disease.

4. The ependymal cell is a glial cell that filters blood to make cerebrospinal fluid (CSF), the fluid that
circulates through the CNS. Because of the privileged blood supply inherent in the BBB, the extracellular
space in nervous tissue does not easily exchange components with the blood. Ependymal cells line four
cavities in the brain. These glial cells appear similar to epithelial cells, making a single layer of cells with
little intracellular space and tight connections between adjacent cells. They also have cilia on their apical
surface to help move the CSF.

2b. Glial Cells of the PNS

Recall that there are two glial cells that are located in the peripheral nervous system (PNS). Below is a review of
each of these two cell types—satellite and Schwann cells.

1. Satellite cells are found in sensory and autonomic ganglia, where they surround the cell bodies of
neurons. This accounts for the name, based on their appearance under the microscope. They provide
support, performing similar functions in the periphery as astrocytes do in the CNS—except, of course, for
establishing the BBB.

2. The second type of glial cell is the Schwann cell, which insulates axons with myelin in the periphery.
Schwann cells are different than oligodendrocytes, in that a Schwann cell wraps around a portion of only
one axon segment and no others. Oligodendrocytes have processes that reach out to multiple axon
segments, whereas the entire Schwann cell surrounds just one axon segment. The nucleus and cytoplasm
of the Schwann cell are on the edge of the myelin sheath. The relationship of these two types of glial cells
to ganglia and nerves in the PNS is seen in the image below.

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 Glial Cells of the PNS - The PNS has satellite cells and Schwann cells.

2c. Myelin
The insulation for axons in the nervous system is provided by glial cells, oligodendrocytes in the CNS, and
Schwann cells in the PNS. Whereas the manner in which either cell is associated with the axon segment, or
segments, that it insulates is different. The means of myelinating an axon segment is mostly the same in the two
situations.

Recall that every cell contains a cell membrane and those cell membranes are primarily composed of
phospholipids. Oligodendrocyte and Schwann cells insulate their respective CNS or PNS neurons by wrapping
extensions of their cell body around the axon. This wrapping is the cell membrane of each cell and is therefore
lipid (mostly phospholipid) rich. The myelin wrapping is known as a myelin sheath (close-fitting covering)
facilitates the transmission of electrical signals along the axon. Myelin, however, is more than just the membrane
of the glial cell. It also includes important proteins that are integral to that membrane. Some of the proteins help
to hold the layers of the glial cell membrane closely together.

 KEY CONCEPT

The appearance of the myelin sheath can be thought of as similar to the pastry wrapped around a hot dog
for “pigs in a blanket” or similar food. The glial cell is wrapped around the axon several times with little to no
cytoplasm between the glial cell layers. For oligodendrocytes, the rest of the cell is separate from the
myelin sheath as a cell process extends back toward the cell body. A few other processes provide the same

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 insulation for other axon segments in the area. For Schwann cells, the outermost layer of the cell membrane
contains cytoplasm and the nucleus of the cell as a bulge on one side of the myelin sheath.

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 The Process of Myelination - Myelinating glia wrap several layers of cell membrane around the cell membrane of an
axon segment. A single Schwann cell insulates a segment of a peripheral nerve, whereas in the CNS, an
oligodendrocyte may provide insulation for a few separate axon segments. EM × 1,460,000.

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 IN CONTEXT
Disorders of the Nervous Tissue

Several diseases can result from the demyelination of axons. The causes of these diseases are not the
same; some have genetic causes, some are caused by pathogens, and others are the result of
autoimmune disorders. Though the causes are varied, the results are largely similar. The myelin
insulation of axons is compromised, making electrical signaling slower.

Multiple sclerosis (MS) is one such disease. It is an example of an autoimmune disease. The antibodies
produced by lymphocytes (a type of white blood cell) mark myelin as something that should not be in
the body. This causes inflammation and the destruction of the myelin in the central nervous system. As
the insulation around the axons is destroyed by the disease, scarring becomes obvious. This is where
the name of the disease comes from; sclerosis means hardening of tissue, which is what a scar is.
Multiple scars are found in the white matter of the brain and spinal cord. The symptoms of MS include
both somatic and autonomic deficits. Control of the musculature is compromised, as is control of
organs such as the bladder.

Guillain-Barré (pronounced gee-YAN bah-RAY) syndrome is an example of a demyelinating disease of

the peripheral nervous system. It is also the result of an autoimmune reaction, but the inflammation is
in peripheral nerves. Sensory symptoms or motor deficits are common, and autonomic failures can
lead to changes in the heart rhythm or a drop in blood pressure, especially when standing, which
causes dizziness.

 TERMS TO KNOW

Blood Brain Barrier (BBB)

A physiological barrier between the blood and central nervous system that restricts the movement of
substances into or out of the CNS.

Cerebrospinal Fluid (CSF)

Fluid produced by ependymal cells that circulates within the central nervous system.

Myelin Sheath
A lipid-rich layer of insulation produced by oligodendrocytes and Schwann cells that surrounds an axon.

 SUMMARY

In this lesson, you learned about the cells of nervous tissue, neurons and glial cells. You learned to
identify the parts of a neuron as well as the types of neurons present in the body. You also learned to
identify the glial cells of the CNS and PNS. Lastly, you learned about the production and formation of
myelin.

© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 25
 Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Axolemma
The cell membrane of the axon of a neuron.

Axon Hillock
A tapering of the neuron cell body towards the axon.

Axoplasm
The cytoplasm within the axon of a neuron.

Bipolar Neuron
A neuron with two processes.

Blood Brain Barrier (BBB)

A physiological barrier between the blood and central nervous system that restricts the movement of
substances into or out of the CNS.

Cerebrospinal Fluid (CSF)

Fluid produced by ependymal cells that circulates within the central nervous system.

Initial Segment
The first portion of the axon of a neuron.

Multipolar Neuron
A neuron with more than two processes.

Myelin Sheath
A lipid-rich layer of insulation produced by oligodendrocytes and Schwann cells that surrounds an axon.

Node of Ranvier
A gap in the myelin surrounding a neuron where the axon is exposed.

Synapse
A cell-to-cell connection by which information is transferred and received.

Synaptic End Bulb

The expansion at the end of the axon where a synapse is formed.

Unipolar Neuron
A neuron with a single process.

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 Membrane Potential
by Sophia

 WHAT'S COVERED

In this lesson, you will learn how certain cells become electrically active and create membrane
potentials. Specifically, this lesson will cover:
1. Electrically Active Cell Membranes
2. The Membrane Potential

1. Electrically Active Cell Membranes

Most cells in the body make use of charged particles, ions, to build up a charge across the cell membrane.
Recall that this was shown to be a part of how muscle cells work. For skeletal muscles to contract, based on
excitation–contraction coupling, input from a neuron is required. Both of the cells make use of the cell
membrane to regulate ion movement between the extracellular fluid and cytosol.

Also, recall that the cell membrane is primarily responsible for regulating what can cross the membrane and
what stays on only one side. The cell membrane is a phospholipid bilayer, so only substances that can pass
directly through the hydrophobic core can diffuse through unaided. Charged particles, which are hydrophilic by
definition, cannot pass through the cell membrane without assistance. Transmembrane proteins, specifically
channel proteins, make this possible. Several passive transport channels, as well as active transport pumps, are
necessary to generate a transmembrane potential and an electrochemical signal. Of special interest is the
carrier protein referred to as the sodium-potassium pump that moves sodium ions (Na⁺) out of a cell and
potassium ions (K⁺) into a cell, thus regulating ion concentration on both sides of the cell membrane.

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 Cell Membrane and Transmembrane Proteins - The cell membrane is composed of a phospholipid bilayer and has
many transmembrane proteins, including different types of channel proteins that serve as ion channels.

 KEY CONCEPT

The sodium-potassium pump requires energy in the form of adenosine triphosphate (ATP). On either side of a
resting neuron, the concentration of Na⁺ is higher outside the cell than inside, and the concentration of K⁺ is
higher inside the cell than outside. That means that this pump is moving the ions against the concentration
gradients for sodium and potassium, which is why it requires energy. In fact, the pump basically maintains those
concentration gradients.
Ion channels are pores that allow specific charged particles to cross the membrane in response to an existing
concentration gradient. Each ion channel is a protein formed by a unique sequence of amino acids. The amino
acids that are present along the pore and the size of the pore will determine the type of ion that channel allows
through. Channels for cations (positive ions) will have negatively charged side chains in the pore. Channels for
anions (negative ions) will have positively charged side chains in the pore. Smaller pores cannot allow larger
ions through while larger pores do not allow smaller ions (associated with water) through. Some ion channels
are selective for charge but not necessarily for size, and thus are called a nonspecific ion channel.

Ion channels do not always freely allow ions to diffuse across the membrane. Some are opened by certain
events, meaning the channels are gated. So another way that channels can be categorized is on the basis of
how they are gated. Although these classes of ion channels are found primarily in the cells of nervous or
muscular tissue, they also can be found in the cells of epithelial and connective tissues.

A ligand-gated channel, also known as a chemically-gated channel, opens because a signaling molecule, a
ligand (chemical), binds to the extracellular region of the channel.

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 Ligand-Gated Channels - When the ligand, in this case the neurotransmitter acetylcholine, binds to a specific location
on the extracellular surface of the channel protein, the pore opens to allow select ions through. The ions, in this case,
are cations of sodium, calcium, and potassium.

A mechanically gated channel opens because of a physical distortion of the cell membrane. Many channels
associated with the sense of touch are mechanically gated. For example, as pressure is applied to the skin,
these channels open and allow ions to enter the cell.

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 Mechanically Gated Channels - When a mechanical change occurs in the surrounding tissue, such as pressure or
touch, the channel is physically opened. Thermoreceptors work on a similar principle. When the local tissue

temperature changes, the protein reacts by physically opening the channel.

A voltage-gated channel is a channel that responds to changes in the electrical properties of the membrane in
which it is embedded. Normally, the inner portion of the membrane is at a negative voltage. When that voltage
becomes less negative, the channel begins to allow ions to cross the membrane.

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 Voltage-Gated Channels - Voltage-gated channels open when the transmembrane voltage changes around them.
Amino acids in the structure of the protein are sensitive to charge and cause the pore to open to the selected ion.

A leakage channel is randomly gated, meaning that it opens and closes at random, hence the reference to
leaking. There is no actual event that opens the channel; instead, it has an intrinsic rate of switching between
the open and closed states.

Leakage Channels - In certain situations, ions need to move across the membrane randomly. The particular electrical

properties of certain cells are modified by the presence of this type of channel.

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  TERMS TO KNOW

Nonspecific Ion Channel

An ion channel that is specific to charge but not size.

Ligand-Gated Channel
A membrane channel that opens once a specific signaling molecule binds to it.

Mechanically-Gated Channel
A membrane channel that opens because of physical distortion of the cell membrane.

Voltage-Gated Channel
A membrane channel that responds to changes in the electrical properties of the membrane.

Leakage Channel
A membrane channel that is randomly gated, opening and closing randomly.

2. The Membrane Potential

The electrical state of the cell membrane can have several variations. These are all variations in the membrane
potential. Recall that a membrane potential is a difference in electrical charge across a cell membrane. This
charge is based on a differential distribution of charge across the cell membrane, measured in millivolts (mV).
The standard is to compare the inside of the cell relative to the outside, so the membrane potential is a value
representing the charge on the intracellular side of the membrane based on the outside being zero, relatively
speaking. As shown in the image below, there is an overall positive charge on the outside of the cell membrane
and an overall negative charge on the inside. Therefore, the membrane potential (inside relative to outside) is
negative. If the indicated charges were swapped (outside was negative and inside was positive), then the
membrane potential would be positive.

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 Measuring Charge across a Membrane with a Voltmeter - A recording electrode is inserted into the cell and a

reference electrode is outside the cell. By comparing the charge measured by these two electrodes, the
transmembrane voltage is determined. It is conventional to express that value for the cytosol relative to the outside.

The concentration of ions in extracellular and intracellular fluids is largely balanced, with a net neutral charge.
However, a slight difference in charge occurs right at the membrane surface, both internally and externally. It is
the difference in this very limited region that has all the power in neurons (and muscle cells) to generate
electrical signals.

 WATCH

Please watch the following video for more information on this topic.

 KEY CONCEPT

Before electrical signals can be generated and propagated, the cell must first find itself at rest. When the
cell is at rest and the ion channels are closed (except for leakage channels which randomly open), ions are
distributed across the membrane in a very predictable way.

The concentration of Na⁺ outside the cell is greater than the concentration inside.
The concentration of K⁺ inside the cell is greater than outside.
The cytosol contains a high concentration of anions (negative ions), in the form of phosphate ions and
negatively charged proteins.

With the ions distributed across the membrane at these concentrations, the difference in charge during
steady-state conditions is described as the resting membrane potential. The exact value measured for the

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 resting membrane potential varies between cells, -70 mV is the most commonly used value for neurons
(skeletal muscle cells are -85 mV).
 HINT

Here’s one way to remember which ion—sodium (Na⁺) or potassium (K⁺) is found in greater concentration
inside or outside the cell membrane at rest. When people come knocking on the door, most people are
more likely to allow their kin (relatives) in and keep strangers out.

Potassium comes knocking, should you let them in? Okay (K, potassium)
Potassium’s chemical symbol is the letter K (i.e., okay).
Kin, come in (K In, potassium in)
Sodium comes knocking, should you let them in? Nah! (Na, sodium)
Sodium’s chemical symbol is the letters Na (i.e., nah!)

 TERM TO KNOW

Resting Membrane Potential

The difference in charge across a cell membrane during steady state conditions.

 SUMMARY

In this lesson, you learned about the components of electrically active cell membranes. You then
learned how these electrically active cells generate a membrane potential.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Leakage Channel
A membrane channel that is randomly gated, opening and closing randomly.

Ligand-Gated Channel
A membrane channel that opens once a specific signaling molecule binds to it.

Mechanically-Gated Channel
A membrane channel that opens because of physical distortion of the cell membrane.

Nonspecific Ion Channel

An ion channel that is specific to charge but not size.

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 Resting Membrane Potential
The difference in charge across a cell membrane during steady state conditions.

Voltage-Gated Channel
A membrane channel that responds to changes in the electrical properties of the membrane.

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 Action Potential
by Sophia

 WHAT'S COVERED

In this lesson, you will learn how electrically active cells transmit electrical signals. Specifically, this
lesson will cover:
1. The Action Potential
2. Propagation of the Action Potential

1. The Action Potential

Resting membrane potential describes the steady state of the cell, which is a dynamic process balanced by ion
leakage and ion pumping. Without any outside influence, it will not change. However, an action potential is a
change in the membrane potential of an excitable cell in response to a stimulus. This stimulus disturbs the
membrane potential and causes the creation of an action potential. The first step of an action potential is the
opening of voltage-gated sodium channels.

 THINK ABOUT IT

Before you read on, see if you can predict the next event in the generation of an action potential.

Based on the known concentrations of sodium and potassium during the resting membrane
potential, what moves in what direction when a sodium channel is opened? +

Sodium moves in. Sodium has a greater concentration outside the cell at rest and will therefore move
down its concentration gradient when a sodium channel opens. A sodium channel is also not able to
transport potassium due to its specificity.

Recall that the concentration of Na⁺ is higher outside the cell. Therefore, when the sodium channel opens,
sodium ions rush into the cell. Because sodium is a positively charged ion, it will change the relative voltage
inside the cell relative to the outside. The resting potential is the state of the membrane at a voltage of -70 mV,
so the sodium cation entering the cell will cause it to become less negative. This is known as depolarization,
meaning the membrane potential moves toward zero.

The concentration gradient for Na⁺ is so strong that it will continue to enter the cell even after the membrane
potential has become zero so that the voltage immediately around the pore begins to become positive. The
membrane potential will reach +30 mV by the time sodium has entered the cell.

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 As the membrane potential reaches +30 mV, multiple voltage-gated channels respond. The voltage-gated
sodium channel that allowed for depolarization of the membrane closes. Additionally, a voltage-gated
potassium channel opens.

 THINK ABOUT IT

Before you read on, see if you can predict the next event in the generation of an action potential.

Based on the known concentrations of sodium and potassium after depolarization, what moves in
+
what direction when a potassium channel is opened?

Potassium moves out. Potassium has a greater concentration inside the cell at rest as well as at +30 mV
and will therefore move down its concentration gradient when a potassium channel opens. A
potassium channel is also not able to transport sodium due to its specificity.

Recall that the concentration of K⁺ is higher inside the cell. Therefore, when the potassium channel opens,
potassium ions rush out of the cell. As K⁺ starts to leave the cell, taking a positive charge with it, the membrane
potential begins to move back toward its resting voltage, -70 mV, a process called repolarization.

The membrane potential decreases as potassium moves out of the cell. When it reaches -70 mV, voltage-gated
potassium channels begin to close. As they slowly close, the membrane eventually reaches -90 mV, the
equilibrium for potassium ions. This period of excessive negative membrane potential is called
hyperpolarization. With both voltage-gated sodium and potassium channels closed, the sodium-potassium
pump, which has been active the entire time, can finally make progress on moving both ions back to their
respective sides. This returns the membrane potential to resting, -70mV, and resets the sodium and potassium
channels in preparation for the next electrical signal.

What has been described above are the steps of an action potential, which are presented as a graph of voltage
over time in the image below. It is the electrical signal that nervous tissue generates for communication.

 DID YOU KNOW

The change in the membrane voltage from -70 mV at rest to +30 mV at the end of depolarization is a 100
mV (0.1 V) change. To put that value in perspective, think about a battery. An AA (double A) battery that you
might find in a television remote has a voltage of 1.5 V, or a 9 V battery (the rectangular battery with two
posts on one end) is, obviously, 9 V. The change seen in the action potential is one or two orders of
magnitude less than the charge in these batteries. In fact, the membrane potential can be described as a
battery. A charge is stored across the membrane that can be released under the correct conditions. A
battery in your remote has stored a charge that is “released” when you push a button.

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 Graph of Action Potential - Plotting voltage measured across the cell membrane against time, the action potential

begins with depolarization, followed by repolarization, which goes past the resting potential into hyperpolarization,
and finally, the membrane returns to rest.

Recall that the action potential was initiated by a “stimulus,” but the description did not explain what that was.
Physiologically, this stimulus is an event that causes the membrane potential to increase slightly above resting,
moving toward zero. Anatomically, it is the opening of a ligand-gated or mechanically-gated sodium channel. A
ligand-gated channel opens when a neurotransmitter binds to its receptor. A mechanically-gated sodium
channel opens when the membrane of the sensory receptor is physically manipulated (like pressure applied to
the skin). In either case, sodium begins to enter the neuron and the membrane potential becomes less
negative.

If the stimulus is strong enough to allow the membrane potential to reach -55 mV, voltage-gated sodium
channels open. This membrane potential is known as the threshold. When reached, an action potential occurs.
If not reached, an action potential does not occur. There is no middle ground. The generation of an action
potential is an all-or-none event. Additionally, the events of an action potential are the same every time—the
peak is at +30 mV, hyperpolarization reaches +90 mV, and the membrane returns to -70 mV.

As we have seen, the depolarization and repolarization of an action potential are dependent on two types of
channels (the voltage-gated Na⁺ channel and the voltage-gated K⁺ channel). The voltage-gated Na⁺ channel
actually has two gates. One is the activation gate, which opens when the membrane potential crosses -55 mV.
The other gate is the inactivation gate, which closes after a specific period of time—on the order of a fraction of

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 a millisecond. When a cell is at rest, the activation gate is closed and the inactivation gate is open. However,
when the threshold is reached, the activation gate opens, allowing Na⁺ to rush into the cell. Timed with the
peak of depolarization, the inactivation gate closes. During repolarization, no more sodium can enter the cell.
When the membrane potential passes -55 mV again, the activation gate closes. After that, the inactivation gate
re-opens, making the channel ready to start the whole process over again. See these two Na⁺ channel gates in
the image below.

The voltage-gated K⁺ channel has only one gate, which is sensitive to a membrane voltage of -50 mV. However,
it does not open as quickly as the voltage-gated Na⁺ channel does. It might take a fraction of a millisecond for
the channel to open once that voltage has been reached. The timing of this coincides exactly with when the
Na⁺ flow peaks, so voltage-gated K⁺ channels open just as the voltage-gated Na⁺ channels are being
inactivated. As the membrane potential repolarizes and the voltage passes -50 mV again, the channel closes—
again, with a little delay. Potassium continues to leave the cell for a short while and the membrane potential
becomes more negative, resulting in the hyperpolarizing overshoot. Then the channel closes again and the
membrane can return to the resting potential because of the ongoing activity of the non-gated channels and
the Na⁺/K⁺ pump.

See the K⁺ channel gate and the two Na⁺ channel gates in action below.

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 Voltage-Gated Channel Gates - The gates of the voltage-gated sodium and potassium channels regulate the
movement of ions during an action potential. A) At resting membrane potential, the sodium channel has the
inactivation gate open and the activation gate closed while the potassium channel has its gate closed. B) When the

membrane reaches threshold, the sodium channel activation gate opens. C) When the membrane reaches +30 mV, the
sodium channel inactivation gate closes and the potassium channel gate opens. D) When the membrane undergoes
hyperpolarization, the potassium channel gate closes.

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 The events of an action potential take place within approximately 2 milliseconds. While an action potential is in
progress, another one cannot be initiated. That effect is referred to as the refractory period. There are two
phases of the refractory period:

Absolute refractory period

Relative refractory period

During the absolute refractory period, another action potential will not start. This is because of the inactivation
gate of the voltage-gated Na⁺ channel. Once that channel is back to its resting conformation (less than -55 mV),
the relative refractory period begins in which a new action potential could be started, but only by a stronger
stimulus than the one that initiated the current action potential. This is because of the flow of K⁺ out of the cell.
Because that ion is rushing out, any Na+ that tries to enter will not depolarize the cell, but will only keep the cell
from hyperpolarizing.

Stages of an Action Potential - Plotting voltage measured across the cell membrane against time, the events of the
action potential can be related to specific changes in the membrane voltage. (1) At rest, the membrane voltage is -70

mV. (2) The membrane begins to depolarize when an external stimulus is applied. (3) The membrane voltage begins a
rapid rise toward +30 mV. (4) The membrane voltage starts to return to a negative value. (5) Repolarization continues
past the resting membrane voltage, resulting in hyperpolarization. (6) The membrane voltage returns to the resting

value shortly after hyperpolarization.

 TERMS TO KNOW

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 Action Potential
The change in the membrane potential of an excitable cell in response to a stimulus.

Depolarize
The reduction of a membrane potential, making the inside less negative.

Repolarize
The return of a membrane potential to its negative state.

Hyperpolarize
The increase of a membrane potential beyond its resting state.

Activation Gate
A portion of the voltage-gated sodium channel that opens when the membrane voltage reaches
threshold.

Inactivation Gate
A portion of the voltage-gated sodium channel that closes when the membrane potential reaches +30
mV.

Refractory Period
The time period during an action potential when a new action potential cannot be generated.

Absolute Refractory Period

The time period during an action potential when a new action potential cannot be generated.

Relative Refractory Period

The time period during an action potential when a new action potential can only be generated by a
stronger stimulus.

2. Propagation of the Action Potential

The action potential is initiated at the initial segment, the beginning of the axon. There is a high density of
voltage-gated Na⁺ channels in this region so rapid depolarization can take place here. Going down the length
of the axon, the action potential is propagated because more voltage-gated Na⁺ channels are opened as the
depolarization spreads. This spreading occurs because Na⁺ enters through the channel and moves along the
inside of the cell membrane. As the Na⁺ moves, or flows, a short distance along the cell membrane, its positive
charge depolarizes a little more of the cell membrane. As that depolarization spreads, new voltage-gated Na⁺
channels open and more ions rush into the cell, spreading the depolarization a little farther.

Because voltage-gated Na⁺ channels are inactivated at the peak of the depolarization, they cannot be opened
again for a brief time—the absolute refractory period. Because of this, depolarization spreading back toward
previously opened channels has no effect. The action potential must propagate toward the axon terminals; as a
result, the polarity of the neuron is maintained, as mentioned above.

In an unmyelinated axon, propagation occurs along the entire length of the axon. This is referred to as
continuous propagation. In a myelinated axon, voltage-gated sodium and potassium channels only exist in the
nodes of Ranvier. This causes the depolarization to jump from node to node which is called saltatory
propagation (saltare, to leap).

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 Continuous propagation is slow because there are always voltage-gated Na⁺ channels opening, and more and
more Na⁺ is rushing into the cell. Saltatory propagation is faster because the action potential jumps from one
node to the next, and the new influx of Na⁺ renews the depolarized membrane. Along with the myelination of
the axon, the diameter of the axon can influence the speed of conduction. Much as water runs faster in a wide
river than in a narrow creek, Na⁺-based depolarization spreads faster down a wide axon than down a narrow
one.

 WATCH

Please watch the following video for more information on this topic.

IN CONTEXT
Homeostatic Imbalances - Potassium Concentration

Glial cells, especially astrocytes, are responsible for maintaining the chemical environment of the CNS
tissue. The concentrations of ions in the extracellular fluid are the basis for how the membrane
potential is established and changes in electrochemical signaling. If the balance of ions is upset,
drastic outcomes are possible.

Normally the concentration of K⁺ is higher inside the neuron than outside. After the repolarizing phase
of the action potential, K⁺ leakage channels and the Na⁺/K⁺ pump ensure that the ions return to their
original locations. Following a stroke or other ischemic event, extracellular K⁺ levels are elevated. The
astrocytes in the area are equipped to clear excess K⁺ to aid the pump. But when the level is far out of
balance, the effects can be irreversible.

Astrocytes can become reactive in cases such as these, which impairs their ability to maintain the local
chemical environment. The glial cells enlarge and their processes swell. They lose their K⁺ buffering
ability and the function of the pump is affected or even reversed. One of the early signs of cell disease
is this "leaking" of sodium ions into the body cells. This sodium/potassium imbalance negatively
affects the internal chemistry of cells, preventing them from functioning normally.

 TERMS TO KNOW

Continuous Propagation
Slow conduction of an action potential along an unmyelinated axon.

Saltatory Propagation
Fast conduction of an action potential along a myelinated axon.

 SUMMARY

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 In this lesson, you learned the steps of the action potential. You then learned that the propagation of
the action potential proceeds differently in myelinated versus unmyelinated axons.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Absolute Refractory Period

The time period during an action potential when a new action potential cannot be generated.

Action Potential
The change in the membrane potential of an excitable cell in response to a stimulus.

Activation Gate
A portion of the voltage-gated sodium channel that opens when the membrane voltage reaches
threshold.

Continuous Propagation
Slow conduction of an action potential along an unmyelinated axon.

Depolarize
The reduction of a membrane potential, making the inside less negative.

Hyperpolarize
The increase of a membrane potential beyond its resting state.

Inactivation Gate
A portion of the voltage-gated sodium channel that closes when the membrane potential reaches +30 mV.

Refractory Period
The time period during an action potential when a new action potential cannot be generated.

Relative Refractory Period

The time period during an action potential when a new action potential can only be generated by a
stronger stimulus.

Repolarize
The return of a membrane potential to its negative state.

Saltatory Propagation
Fast conduction of an action potential along a myelinated axon.

© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 44
 Graded Potentials
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about cell-to-cell connections and how they transfer and regenerate an
action potential. Specifically, this lesson will cover:
1. Graded Potentials
1a. Types of Graded Potentials
1b. Summation

 BEFORE YOU START

The electrical changes taking place within a neuron, as described in the previous section, are similar to a
light switch being turned on. A stimulus starts the depolarization, but the action potential runs on its own
once a threshold has been reached. The question is now, “What flips the light switch on?” Temporary
changes to the cell membrane voltage can result from neurons receiving information from the environment,
or from the action of one neuron on another. These special types of potentials influence a neuron and
determine whether an action potential will occur or not. Many of these transient signals originate at the
synapse.

1. Graded Potentials
Local changes in the membrane potential are called graded potentials and are usually associated with the
dendrites of a neuron. The amount of change in the membrane potential is determined by the size of the
stimulus that causes it.

EXAMPLE Testing the temperature of the shower, slightly warm water would only initiate a small change
in a heat-sensitive receptor, whereas hot water would cause a large amount of change in the membrane
potential.
Graded potentials can be of two sorts:

Depolarizing
Hyperpolarizing

For a neuronal membrane at the resting potential, a graded potential represents a change in that voltage either
above -70 mV or below -70 mV.

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 Depolarizing graded potentials are often the result of Na⁺ or Ca²⁺ entering the cell. Both of these ions have
higher concentrations outside the cell than inside; because they have a positive charge, they will move into the
cell causing it to become less negative relative to the outside.

Hyperpolarizing graded potentials can be caused by K⁺ leaving the cell or Cl⁻ entering the cell. If a positive
charge moves out of a cell, the cell becomes more negative; if a negative charge enters the cell, the same thing
happens.

Graded Potentials - Graded potentials are temporary changes in the membrane voltage, the characteristics of which
depend on the size of the stimulus. Some types of stimuli cause depolarization of the membrane, whereas others

cause hyperpolarization. It depends on the specific ion channels that are activated in the cell membrane.

1a. Types of Graded Potentials

A postsynaptic potential (PSP) is the graded potential in the dendrites of a neuron that is receiving synapses
from other cells. Postsynaptic potentials can be depolarizing or hyperpolarizing. Depolarization in a
postsynaptic potential is called an excitatory postsynaptic potential (EPSP) because it causes the membrane
potential to move toward threshold. Hyperpolarization in a postsynaptic potential is an inhibitory postsynaptic
potential (IPSP) because it causes the membrane potential to move away from threshold.

1b. Summation

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 All types of graded potentials will result in small changes of either depolarization or hyperpolarization in the
voltage of a membrane. These changes can lead to the neuron reaching threshold if the changes add together,
or summate. The combined effects of different types of graded potentials are illustrated in the image below. If
the total change in voltage in the membrane is a positive 15 mV, meaning that the membrane depolarizes from
-70 mV to -55 mV, then the graded potentials will result in the membrane reaching threshold.

Graded potentials summate at a specific location at the beginning of the axon to initiate the action potential,
namely the initial segment. This location has a high density of voltage-gated Na⁺ channels that initiate the
depolarizing phase of the action potential.

Summation can be spatial or temporal, meaning it can be the result of multiple graded potentials at different
locations on the neuron, or all at the same place but separated in time. Spatial summation is the combination of
various inputs to a neuron with each other. Temporal summation is the combination of multiple action potentials
from a single cell resulting in a significant change in the membrane potential. Spatial and temporal summation
can act together, as well.

Postsynaptic Potential Summation - The result of summation of postsynaptic potentials is the overall change in the

membrane potential. At point A, several different excitatory postsynaptic potentials add up to a large depolarization.

At point B, a mix of excitatory and inhibitory postsynaptic potentials result in a different end result for the membrane
potential.

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  TERMS TO KNOW

Graded Potential
Local changes in the membrane potential.

Postsynaptic Potential (PSP)

The graded potential in the dendrites of a neuron that is receiving synapses from other cells.

Excitatory Postsynaptic Potential (EPSP)

A depolarizing graded potential in the postsynaptic membrane.

Inhibitory Postsynaptic Potential (IPSP)

A hyperpolarizing graded potential in the postsynaptic membrane.

Summate
Add together.

Spatial Summation
The combination of various inputs to a neuron with each other.

Temporal Summation
The combination of multiple action potentials from a single cell resulting in a significant change in the
membrane potential.

 SUMMARY

In this lesson, you learned how graded potentials are created, how the types of graded potentials
affect membrane potential, and how summation functions to allow graded potentials to create an
action potential.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Excitatory Postsynaptic Potential (EPSP)

A depolarizing graded potential in the postsynaptic membrane.

Graded Potential
Local changes in the membrane potential.

Inhibitory Postsynaptic Potential (IPSP)

A hyperpolarizing graded potential in the postsynaptic membrane.

Postsynaptic Potential (PSP)

The graded potential in the dendrites of a neuron that is receiving synapses from other cells.

© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 48
 Spatial Summation
The combination of various inputs to a neuron with each other.

Summate
Add together.

Temporal Summation
The combination of multiple action potentials from a single cell resulting in a significant change in the
membrane potential.

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 Communication Between Neurons
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about how neurons are connected and communicate. Specifically, this
lesson will cover:
1. Synapses
1a. Neurotransmitter Release
1b. Neurotransmitter Systems

1. Synapses
Recall that a synapse is a cell-to-cell connection by which information is transferred and received. There are two
types of connections between electrically active cells:

Chemical Synapses
Electrical Synapses

In a chemical synapse, a chemical signal—namely, a neurotransmitter—is released from one cell the presynaptic
cell) and it affects the other cell (the postsynaptic cell).

In an electrical synapse, there is a direct connection between the two cells so that ions can pass directly from
one cell to the next. If one cell is depolarized in an electrical synapse, the joined cell also depolarizes because
the ions pass between the cells.

Chemical synapses involve the transmission of chemical information from one cell to the next. This section will
concentrate on the chemical type of synapse.

An example of a chemical synapse is the neuromuscular junction (NMJ) described in muscle tissue. In the
nervous system, there are many more synapses that are fundamentally the same as the NMJ. All synapses have
common characteristics, which can be summarized in this list:

Presynaptic Element
Neurotransmitter (packaged in vesicles)
Synaptic Cleft

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 Receptor Proteins
Postsynaptic Element
Neurotransmitter Elimination or Re-Uptake

For the NMJ, these characteristics are as follows. Other synapses contain the same characteristics with only
specific differences.

The presynaptic element is the motor neuron's axon terminals.

The neurotransmitter is acetylcholine.
The synaptic cleft is the space between the cells where the neurotransmitter diffuses.
The receptor protein is the nicotinic acetylcholine receptor.
The postsynaptic element is the sarcolemma of the muscle cell.
The neurotransmitter is eliminated by acetylcholinesterase.

1a. Neurotransmitter Release

When an action potential reaches the axon terminals, voltage-gated Ca²⁺ channels in the membrane of the
synaptic end bulb open. The concentration of Ca²⁺ increases inside the end bulb, and the Ca²⁺ ion associates
with proteins in the outer surface of neurotransmitter vesicles. The Ca²⁺ facilitates the merging of the vesicle
with the presynaptic membrane so that the neurotransmitter is released through exocytosis into the small gap
between the cells, known as the synaptic cleft.

Once in the synaptic cleft, the neurotransmitter diffuses the short distance to the postsynaptic membrane and
can interact with neurotransmitter receptors. Receptors are specific for the neurotransmitter, and the two fit
together like a key and lock. One neurotransmitter binds to its receptor and will not bind to receptors for other
neurotransmitters, making the binding a specific chemical event.

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 The Synapse - The synapse is a connection between a neuron and its target cell (which is not necessarily a neuron).
The presynaptic element is the synaptic end bulb of the axon where Ca²⁺ enters the bulb to cause vesicle fusion and

neurotransmitter release. The neurotransmitter diffuses across the synaptic cleft to bind to its receptor. The

neurotransmitter is cleared from the synapse either by enzymatic degradation, neuronal reuptake, or glial reuptake.

1b. Neurotransmitter Systems

There are several systems of neurotransmitters that are found at various synapses in the nervous system.
These groups refer to the chemicals that are the neurotransmitters, and within the groups are specific systems.

The first group, which is a neurotransmitter system of its own, is the cholinergic system. It is a system based on
acetylcholine. This includes the NMJ as an example of a cholinergic synapse, but cholinergic synapses are

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 found in other parts of the nervous system. They are in the autonomic nervous system, as well as distributed
throughout the brain.

The cholinergic system has two types of receptors, the nicotinic receptor is found in the NMJ as well as other
synapses. There is also an acetylcholine receptor known as the muscarinic receptor. Both of these receptors
are named for drugs that interact with the receptor in addition to acetylcholine. Nicotine will bind to the nicotinic
receptor and activate it similar to acetylcholine. Muscarine, a product of certain mushrooms, will bind to the
muscarinic receptor. However, nicotine will not bind to the muscarinic receptor and muscarine will not bind to
the nicotinic receptor.

Multiple other groups of neurotransmitters also play a role in the regulation and/or transmission of action
potentials and information throughout the body.

 LEARN MORE

To see more in-depth and additional information about these systems, please visit the supplemental 
Neurotransmitter Systems.pdf.

IN CONTEXT
Disorders of the Nervous System

The underlying cause of some neurodegenerative diseases, such as Alzheimer’s and Parkinson’s,
appears to be related to proteins—specifically, to proteins behaving badly. One of the strongest
theories of what causes Alzheimer’s disease is based on the accumulation of beta-amyloid plaques,
dense conglomerations of a protein that is not functioning correctly. Parkinson’s disease is linked to an
increase in a protein known as alpha-synuclein that is toxic to the cells of the substantia nigra nucleus
in the midbrain.

For proteins to function correctly, they are dependent on their three-dimensional shape. The linear
sequence of amino acids folds into a three-dimensional shape that is based on the interactions
between and among those amino acids. When the folding is disturbed and proteins take on a different
shape, they stop functioning correctly. But the disease is not necessarily the result of functional loss of
these proteins; rather, these altered proteins start to accumulate and may become toxic.

EXAMPLE In Alzheimer’s, the hallmark of the disease is the accumulation of these amyloid
plaques in the cerebral cortex. The term coined to describe this sort of disease is “proteopathy”
and it includes other diseases.

Creutzfeld-Jacob disease, the human variant of the prion disease known as mad cow disease in
the bovine, also involves the accumulation of amyloid plaques, similar to Alzheimer’s.

Diseases of other organ systems can fall into this group as well, such as cystic fibrosis or type 2
diabetes.

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 Recognizing the relationship between these diseases has suggested new therapeutic possibilities.
Interfering with the accumulation of the proteins, and possibly as early as their original production
within the cell, may unlock new ways to alleviate these devastating diseases.

 MAKE THE CONNECTION

If you're taking the Anatomy & Physiology I Lab course simultaneously with this lecture, it's a good time to
try the Lab Gross Function of the Nervous System: Let your brain learn about itself in Unit 5 of the Lab
course. Review the lab-to-lecture crosswalk if you need more information. Good luck!

 TERMS TO KNOW

Chemical Synapse
A cell-to-cell connection in which a neurotransmitter is released from one cell into a synaptic cleft and
interacts with another cell.

Electrical Synapse
A cell-to-cell connection in which there is a direct connection between the two cells so that ions can
pass directly from one cell to the next.

Cholinergic System
A neurotransmitter system in which acetylcholine is the neurotransmitter used at the chemical synapse.

Nicotinic Receptor
An acetylcholine receptor that is also activated by nicotine.

Muscarinic Receptor
An acetylcholine receptor that is also activated by muscarine.

 SUMMARY

In this lesson, you learned about the types of synapses that exist between electrically active cells. Then
you learned more details about how neurotransmitter release occurs at the axon terminal and the
types of neurotransmitter systems that exist throughout the body.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Chemical Synapse
A cell-to-cell connection in which a neurotransmitter is released from one cell into a synaptic cleft and
interacts with another cell.

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 Cholinergic System
A neurotransmitter system in which acetylcholine is the neurotransmitter used at the chemical synapse.

Electrical Synapse
A cell-to-cell connection in which there is a direct connection between the two cells so that ions can pass
directly from one cell to the next.

Muscarinic Receptor
An acetylcholine receptor that is also activated by muscarine.

Nicotinic Receptor
An acetylcholine receptor that is also activated by nicotine.

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 Central Nervous System: Spinal Cord
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of the spinal cord. Specifically, this lesson
will cover:
1. The Spinal Cord
1a. Gross Anatomy of the Spinal Cord
1b. Cross-Sectional Anatomy of the Spinal Cord
1c. Roots, Spinal Nerves, and Rami

 BEFORE YOU START

The brain and the spinal cord are the central nervous system, and they represent the main organs of the
nervous system. The spinal cord is a single structure, whereas the adult brain is described in terms of four
major regions: the cerebrum, the diencephalon, the brainstem, and the cerebellum. A person’s conscious
experiences are based on neural activity in the brain. The regulation of homeostasis is governed by a
specialized region in the brain. The coordination of reflexes depends on the integration of sensory and

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 motor pathways in both the brain and the spinal cord.

The Central Nervous system - The central nervous system is composed of the brain and spinal cord. The brain is

subdivided into the cerebrum, diencephalon, brainstem, and cerebellum.

1. The Spinal Cord

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 1a. Gross Anatomy of the Spinal Cord
Recall that the spinal cord is an organ of the nervous system primarily made of nervous tissue and located in the
vertebral canal formed by the vertebrae. At the superior end, the spinal cord connects to the brain through the
foramen magnum in the occipital bone. In the adult body, the inferior end of the spinal cord extends to
approximately vertebrae L1 or L2 because the spinal cord stops growing before the vertebrae do. The inferior
tip of the spinal cord is cone-shaped and is called the conus medullaris. Beyond this, a strand of fibrous
connective tissue called the filum terminale connects the conus medullaris to the sacrum to hold it in place.

Along the length of the spinal cord, 31 pairs of spinal nerves (collections of axons in the peripheral nervous
system) exit the vertebral space between each vertebra and extend out to various points throughout the body.
Each vertebra has its own set, which are named after the vertebral region where they originate—cervical,
thoracic, lumbar, and sacral spinal nerves. At the inferior end of the spinal cord, beyond the conus medullaris,
the lumbar and sacral spinal nerves extend into the vertebral cavity towards their exit. This collection of inferior
spinal nerves resembles a horse's tail and is known as the cauda equina (cauda, tail; equine, horse).

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© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 59
 Gross Anatomy of the Spinal Cord - The spinal cord is an organ of the CNS located in the vertebral canal created by

the vertebrae. The superior end is connected to the brain. The inferior end extends to the level of L1/L2 and forms the
conus medullaris. A connective tissue extension called the filum terminale connects the inferior spinal cord to the

coccyx. Between each vertebra, a pair of spinal nerves exit, connecting the CNS to the peripheral body.

1b. Cross-Sectional Anatomy of the Spinal Cord

The anterior midline is marked by the deep anterior median fissure, and the posterior midline is marked by the
shallow posterior median sulcus. These two grooves allow you to orient any view of the spinal cord to
determine the anterior and posterior sides. In the center of the spinal cord is a longitudinal hollow tube called
the central canal which circulates cerebrospinal fluid (CSF). CSF is produced in the brain and functions much
like synovial fluid to circulate nutrients and remove wastes from a region that does not have direct access to
blood.

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 Cross-section of Spinal Cord - The cross-section of a thoracic spinal cord segment shows the posterior, anterior, and

lateral horns of gray matter, as well as the posterior, anterior, and lateral columns of white matter. LM × 40.

Recall that nervous tissue can be grouped into gray and white matter. Gray matter is primarily composed of cell
bodies, dendrites, and non-myelinated axons while white matter is primarily composed of myelinated axons.

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 The spinal cord is organized with the white matter being superficial and the gray matter being mostly deep. As
you can see in the image above, the gray matter is organized in what some call a letter H, a butterfly, or an ink-
blot test. The gray matter can be further subdivided into regions referred to as horns. The posterior horn is
responsible for sensory processing. The anterior horn sends out motor signals to the skeletal muscles. The
lateral horn, which is only found in the thoracic, upper lumbar, and sacral regions, contains cell bodies of motor
neurons of the autonomic nervous system.

 DID YOU KNOW

Some of the largest neurons of the spinal cord are the multipolar motor neurons in the anterior horn. The
fibers that cause contraction of skeletal muscles are the axons of these neurons. The motor neuron that
causes contraction of the big toe, for example, is located in the sacral spinal cord. The axon that has to
reach all the way to the belly of that muscle may be a meter in length. The neuronal cell body that maintains
that long fiber must be quite large, possibly several hundred micrometers in diameter, making it one of the
largest cells in the body.
Just as the gray matter is separated into horns, the white matter of the spinal cord is separated into columns.
These columns contain tracts (groups of axons in the central nervous system) that transport action potentials up
(ascending) or down (descending) the spinal cord. Ascending tracts of the spinal cord of nervous system fibers
in these columns carry sensory information up to the brain, whereas descending tracts of the spinal cord carry
motor commands from the brain. Looking at the spinal cord longitudinally, the columns extend along its length
as continuous bands of white matter. Between the two posterior horns of gray matter are the posterior columns.
Between the two anterior horns, and bounded by the axons of motor neurons emerging from that gray matter
area, are the anterior columns. The white matter on either side of the spinal cord, between the posterior horn
and the axons of the anterior horn neurons, are the lateral columns. The posterior columns are composed of
axons of ascending tracts. The anterior and lateral columns are composed of many different groups of axons of
both ascending and descending tracts—the latter carrying motor commands down from the brain to the spinal
cord to control output to the periphery.

1c. Roots, Spinal Nerves, and Rami

Attached to the spinal cord are the axons which connect the central nervous system to all peripheral structures
such as the internal organs, muscles, skin, and more. Axons enter the posterior side through the posterior
(dorsal) root. Axons emerge from the anterior side through the anterior (ventral) root. The directionality of
information flow through the nerve roots is one-way traffic in which incoming sensory information travels
through the posterior root to enter the spinal cord and outgoing motor information travels through the anterior
root to travel to the effector. Along the posterior root is a ganglion (collection of neuron cell bodies in the
peripheral nervous system) called the posterior root ganglion. This ganglion contains the cell bodies of the
sensory neurons that make up the posterior root.

Where the posterior and anterior root meet, they combine together for a short time before separating again.
This short segment is called the spinal nerve and contains both sensory and motor information. There are 31
pairs of spinal nerves named for the vertebral region where they are located: cervical, thoracic, lumbar, and
sacral. Beyond the spinal nerve, the axons that make it up separate into three different pathways, each referred
to as a ramus (ramus, branch). Each ramus (plural, rami) serves a different region of the body by transporting
sensory signals from that region to the spinal cord and motor signals from the spinal cord to that region. The

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 anterior rami serve the anterior and lateral parts of the body as well as the limbs. The posterior rami serve the
posterior side of the body. The rami communicantes (communicating branch) transports signals to and from the
internal organs and structures within the trunk.

Roots, Spinal Nerves, and Rami - The axons connected to the spinal cord form a network of pathways for incoming
and outgoing signals. The roots are connected directly to the spinal cord and only carry one type of signal—posterior

carries sensory and anterior carries motor. The roots combine to form a spinal nerve which then branches into three

rami which each serve unique regions of the body.

 TRY IT

Use the information about the pathway that sensory and motor information take to and from the spinal cord
to identify the pathway (rami, nerve, and roots) that the following signals take in chronological order.

Define the sensory and motor pathway for the sensation of a handshake to which your body
responds by squeezing your hand. +

Sensory: Anterior rami, spinal nerve, posterior root, spinal cord

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 Motor: Spinal cord, anterior root, spinal nerve, anterior rami

Define the sensory and motor pathway for the sensation of something crawling on your back to
which your body responds by swatting at it with your hand. +

Sensory: Posterior rami, spinal nerve, posterior root, spinal cord

Motor: Spinal cord, anterior root, spinal nerve, anterior rami

Define the sensory and motor pathway for the sensation of the stomach expanding to which your
body responds by relaxing the muscles around your stomach. +

Sensory: Rami communicantes, spinal nerve, posterior root, spinal cord

Motor: Spinal cord, anterior root, spinal nerve, rami communicantes

Define the sensory and motor pathway for the sensation of a the hot sun on the back of your neck
+
to which your body responds by releasing sweat on that region of the skin.

Sensory: Posterior rami, spinal nerve, posterior root, spinal cord

Motor: Spinal cord, anterior root, spinal nerve, posterior rami

 TERMS TO KNOW

Conus Medullaris
The cone-shaped inferior end of the spinal cord.

Filum Terminale
A strand of connective tissue that connects the inferior spinal cord to the sacrum.

Spinal Nerve
One of thirty-one sets of nerves transporting sensory and motor information into and out of the spinal
cord through the anterior and posterior roots.

Cauda Equina
A bundle of spinal nerves that resembles a horse’s tail and extends beyond the conus medullaris in the
vertebral cavity.

Anterior Median Fissure

A deep midline groove on the ventral side of the spinal cord.

Posterior Median Sulcus

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 A shallow midline groove on the dorsal side of the spinal cord.

Central Canal of the Spinal Cord

A hollow tube in the center of the spinal cord which allows for the circulation of cerebrospinal fluid.

Posterior Horn
Posterior gray matter region of the spinal cord.

Anterior Horn
Anterior gray matter region of the spinal cord.

Lateral Horn
Gray matter region of the spinal cord only found in the thoracic, upper lumber, and sacral regions.

Ascending Tracts of the Spinal Cord

Axons of the spinal columns carrying sensory signals.

Descending Tracts of the Spinal Cord

Axons of the spinal column carrying motor signals.

Posterior Column
White matter region of the spinal cord.

Anterior Column
White matter region of the spinal cord.

Lateral Column
White matter region of the spinal cord.

Posterior Root
Sensory axons entering the spinal cord at the posterior horn.

Anterior Root
Motor axons exiting the spinal cord at the anterior or lateral horn.

Posterior Root Ganglion

A collection of sensory neuron cell bodies along the posterior root.

Anterior Rami
The branch of the spinal nerve which serves the anterior and lateral portions of the trunk as well as the
limbs.

Posterior Rami
The branch of the spinal nerve which serves the posterior portions of the trunk.

Rami Communicantes
The branch of the spinal nerve which serves the internal organs of the trunk.

 SUMMARY

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 In this lesson, you learned about the anatomy and function of the spinal cord and brainstem. You
learned the structures of both the gross anatomy and cross-sectional anatomy of the spinal cord as
well as the roots, spinal nerves, and rami that transport action potentials into and out of the spinal cord.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Anterior Column
White matter region of the spinal cord.

Anterior Horn
Anterior gray matter region of the spinal cord.

Anterior Median Fissure

A deep midline groove on the ventral side of the spinal cord.

Anterior Rami
The branch of the spinal nerve which serves the anterior and lateral portions of the trunk as well as the
limbs.

Anterior Root
Motor axons exiting the spinal cord at the anterior or lateral horn.

Ascending Tracts of the Spinal Cord

Axons of the spinal columns carrying sensory signals.

Cauda Equina
A bundle of spinal nerves that resembles a horse’s tail and extends beyond the conus medullaris in the
vertebral cavity.

Central Canal of the Spinal Cord

A hollow tube in the center of the spinal cord which allows for the circulation of cerebrospinal fluid.

Conus Medullaris
The cone-shaped inferior end of the spinal cord.

Descending Tracts of the Spinal Cord

Axons of the spinal column carrying motor signals.

Filum Terminale
A strand of connective tissue that connects the inferior spinal cord to the sacrum.

Lateral Column

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 White matter region of the spinal cord.

Lateral Horn
Gray matter region of the spinal cord only found in the thoracic, upper lumber, and sacral regions.

Posterior Column
White matter region of the spinal cord.

Posterior Horn
Posterior gray matter region of the spinal cord.

Posterior Median Sulcus

A shallow midline groove on the dorsal side of the spinal cord.

Posterior Rami
The branch of the spinal nerve which serves the posterior portions of the trunk.

Posterior Root
Sensory axons entering the spinal cord at the posterior horn.

Posterior Root Ganglion

A collection of sensory neuron cell bodies along the posterior root.

Rami Communicantes
The branch of the spinal nerve which serves the internal organs of the trunk.

Spinal Nerve
One of thirty-one sets of nerves transporting sensory and motor information into and out of the spinal cord
through the anterior and posterior roots.

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 Central Nervous System: Brainstem and
Cerebellum
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of the brainstem and cerebellum in the
brain. Specifically, this lesson will cover:
1. Brainstem
1a. Medulla Oblongata
1b. Pons
1c. Midbrain
2. The Cerebellum

1. Brainstem
The superior end of the spinal cord connects to the inferior portion of the brain called the brainstem. This
region is subdivided into three regions, the medulla oblongata, pons, and midbrain. Collectively, the brainstem
provides a pathway for the transport of sensory and motor signals to higher or lower structures and maintains
homeostasis, including the cardiovascular and respiratory systems and rates.

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 This diagram shows a cross section of the a vertebrae, including the nerve pathways that carry information to and
from the central nervous system.

1a. Medulla Oblongata

The medulla oblongata, commonly referred to as the medulla, is the inferior portion of the brainstem. This
region is composed of significant amounts of white matter which is continuous with the white matter of the
spinal cord. The medulla serves as a multi-lane information highway with all sensory information traveling to the
brain and all motor information leaving the brain traveling through the medulla. This also means that damage to
the medulla can severely impair sensory or motor function. A diffuse region of gray matter throughout the
brainstem, known as the reticular formation, is related to sleep and wakefulness, such as general brain activity
and attention. Additional centers in the medulla regulate homeostatic mechanisms including heart rate and
strength as well as breathing rate and depth.

1b. Pons
The pons is the middle portion of the brainstem which has a bulbous anterior portion. The word pons comes
from the Latin word for bridge. It is visible on the anterior surface of the brainstem as the thick bundle of white
matter attached to the cerebellum. The pons is the main connection between the cerebellum and the

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 brainstem. In terms of sensory and motor signals, the pons acts like a point where multiple highways intersect.
Like the medulla, the pons allows sensory and motor information to travel through to or from the brain, but it
also redirects that information to the cerebellum which sits posteriorly. Additionally, the pons contains centers
that regulate those in the medulla.

1c. Midbrain
The midbrain is the superior portion of the brainstem, just below the diencephalon (you will learn about the
diencephalon in the next lesson). It is separated into the tectum and tegmentum, from the Latin words for roof
and floor, respectively. A canal for cerebrospinal fluid referred to as the cerebral aqueduct passes through the
center of the midbrain, such that these regions are the roof and floor of that canal.

The tectum contains four bumps collectively known as the corpora quadrigemina, or four paired bodies (quad,
four; geminus, paired; corpora, body). Each bump is referred to as a colliculus (plural, colliculi) which means
“little hill” in Latin. The colliculi come in inferior and superior pairs, both process sensory information and both
control involuntary reflex movements. The inferior colliculus is the inferior pair of these bumps and is part of the
auditory brainstem pathway. The primary function of these structures is to process auditory sensation (sound).

EXAMPLE If you hear a noise such as a loud bang or someone calling your name, the inferior colliculus
receives that signal and controls the movement of the head and neck in order to turn towards the noise
reflexively. This response helps you to identify noises quickly.
The superior colliculus is the superior pair of bumps. Similar to its inferior counterpart, the superior colliculus
controls involuntary reflexive movements of the head, neck, and eyes in order to respond to visual stimuli.

EXAMPLE If you see something move in your peripheral vision, you may have the reflex to turn towards
it while turning on a blinding light that closes the eyes and turns the head away.
Additionally, it also combines sensory information about visual space, auditory space, and somatosensory
space.

EXAMPLE If you are walking along the sidewalk on campus and you hear chirping, the superior
colliculus coordinates that information with your awareness of the visual location of the tree right above
you. That is the correlation between auditory and visual maps. If you suddenly feel something wet fall on
your head, your superior colliculus integrates that with the auditory and visual maps and you know that the
chirping bird just relieved itself on you. You want to look up to see the culprit but do not.
The tegmentum is continuous with the gray matter of the rest of the brainstem. Throughout the midbrain, pons,
and medulla, the tegmentum contains the nuclei that receive and send information through the cranial nerves,
as well as regions that regulate important functions such as those of the cardiovascular and respiratory systems.

 TERMS TO KNOW

Brainstem
The inferior portion of the brain which contains the midbrain, pons, and medulla oblongata.

Medulla Oblongata
The inferior portion of the brainstem.

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 Reticular Formation
A region of gray matter throughout the brainstem that regulates sleep and wakefulness.

Pons
The middle portion of the brainstem.

Midbrain
The superior portion of the brainstem.

Tectum
The roof of the midbrain.

Tegmentum
The floor of the midbrain.

Corpora Quadrigemina
A group of two pairs of enlargements (colliculi).

Inferior Colliculi
A pair of enlargements in the midbrain along the auditory pathway.

Superior Colliculi
A pair of enlargements in the midbrain that integrates visual, auditory, and somatosensory information.

2. The Cerebellum
The cerebellum (little brain) is located posterior to the pons of the brainstem and accounts for ~10% of the mass
of the brain. The surface of the cerebellum, like the larger cerebrum, is covered in nervous tissue folds called
gyri (singular, gyrus) and shallow grooves called sulci (singular, sulcus). As previously mentioned, the cerebellum
receives input from the pons which redirects ingoing sensory or outgoing motor signals to or from the
cerebrum. The cerebellum is largely responsible for comparing information from the cerebrum with sensory
feedback from the periphery through the spinal cord. The various regions of the cerebellum are connected to
one another and the brainstem through the white matter tracts collectively known as the arbor vitae (arbor, tree;
vitae, life).

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 The Cerebellum - The cerebellum is situated on the posterior surface of the brainstem. Descending input from the

cerebellum enters through the large white matter structure of the pons. Ascending input from the periphery and spinal

cord enters through the medulla oblongata and pons. Output goes to the midbrain, which sends a descending signal
to the spinal cord.

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 The cerebellum receives descending motor signals from the cerebrum and ascending sensory signals from the
spinal cord. The cerebellum’s job is to compare motor commands to current sensory information and, when the
motor command is inadequate, send out a corrective command to compensate.

EXAMPLE If walking is not coordinated, perhaps because the ground is uneven or a strong wind is
blowing, then the cerebellum sends out a corrective command to compensate for the difference between
the original command and the sensory feedback.

 TERMS TO KNOW

Cerebellum
The posterior portion of the brain which functions to coordinate motor and sensory signals; little brain.

Arbor Vitae
The branching white matter of the cerebellum.

 SUMMARY

In this lesson, you learned the location and function of the regions of the brainstem—the medulla
oblongata, pons, and midbrain. Then you learned about how the cerebellum is connected to the
brainstem and functions to coordinate signals of the higher functional regions of the brain.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Arbor Vitae
The branching white matter of the cerebellum.

Brainstem
The inferior portion of the brain which contains the midbrain, pons, and medulla oblongata.

Cerebellum
The posterior portion of the brain which functions to coordinate motor and sensory signals; little brain.

Corpora Quadrigemina
A group of two pairs of enlargements (colliculi).

Inferior Colliculi
A pair of enlargements in the midbrain along the auditory pathway.

Medulla Oblongata

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 The inferior portion of the brainstem.

Midbrain
The superior portion of the brainstem.

Pons
The middle portion of the brainstem.

Reticular Formation
A region of gray matter throughout the brainstem that regulates sleep and wakefulness.

Superior Colliculi
A pair of enlargements in the midbrain that integrates visual, auditory, and somatosensory information.

Tectum
The roof of the midbrain.

Tegmentum
The floor of the midbrain.

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 Central Nervous System: Cerebrum and
Diencephalon
by Sophia

 WHAT'S COVERED

In this lesson, you will learn the anatomy and physiology of the diencephalon and cerebrum.
Specifically, this lesson will cover:
1. The Diencephalon
1a. Thalamus
1b. Hypothalamus
1c. Epithalamus
2. The Cerebrum
2a. Cerebral Cortex
2b. Subcortical structures

1. The Diencephalon
The diencephalon is a region of the brain deep to the cerebrum and superior to the midbrain. Translated as
“through brain,” the diencephalon connects the cerebrum to the rest of the nervous system except for the
olfaction (smell) pathway. This region is subdivided into three parts:

Thalamus
Hypothalamus
Epithalamus

1a. Thalamus

The thalamus is a collection of nuclei that relay information between the cerebral cortex and the periphery,
spinal cord, or brainstem. All sensory information, except for the sense of smell, projects to the thalamus. The
thalamus then processes the information, influencing what stimuli are important, and projects the processed
information onto the correct region of the cerebrum. The cerebrum also sends motor information down to the
thalamus. This involves interactions with the cerebellum and other nuclei in the brainstem. This process
functions much like old telephone switchboard operators that would receive a call, ask who the caller wants to

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 be connected to, and then connect them by removing and inserting cables
into their appropriate plugs. The thalamus functions as a signal switchboard
between lower and higher centers in the brain.

1b. Hypothalamus
The hypothalamus is inferior (hypo, below) and slightly anterior to the
thalamus. This region of the diencephalon is a collection of nuclei that are
largely involved in regulating homeostasis. The hypothalamus is the
executive region in charge of the subconscious nervous system control
(autonomic nervous system) and the endocrine system (hormones) which
will be discussed in detail in the A&P II course. Other parts of the
hypothalamus are involved in memory and emotion as part of the limbic
system.

1c. Epithalamus
The epithalamus is located superior (epi, above) and slightly posterior to the thalamus. This region of the
diencephalon plays a role in the limbic system and the production of cerebrospinal fluid. It also contains the
pineal gland, which produces and releases melatonin to regulate circadian rhythms (sleep-wake cycles).

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 The Diencephalon - The diencephalon is composed primarily of the thalamus and hypothalamus, which together

define the walls of the third ventricle. The thalami are two elongated, ovoid structures on either side of the midline

that make contact in the middle. The hypothalamus is inferior and anterior to the thalamus, culminating in a sharp
angle to which the pituitary gland is attached.

 TERMS TO KNOW

Diencephalon
Region of the brain deep to the cerebrum and superior to the midbrain.

Thalamus
A collection of nuclei that relay information between the cerebral cortex and the periphery, spinal cord,
or brainstem.

Hypothalamus
Executive region in charge of the subconscious nervous system control (ANS) and the endocrine
system.

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 Epithalamus
Plays a role in the limbic system and the production of cerebrospinal fluid.

Pineal Gland
Produces and releases melatonin to regulate circadian rhythms (sleep-wake cycles).

2. The Cerebrum
The iconic superficial gray mantle of the human brain, which appears to make up most of the mass of the brain,
is the cerebrum. The wrinkled portion is the cerebral cortex, and the rest of the structure is beneath that outer
covering. Many of the higher neurological functions, such as memory, emotion, and consciousness, are the
result of cerebral function. The complexity of the cerebrum is different across vertebrate species. The cerebrum
of the most primitive vertebrates is not much more than the connection for the sense of smell. In mammals, the
cerebrum comprises the outer gray matter that is the cortex (from the Latin word meaning “bark of a tree”) and
several deep nuclei.

2a. Cerebral Cortex

The cerebrum is covered by a continuous layer of gray matter—the cerebral cortex. This thin, extensive region
of wrinkled gray matter is responsible for the higher functions of the nervous system. Like the cerebellum, it
contains gyri (folds) and sulci (grooves) on its surface. The pattern of these folds of tissue indicates specific
regions of the cerebral cortex.

 DID YOU KNOW

The head is limited by the size of the birth canal, and the brain must fit inside the cranial cavity of the skull.
Extensive folding in the cerebral cortex enables more gray matter to fit into this limited space. If the gray
matter of the cortex were peeled off of the cerebrum and laid out flat, its surface area would be roughly
equal to one square meter.
The surface of the cerebrum provides many features which can be used to map various functional regions.
Along the midline is deep groove or fissure called the longitudinal fissure which splits the cerebrum into two
halves, the right and left cerebral hemispheres. These two structures are connected by a large white matter
tract known as the corpus callosum.

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 The Cerebrum - The cerebrum is a large component of the CNS in humans, and the most obvious aspect of it is the

folded surface called the cerebral cortex.

If you recall the cranial bones that form the majority of the cranial cavity (frontal, parietal, occipital, and
temporal), you will be able to identify the four major regions, or lobes, of the cerebral cortex (see image below).

The frontal lobes are located underneath the frontal bone.

The parietal lobes are located underneath the parietal bones.
The occipital lobes are located underneath the occipital bone.
The temporal lobes are located underneath the temporal bones.
The insula (not shown) is the floor of the lateral sulcus, underneath the frontal, parietal, and temporal lobes.

As you can see in the image below, these lobes are furthermore defined by gyri and sulci on the cerebral
surface. The central sulcus runs perpendicular to the longitudinal fissure and separates the frontal and parietal
lobes. The lateral sulcus runs along the lateral side of the cerebrum and separates the temporal lobe from the
frontal and parietal lobes. The medial portion of the occipital lobe is separated from the parietal lobe by the
parieto-occipital sulcus. The fact that the lateral region where the parietal, occipital, and temporal lobe meet
has no obvious anatomical border between them is consistent with the functions of these regions being
interrelated.

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 Lobes of the Cerebral Cortex - The cerebral cortex is divided into four lobes. Extensive folding increases the surface

area available for cerebral functions.

 KEY CONCEPT

Different regions of the cerebral cortex can be associated with particular functions, a concept known as
localization of function. In the early 1900s, a German neuroscientist named Korbinian Brodmann performed
an extensive study of the microscopic anatomy—the cytoarchitecture—of the cerebral cortex and divided
the cortex into 52 separate regions on the basis of the histology of the cortex. His work resulted in a system
of classification known as Brodmann’s areas, which is still used today to describe the anatomical
distinctions within the cortex. The results from Brodmann’s work on the anatomy align very well with the
functional differences within the cortex.
The frontal lobe is associated with motor function as well as executive functions such as contemplation and
decision-making. Brodmann area 4 is at the posterior portion of the temporal lobe and anterior to the central
sulcus. This region is anatomically referred to as the precentral gyrus and physiologically as the primary motor

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 cortex. Neurons from this region instruct cells in the spinal cord to move skeletal muscles. Anterior to this
region are a few areas that are associated with planned movements. Anterior to the primary motor cortex is the
premotor area which is responsible for planning the many components of a movement to be made.

Brodmann’s areas 44 and 45 are known as Broca’s area and are responsible for the production of language
and controlling movements responsible for speech. For most people, this area is only located in the left cerebral
hemisphere. Anterior to Broca’s area and the premotor cortex is the prefrontal lobe, which serves cognitive
functions that can be the basis of personality, short-term memory, and consciousness. The prefrontal lobotomy
is an outdated mode of treatment for personality disorders (psychiatric conditions) that profoundly affected the
personality of the patient.

The parietal lobe contains Brodmann’s areas 1, 2, and 3 just posterior to the central sulcus. This gyrus is known
anatomically as the postcentral gyrus and physiologically as the primary somatosensory cortex. This region
processes tactile sensory signals for touch (pressure, pain, itch, vibration) as well as proprioception (body
position) and kinesthesia (body movement).

The temporal lobe contains Brodmann’s areas 41 and 42, known as the primary auditory cortex which
processes auditory sensory signals from the ear. Brodmann’s area 22, 39, and 40 (not shown in the image
below) make up what is known as the Wernicke’s area, another language region. This area is responsible for
processing written and spoken language and allowing us to comprehend meaning. Because regions of the
temporal lobe are part of the limbic system, memory is an important function associated with that lobe. Memory
is essentially a sensory function; memories are recalled sensations such as the smell of Mom’s baking or the
sound of a barking dog. Even memories of movement are really the memory of sensory feedback from those
movements, such as stretching muscles or the movement of the skin around a joint. Structures in the temporal
lobe are responsible for establishing long-term memory, but the ultimate location of those memories is usually
in the region in which the sensory perception was processed.

The insula is the floor of the lateral sulcus. If the temporal, frontal, and parietal lobes are retracted, the insula is
located underneath the area where they meet. This region of the cerebral cortex contains the primary gustatory
cortex which is responsible for processing gustatory (taste) sensory information.

The occipital lobe contains Brodmann areas 17, known as the primary visual cortex which is responsible for
processing visual sensory information from the eye. However, visual information is complex, so once it is
processed, it is further integrated in the temporal and parietal lobes as well.

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 Brodmann's Areas of the Cerebral Cortex - Brodmann mapping of functionally distinct regions of the cortex was based
on its cytoarchitecture at a microscopic level.

2b. Subcortical structures

Beneath the cerebral cortex are sets of nuclei known as basal nuclei that augment cortical processes. These
groups of neuron cell bodies in the central nervous system are responsible for comparing cortical processing
with the general state of activity in the nervous system to influence the likelihood of movement taking place.

EXAMPLE While a student is sitting in a classroom listening to a lecture, the basal nuclei will keep the
urge to jump up and scream from actually happening. (The basal nuclei are also referred to as the basal
ganglia, although that is potentially confusing because the term ganglia is typically used for peripheral
structures.)

IN CONTEXT
Everyday Connection - The Myth of Left Brain/Right Brain

There is a persistent myth that people are “right-brained” or “left-brained,” which is an

oversimplification of an important concept about the cerebral hemispheres. There is some
lateralization of function, in which the left side of the brain is devoted to language function and the
right side is devoted to spatial and nonverbal reasoning. Whereas these functions are predominantly
associated with those sides of the brain, there is no monopoly by either side on these functions. Many

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 pervasive functions, such as language, are distributed globally around the cerebrum.

Some of the support for this misconception has come from studies of split brains. A drastic way to deal
with a rare and devastating neurological condition (intractable epilepsy) is to separate the two
hemispheres of the brain. After sectioning the corpus callosum, a split-brained patient will have
trouble producing verbal responses on the basis of sensory information processed on the right side of
the cerebrum, leading to the idea that the left side is responsible for language function.

However, there are well-documented cases of language functions lost from damage to the right side
of the brain. The deficits seen in damage to the left side of the brain are classified as aphasia, a loss of
speech function; damage on the right side can affect the use of language. Right-side damage can
result in a loss of ability to understand figurative aspects of speech, such as jokes, irony, or metaphors.
Nonverbal aspects of speech can be affected by damage to the right side, such as facial expression or
body language, and right-side damage can lead to a “flat affect” in speech, or a loss of emotional
expression in speech—sounding like a robot when talking.

IN CONTEXT
Disorders of the Basal Nuclei

Parkinson’s disease is a disorder of the basal nuclei, specifically of the substantia nigra, that
demonstrates the effects of the direct and indirect pathways. Parkinson’s disease is the result of
neurons in the substantia nigra pars compacta dying. These neurons release dopamine into the
striatum. Without that modulatory influence, the basal nuclei are stuck in the indirect pathway, without
the direct pathway being activated. The direct pathway is responsible for increasing cortical
movement commands. The increased activity of the indirect pathway results in the hypokinetic
disorder of Parkinson’s disease.

Parkinson’s disease is neurodegenerative, meaning that neurons die that cannot be replaced, so there
is no cure for the disorder. Treatments for Parkinson’s disease are aimed at increasing dopamine
levels in the striatum. Currently, the most common way of doing that is by providing the amino acid L-
DOPA, which is a precursor to the neurotransmitter dopamine and can cross the blood-brain barrier.
With levels of the precursor elevated, the remaining cells of the substantia nigra pars compacta can
make more neurotransmitter and have a greater effect. Unfortunately, the patient will become less
responsive to L-DOPA treatment as time progresses, and it can cause increased dopamine levels
elsewhere in the brain, which are associated with psychosis or schizophrenia.

 TERMS TO KNOW

Cerebrum
The superior region of the brain which controls higher neurological functions.

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 Cerebral Cortex
The outer gray matter region of the cerebrum.

Longitudinal Fissure
The sagittal deep groove which separates the cerebrum into right and left hemispheres.

Cerebral Hemisphere
Right and left halves of the cerebrum.

Frontal Lobe
The region of the cerebrum located under the frontal bone and responsible for motor and cognitive
functions.

Parietal Lobe
The region of the cerebrum located under the parietal bone and responsible for somatosensory
function.

Temporal Lobe
The region of the cerebrum located under the temporal bone and responsible for auditory function.

Occipital Lobe
The region of the cerebrum located under the occipital bone and responsible for visual function.

Insula
The region of the cerebrum located deep to the temporal and frontal lobes and responsible for
gustatory function.

Central Sulcus
The groove which separates the frontal and parietal lobes of the cerebrum.

Lateral Sulcus
The groove that separates the temporal lobe from the frontal and anterior parietal lobes of the
cerebrum.

Parieto-Occipital Sulcus
The groove that separates the medial occipital lobe from the parietal lobe of the cerebrum.

Precentral Gyrus
The cerebral gyrus located immediately anterior to the central sulcus.

Primary Motor Cortex

The region of the frontal lobe that is responsible for voluntary motor movement in the body, located in
the precentral gyrus.

Broca’s Area
The region of the frontal lobe that is responsible for production of meaningful language.

Prefrontal Cortex
The anterior region of the frontal lobe that is responsible for cognitive functions.

Postcentral Gyrus

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 The cerebral gyrus located immediately posterior to the central sulcus.

Primary Somatosensory Cortex

The region of the parietal lobe that is responsible for processing somatosensory sensory information,
located in the postcentral gyrus.

Proprioception
The sensation of where the body is in space.

Kinesthesia
The sensation of body movement.

Primary Auditory Cortex

The region of the temporal lobe that is responsible for processing auditory sensory information.

Wernicke’s Area
The region of the temporal lobe that is responsible for the comprehension of written and spoken
language.

Primary Gustatory Cortex

The region of the insula that is responsible for processing gustatory sensory information.

Primary Visual Cortex

The region of the occipital lobe that is responsible for processing visual sensory information.

Basal Nuclei
Subcortical groups of cell bodies that influence cortical motor signaling to maintain body position and
coordination.

 SUMMARY

In this lesson, you first learned about the subdivided structures of the diencephalon, the thalamus,
hypothalamus, and epithalamus as well as their unique functions. You then learned about the various
regions of the cerebrum. You learned about the lobes and surface landmarks in the cerebral cortex as
well as the functions of the various lobes and Brodmann’s areas. Lastly, you learned about the
subcortical structures that play a role in modifying cerebral activity.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Basal Nuclei

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 Subcortical groups of cell bodies that influence cortical motor signaling to maintain body position and
coordination.

Broca’s Area
The region of the frontal lobe that is responsible for production of meaningful language.

Central Sulcus
The groove which separates the frontal and parietal lobes of the cerebrum.

Cerebral Cortex
The outer gray matter region of the cerebrum.

Cerebral Hemisphere
Right and left halves of the cerebrum.

Cerebrum
The superior region of the brain which controls higher neurological functions.

Diencephalon
Region of the brain deep to the cerebrum and superior to the midbrain.

Epithalamus
Plays a role in the limbic system and the production of cerebrospinal fluid.

Frontal Lobe
The region of the cerebrum located under the frontal bone and responsible for motor and cognitive
functions.

Hypothalamus
Executive region in charge of the subconscious nervous system control (ANS) and the endocrine system.

Insula
The region of the cerebrum located deep to the temporal and frontal lobes and responsible for gustatory
function.

Kinesthesia
The sensation of body movement.

Lateral Sulcus
The groove that separates the temporal lobe from the frontal and anterior parietal lobes of the cerebrum.

Longitudinal Fissure
The sagittal deep groove which separates the cerebrum into right and left hemispheres.

Occipital Lobe
The region of the cerebrum located under the occipital bone and responsible for visual function.

Parietal Lobe
The region of the cerebrum located under the parietal bone and responsible for somatosensory function.

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 Parieto-Occipital Sulcus
The groove that separates the medial occipital lobe from the parietal lobe of the cerebrum.

Pineal Gland
Produces and releases melatonin to regulate circadian rhythms (sleep-wake cycles).

Postcentral Gyrus
The cerebral gyrus located immediately posterior to the central sulcus.

Precentral Gyrus
The cerebral gyrus located immediately anterior to the central sulcus.

Prefrontal Cortex
The anterior region of the frontal lobe that is responsible for cognitive functions.

Primary Auditory Cortex

The region of the temporal lobe that is responsible for processing auditory sensory information.

Primary Gustatory Cortex

The region of the insula that is responsible for processing gustatory sensory information.

Primary Motor Cortex

The region of the frontal lobe that is responsible for voluntary motor movement in the body, located in the
precentral gyrus.

Primary Somatosensory Cortex

The region of the parietal lobe that is responsible for processing somatosensory sensory information,
located in the postcentral gyrus.

Primary Visual Cortex

The region of the occipital lobe that is responsible for processing visual sensory information.

Proprioception
The sensation of where the body is in space.

Temporal Lobe
The region of the cerebrum located under the temporal bone and responsible for auditory function.

Thalamus
A collection of nuclei that relay information between the cerebral cortex and the periphery, spinal cord, or
brainstem.

Wernicke’s Area
The region of the temporal lobe that is responsible for the comprehension of written and spoken
language.

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 Meninges
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the three tissue layers that protect the central nervous system.
Specifically, this lesson will cover:
1. The Meninges: Protective Coverings of the Brain and Spinal Cord
1a. Dura Mater
1b. Arachnoid Mater
1c. Pia Mater

 BEFORE YOU START

The CNS is crucial to the operation and survival of the body, and any compromise in the brain and spinal
cord can lead to severe difficulties. Therefore, it is important to maintain its access to nutrients and waste
removal as well as protect it from harmful substances such as toxins or pathogens. The body maintains this
in two ways. First, the CNS has an extensive, privileged blood supply. Second, the CNS produces and
circulates cerebrospinal fluid (CSF).

1. The Meninges: Protective Coverings of the

Brain and Spinal Cord
The outer surface of the CNS is covered by a series of membranes composed of connective tissue called the
meninges. These membranes (dura mater, arachnoid mater, and pia mater, superficial to deep) serve to protect
the brain and spinal cord.

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 Meningeal Layers of Superior Sagittal Sinus - The layers of the meninges in the longitudinal fissure of the superior

sagittal sinus are shown with the dura mater adjacent to the inner surface of the cranium, the pia mater adjacent to
the surface of the brain, and the arachnoid and subarachnoid space between them. An arachnoid granulation villus is

shown emerging into the dural sinus to allow CSF to filter back into the blood for drainage.

1a. Dura Mater

The dura mater (dura, tough; mater, mother) is a thick, fibrous, outer layer and a strong protective sheath over
the entire brain and spinal cord. It is anchored to the inner surface of the cranium and vertebral cavity. It
encloses the entire CNS and the major blood vessels that enter the cranium and vertebral cavity.

There are infoldings of the dura that fit into large crevasses of the brain. Two infoldings go through the midline
separations of the cerebrum and cerebellum; one forms a shelf-like tent between the occipital lobes of the
cerebrum and the cerebellum, and the other surrounds the pituitary gland. The dura also surrounds and
supports the venous sinuses.

1b. Arachnoid Mater

The arachnoid mater is the middle layer of the meninges. The membrane is composed of a thin fibrous tissue
that forms a loose sac around the CNS. Beneath the arachnoid is a thin, filamentous mesh called the arachnoid
trabeculae, which looks like a spider web and gives this layer its name (arachnid, arthropod such as spider).

The space that the trabeculae extend through is called the subarachnoid space. This space is filled with
cerebrospinal fluid (CSF) that is circulating around the CNS. Around the brain, the arachnoid membrane forms
arachnoid granulations, periodic extensions into the superficial dural sinus (space within the dura mater), where

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 the CSF is filtered back into the blood for drainage from the nervous system.

Similar to clinical blood work, a sample of CSF can be withdrawn through a procedure called a lumbar puncture
or spinal tap. In a lumbar puncture, a needle is inserted most commonly into the space between L3–L4 until it
reaches the subarachnoid space. At this point, CSF is allowed to drain from this space. The CSF can then be
tested to see if it contains bacteria, a virus, various proteins or electrolytes, and more.

1c. Pia Mater

Directly superficial to the surface of the CNS is the pia mater (pia, tender), a thin fibrous membrane that follows
the convolutions of gyri and sulci in the cerebral cortex and fits into other grooves and indentations.

IN CONTEXT
Disorders of the Meninges

Meningitis is an inflammation of the meninges, the three layers of fibrous membrane that surround the
CNS. Meningitis can be caused by infection by bacteria or viruses. The particular pathogens are not
special to meningitis; it is just an inflammation of that specific set of tissues from what might be a
broader infection. Bacterial meningitis can be caused by Streptococcus, Staphylococcus, or the
tuberculosis pathogen, among many others. Viral meningitis is usually the result of common
enteroviruses (such as those that cause intestinal disorders) but may be the result of the herpes virus
or West Nile virus. Bacterial meningitis tends to be more severe.

The symptoms associated with meningitis can be fever, chills, nausea, vomiting, light sensitivity,
soreness of the neck, or severe headache. More important are the neurological symptoms, such as
changes in mental state (confusion, memory deficits, and other dementia-type symptoms). A serious
risk of meningitis can be damage to peripheral structures because of the nerves that pass through the
meninges. Hearing loss is a common result of meningitis.

The primary test for meningitis is a lumbar puncture. A needle inserted into the lumbar region of the
spinal column through the dura mater and arachnoid membrane into the subarachnoid space can be
used to withdraw the fluid for chemical testing. Fatality occurs in 5 to 40% of children and 20 to 50%
of adults with bacterial meningitis. Treatment of bacterial meningitis is through antibiotics, but viral
meningitis cannot be treated with antibiotics because viruses do not respond to that type of drug.
Fortunately, the viral forms are milder.

 TERMS TO KNOW

Meninges
Three connective tissue membranes which surround and protect the CNS.

Dura Mater
The outer, fibrous layer of the meninges.

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 Arachnoid Mater
The middle, spider-like layer of the meninges.

Arachnoid Trabeculae
Extensions of the arachnoid mater into the subarachnoid space that form a spider-like web.

Subarachnoid Space
The space in which the arachnoid trabeculae are located and cerebrospinal fluid circulates around the
CNS.

Arachnoid Granulations
Extensions of the arachnoid mater into the dural sinus in the cranial cavity.

Lumbar Puncture
A medical procedure that obtains a small amount of cerebrospinal fluid from the body.

Pia Mater
The inner, delicate layer of the meninges.

 SUMMARY

In this lesson, you learned about how the central nervous system is protected. You then learned to
identify the protective coverings of the brain and spinal cord called the meninges: dura mater,
arachnoid mater, and pia mater. You also learned to identify the spaces that they create and how
they serve to protect the various regions of the central nervous system.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Arachnoid Granulations
Extensions of the arachnoid mater into the dural sinus in the cranial cavity.

Arachnoid Mater
The middle, spider-like layer of the meninges.

Arachnoid Trabeculae
Extensions of the arachnoid mater into the subarachnoid space that form a spider-like web.

Dura Mater
The outer, fibrous layer of the meninges.

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 Lumbar Puncture
A medical procedure that obtains a small amount of cerebrospinal fluid from the body.

Meninges
Three connective tissue membranes which surround and protect the CNS.

Pia Mater
The inner, delicate layer of the meninges.

Subarachnoid Space
The space in which the arachnoid trabeculae are located and cerebrospinal fluid circulates around the
CNS.

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 Ventricular System of the Central Nervous
System
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about how the body supplies nutrients to and waste removal from the
central nervous system. Specifically, this lesson will cover:
1. Blood Supply to the Brain
2. The Ventricular System
2a. The Ventricles
2b. Cerebrospinal Fluid Circulation

1. Blood Supply to the Brain

A lack of oxygen in the CNS can be devastating. To counteract even short-term losses, the blood vessels
supplying blood to the brain have been designed with multiple pathways to the brain as well as regulatory
reflexes that ensure blood supply is not interrupted.

 THINK ABOUT IT

If you have ever taken the ‘back roads’ while driving due to traffic, you would likely agree that the more
pathways that exist to a given destination, the smoother traffic will travel at all times of the day. If, however,
there is only one road into or out of your destination, it only takes one event of flooding, traffic accident, or
rush hour to cause a backup.
In the CNS, there are multiple pathways for blood to get to the brain. You may have heard of the carotid or
vertebral arteries. If one gets blocked, blood can travel by the others to continue supplying the brain with
nutrients. Furthermore, these pathways fuse and diverge multiple times so that a blockage or narrowing in one
does not affect the perfusion of the CNS.

Additionally, when you stand up, blood tends to rush downwards, away from the brain, due to gravity. A reflex,
called the orthostatic reflex, exists to maintain blood pressure so that the brain continues to receive blood.

2. The Ventricular System

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 Cerebrospinal fluid (CSF) circulates throughout and around the CNS. In other tissues, water and small molecules
are filtered through blood vessels. However, in the CNS, recall that the blood-brain barrier makes it difficult for
many of those items to filter through blood vessels. Instead, CSF is produced in the brain in special structures to
perfuse through the CNS. Specifically, CSF circulates to remove metabolic wastes from nervous tissues and
return them to the bloodstream. The ventricles are four central cavities within the brain where CSF circulates. In
some of these spaces, CSF is produced by filtering of the blood that is performed by a specialized membrane
formed by ependymal cells. The CSF circulates through all of the ventricles to eventually emerge into the
subarachnoid space where it will be reabsorbed into the blood.

2a. The Ventricles

There are four ventricles within the brain. The first two are named the lateral ventricles, are deep within the
cerebrum, and look like stretched-out forms of the letter “C” with a tail. These ventricles are connected to the
third ventricle by two openings called the interventricular foramina (singular, foramen). The third ventricle is the
space between the left and right sides of the diencephalon, which opens into the cerebral aqueduct that
passes through the midbrain. The aqueduct opens into the fourth ventricle, which is the space between the
cerebellum and the pons and upper medulla. The fourth ventricle then narrows into the central canal of the
spinal cord. CSF also moves from the fourth ventricle into the subarachnoid space through three small
openings, the single median aperture or two lateral apertures.

Ventricular System - The ventricles are four interconnected cavities in the brain where cerebrospinal fluid is produced

and circulated.

2b. Cerebrospinal Fluid Circulation

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 Cerebrospinal fluid is produced within the ventricles by a type of specialized membrane called a choroid
plexus. Ependymal cells (one of the types of glial cells described in the introduction to the nervous system)
surround blood capillaries and filter the blood to make CSF. The fluid is a clear solution with a limited amount of
the constituents of blood. It is essentially water, small molecules, and electrolytes. Oxygen and carbon dioxide
are dissolved into the CSF, as they are in blood, and can diffuse between the fluid and the nervous tissue. The
choroid plexuses are found in all four ventricles. Observed in dissection, they appear as soft, fuzzy structures
that may still be pink, depending on how well the circulatory system is cleared in preparation of the tissue.
Because the CSF is produced from components extracted from the blood, its flow out of the ventricles is tied to
the pulse of cardiovascular circulation.

From the lateral ventricles, the CSF flows into the third ventricle, where more CSF is produced, and then
through the cerebral aqueduct into the fourth ventricle where even more CSF is produced. A very small amount
of CSF is filtered at any one of the plexuses, for a total of about 500 milliliters daily, but it is continuously made
and pulses through the ventricular system, keeping the fluid moving. From the fourth ventricle, CSF can
continue down the central canal of the spinal cord, but this is essentially a cul-de-sac, so more of the fluid
leaves the ventricular system and moves into the subarachnoid space through the median and lateral apertures.

Cerebrospinal Fluid Circulation - The choroid plexus in the four ventricles produce CSF, which is circulated through

the ventricular system and then enters the subarachnoid space through the median and lateral apertures. The CSF is

then reabsorbed into the blood at the arachnoid granulations, where the arachnoid membrane emerges into the dural

sinuses.

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 Within the subarachnoid space, the CSF flows around all of the CNS, providing two important functions. As with
elsewhere in its circulation, the CSF picks up metabolic wastes from the nervous tissue and moves it out of the
CNS. It also acts as a liquid cushion for the brain and spinal cord. By surrounding the entire system in the
subarachnoid space, it provides a thin buffer around the organs within the strong, protective dura mater. The
arachnoid granulations are outpocketings of the arachnoid membrane into the dural sinuses so that CSF can be
reabsorbed into the blood, along with the metabolic wastes. From the dural sinuses, blood drains out of the
head and neck, along with the rest of the circulation for blood, to be reoxygenated by the lungs and wastes to
be filtered out by the kidneys.

Components of CSF Circulation

Lateral Third Cerebral Central Subarachnoid

Fourth ventricle
ventricles ventricle aqueduct canal space

Between pons/upper
Location in Spinal
Cerebrum Diencephalon Midbrain medulla and Meninges
CNS cord
cerebellum

Blood vessel Choroid Choroid Arachnoid

None Choroid plexus None
structure plexus plexus granulations

IN CONTEXT
Disorders of the Central Nervous System

The supply of blood to the brain is crucial to its ability to perform many functions. Without a steady
supply of oxygen, and to a lesser extent glucose, the nervous tissue in the brain cannot keep up its
extensive electrical activity. These nutrients get into the brain through the blood, and if blood flow is
interrupted, neurological function is compromised.

The common name for a disruption of blood supply to the brain is a stroke. It is caused by a blockage
of an artery in the brain. The blockage is from some type of embolus: a blood clot, a fat embolus, or an
air bubble. When the blood cannot travel through the artery, the surrounding tissue that is deprived
starves and dies. Strokes will often result in the loss of very specific functions. A stroke in the lateral
medulla, for example, can cause a loss in the ability to swallow. Sometimes, seemingly unrelated
functions will be lost because they are dependent on structures in the same region. Along with the
swallowing in the previous example, a stroke in that region could affect sensory functions from the
face or extremities because important white matter pathways also pass through the lateral medulla.
Loss of blood flow to specific regions of the cortex can lead to the loss of specific higher functions,
from the ability to recognize faces to the ability to move a particular region of the body. Severe or
limited memory loss can be the result of a temporal lobe stroke.

Related to strokes are transient ischemic attacks (TIAs), which can also be called “mini-strokes.” These
are events in which a physical blockage may be temporary, cutting off the blood supply and oxygen to
a region, but not to the extent that it causes cell death in that region. While the neurons in that area

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 are recovering from the event, neurological function may be lost. Function can return if the area is able
to recover from the event. Recovery from a stroke (or TIA) is strongly dependent on the speed of
treatment. Often, the person who is present and notices something is wrong must then make a
decision. The mnemonic FAST helps people remember what to look for when someone is dealing with
sudden losses of neurological function. If someone complains of feeling “funny,” check these things
quickly: Look at the person’s face. Do they have problems moving Face muscles and making regular
facial expressions? Ask the person to raise their Arms above the head. Can the person lift one arm but
not the other? Has the person’s Speech changed? Are they slurring words or having trouble saying
things? If any of these things have happened, then it is Time to call for help.

Sometimes, treatment with blood-thinning drugs can alleviate the problem, and recovery is possible. If
the tissue is damaged, the amazing thing about the nervous system is that it is adaptable. With
physical, occupational, and speech therapy, victims of strokes can recover, or more accurately relearn,
functions.

 MAKE THE CONNECTION

If you're taking the Anatomy & Physiology I Lab course simultaneously with this lecture, it's a good time to
try the Lab Introduction to the Central Nervous System: Explore your body’s command center! in Unit 5 of
the Lab course. Review the lab-to-lecture crosswalk if you need more information. Good luck!

 TERMS TO KNOW

Ventricle
An open space within the brain where CSF is produced and circulates.

Lateral Ventricle
One of two large ventricles in the cerebrum.

Interventricular Foramina
Openings between the lateral ventricles and the third ventricle that allow for the passage of CSF.

Third Ventricle
A ventricle located in the diencephalon.

Cerebral Aqueduct
An opening between the third and fourth ventricles that allows for the passage of CSF.

Fourth Ventricle
A ventricle located in the brainstem.

Median Aperture
A single midline opening in the fourth ventricle that allows for passage of CSF into the subarachnoid
space.

Lateral Aperture

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 One of a pair of openings in the fourth ventricle that allows for passage of CSF into the subarachnoid
space.

Choroid Plexus
A tissue containing ependymal cells that filters blood to form CSF in the ventricles.

 SUMMARY

In this lesson, you learned about the specialized blood supply to the brain which maintains a
constant supply of blood to the CNS. You also learned about the ventricular system, the ventricles
of the brain which allow for cerebrospinal fluid circulation.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Cerebral Aqueduct
An opening between the third and fourth ventricles that allows for the passage of CSF.

Choroid Plexus
A tissue containing ependymal cells that filters blood to form CSF in the ventricles.

Fourth Ventricle
A ventricle located in the brainstem.

Interventricular Foramina
Openings between the lateral ventricles and the third ventricle that allow for the passage of CSF.

Lateral Aperture
One of a pair of openings in the fourth ventricle that allows for passage of CSF into the subarachnoid
space.

Lateral Ventricle
One of two large ventricles in the cerebrum.

Median Aperture
A single midline opening in the fourth ventricle that allows for passage of CSF into the subarachnoid
space.

Third Ventricle
A ventricle located in the diencephalon.

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 Ventricle
An open space within the brain where CSF is produced and circulates.

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 Peripheral Nervous System: Spinal Nerve
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of the peripheral nervous system including
spinal nerves. Specifically, this lesson will cover:
1. Nerves
2. Ganglia
3. Spinal Nerves

 BEFORE YOU START

The PNS is not as contained as the CNS because it is defined as everything that is not the CNS. Some
peripheral structures are even incorporated into the other organs of the body. In describing the anatomy of
the PNS, it is necessary to describe the common structures, the nerves, and the ganglia, as they are found
in various parts of the body. Many of the neural structures that are incorporated into other organs are
features of the digestive system; these structures are known as the enteric nervous system and are a
special subset of the PNS.

1. Nerves
 KEY CONCEPT

Recall that the term ‘nerves’ refers to a bundle of axons in the PNS whereas in the CNS, they are called a
tract. Nerves are composed of more than just nervous tissue. They have connective tissues invested in their
structure as well as blood vessels supplying the tissues with nourishment. The structure of a nerve is also
similar to that of a muscle if you recall how muscle fibers, muscle fascicles, and muscles are organized.
Each individual axon, whether myelinated or not, is wrapped by loose connective tissue called the
endoneurium. Multiple wrapped axons are then bundled together into a grouping called a neuron fascicle
which is wrapped in a fibrous connective tissue called perineurium. Finally, multiple fascicles are collected
together and wrapped one last time by a loose connective tissue called epineurium. These three layers are
similar to the connective tissue sheaths for muscles. Nerves are associated with the region of the CNS to which
they are connected, either as cranial nerves connected to the brain or spinal nerves connected to the spinal
cord.

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 Nerve Structure - The structure of a nerve is organized by the layers of connective tissue on the outside, around each
fascicle, and surrounding the individual nerve fibers (tissue source: simian). LM × 40.

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 Close-Up of Nerve Trunk - Zoom in on this slide of a nerve trunk to examine the endoneurium, perineurium, and

epineurium in greater detail (tissue source: simian). LM × 1600

 TERMS TO KNOW

Endoneurium
A loose connective tissue wrapping around axons in a nerve.

Neuron Fascicle
A bundle of axon fibers wrapped collectively in perineurium.

Perineurium
A fibrous connective tissue wrapping around a neuron fascicle in a nerve.

Epineurium
A loose connective tissue wrapping around a nerve.

2. Ganglia
Recall that a ganglia is a group of neuron cell bodies in the PNS whereas in the CNS it is called a nuclei. Ganglia
can be categorized, for the most part, as either sensory ganglia or autonomic ganglia, referring to their primary
functions. Sensory ganglia relay sensory information into the CNS while autonomic ganglia process motor
responses headed away from the CNS and towards their effector targets.

The most common type of sensory ganglion is a dorsal (posterior) root ganglion. Recall that these ganglia are
the cell bodies of neurons with axons that are sensory endings in the periphery, such as in the skin, and that
extend into the CNS through the posterior nerve root. The ganglion is an enlargement of the nerve root. Under

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 microscopic inspection, it can be seen to include the cell bodies of the neurons, as well as bundles of fibers that
are the posterior nerve root. The cells of the dorsal root ganglion are unipolar cells, classifying them by shape.
Also, the small round nuclei of satellite cells can be seen surrounding—as if they were orbiting—the neuron cell
bodies.

Posterior Root Ganglion - The cell bodies of sensory neurons, which are unipolar neurons by shape, are seen in this

photomicrograph. Also, the fibrous region is composed of the axons of these neurons that are passing through the

ganglion to be part of the posterior nerve root (tissue source: canine). LM × 40

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 Spinal Cord and Root Ganglion - The slide includes both a cross-section of the lumbar spinal cord and a section of the

dorsal root ganglion (tissue source: canine). LM × 1600.

Another type of sensory ganglion is a cranial nerve ganglion. This is analogous to the dorsal root ganglion,
except that it is associated with a cranial nerve instead of a spinal nerve. The roots of cranial nerves are within
the cranium, whereas the ganglia are outside the skull. The neurons of cranial nerve ganglia are also unipolar in
shape with associated satellite cells.

 TERM TO KNOW

Cranial Nerve Ganglion

A sensory ganglion of a cranial nerve.

3. Spinal Nerves
Spinal nerves are nerves connected to the spinal cord. Recall that all of the spinal nerves are combined sensory
and motor axons that separate into two nerve roots. The sensory axons enter the spinal cord as the posterior
root. The motor fibers emerge as the anterior nerve root.

 KEY CONCEPT

Recall that there are 31 spinal nerves, named for the level of the spinal cord at which each one emerges.
There are eight pairs of cervical nerves designated C1 to C8, twelve thoracic nerves designated T1 to T12,
five pairs of lumbar nerves designated L1 to L5, five pairs of sacral nerves designated S1 to S5, and one pair
of coccygeal nerves. The nerves are numbered from the superior to inferior positions and each emerges

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 from the vertebral column through the intervertebral foramen at its level. The first nerve, C1, emerges
between the first cervical vertebra and the occipital bone. The second nerve, C2, emerges between the first
and second cervical vertebrae. The same occurs for C3 to C7, but C8 emerges between the seventh
cervical vertebra and the first thoracic vertebra. For the thoracic and lumbar nerves, each one emerges
between the vertebra that has the same designation and the next vertebra in the column. The sacral nerves
emerge from the sacral foramina (small holes) along the length of that unique vertebra.
Spinal nerves extend outward from the vertebral column to innervate the periphery. However, the nerves in the
periphery are not straight continuations of the spinal nerves but rather the reorganization of the axons in those
nerves to follow different courses. Axons from different spinal nerves will combine with one another to become
a network of nerve fibers with no associated cell bodies, referred to as a nerve plexus. This occurs at four
locations along the length of the vertebral column. Distal to the nerve plexus, the nerves separate into systemic
nerves, major nerves of the peripheral system. In this instance, the word plexus is used to describe networks

Of the four nerve plexuses, two are found at the cervical level, one at the lumbar level, and one at the sacral
level. The cervical plexus is composed of axons from spinal nerves C1 through C5 and branches into nerves in
the posterior neck and head, as well as the phrenic nerve, which connects to the diaphragm at the base of the
thoracic cavity. The other plexus from the cervical level is the brachial plexus. Spinal nerves C4 through T1
reorganize through this plexus to give rise to the nerves of the arms, as the name brachial suggests. A large
nerve from this plexus is the radial nerve from which the axillary nerve branches to go to the armpit region. The
lumbar plexus arises from all the lumbar spinal nerves and gives rise to nerves innervating the pelvic region
and the anterior leg. The femoral nerve is one of the major nerves from this plexus. The sacral plexus comes
from the lower lumbar nerves L4 and L5 and the sacral nerves S1 to S4. The most significant systemic nerve to
come from this plexus is the sciatic nerve. The sciatic nerve extends across the hip joint and is most commonly
associated with the condition sciatica, which is the result of compression or irritation of the nerve or any of the
spinal nerves giving rise to it.

These plexuses are described as arising from spinal nerves and giving rise to certain systemic nerves, but they
contain fibers that serve sensory functions or fibers that serve motor functions. This means that some fibers
extend from cutaneous or other peripheral sensory surfaces and send action potentials into the CNS. Those are
axons of sensory neurons in the posterior root ganglia that enter the spinal cord through the posterior nerve
root. Other fibers are the axons of motor neurons of the anterior horn of the spinal cord, which emerge in the
anterior nerve root and send action potentials to cause skeletal muscles to contract in their target regions.

EXAMPLE The radial nerve contains fibers of cutaneous sensation in the arm, as well as motor fibers
that move muscles in the arm.
Spinal nerves of the thoracic region, T2 through T11, are not part of the plexuses but rather emerge and give
rise to the intercostal nerves found between the ribs, which articulate with the vertebrae surrounding the spinal
nerve.

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© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 106
 Nerve Plexuses of the Body - There are four main nerve plexuses in the human body. The cervical plexus supplies

nerves to the posterior head and neck, as well as to the diaphragm. The brachial plexus supplies nerves to the arm.
The lumbar plexus supplies nerves to the anterior leg. The sacral plexus supplies nerves to the posterior leg

 WATCH

Please watch the following video for more information on this topic.

 TERMS TO KNOW

Nerve Plexus
A network of nerves without cell bodies.

Systemic Nerve
A major nerve of the peripheral system, distal to a nerve plexus.

Cervical Plexus
A nerve plexus associated with the upper cervical spinal nerves.

Brachial Plexus
A nerve plexus associated with the lower cervical spinal nerves and first thoracic spinal nerve.

Lumbar Plexus
A nerve plexus associated with the lumbar spinal nerves.

Sacral Plexus
A nerve plexus associated with the sacral spinal nerves.

 SUMMARY

In this lesson, you learned about the structure of the peripheral nervous system. First, you learned
about the structure and organization of nerves. Then you learned about ganglia that play roles in the
function of both spinal and cranial nerves. Lastly, you learned more about the spinal nerves and how
they serve the peripheral body, including their function in providing spinal reflexes.

© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 107
 Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Brachial Plexus
A nerve plexus associated with the lower cervical spinal nerves and first thoracic spinal nerve.

Cervical Plexus
A nerve plexus associated with the upper cervical spinal nerves.

Cranial Nerve Ganglion

A sensory ganglion of a cranial nerve.

Endoneurium
A loose connective tissue wrapping around axons in a nerve.

Epineurium
A loose connective tissue wrapping around a nerve.

Lumbar Plexus
A nerve plexus associated with the lumbar spinal nerves.

Nerve Plexus
A network of nerves without cell bodies.

Neuron Fascicle
A bundle of axon fibers wrapped collectively in perineurium.

Perineurium
A fibrous connective tissue wrapping around a neuron fascicle in a nerve.

Sacral Plexus
A nerve plexus associated with the sacral spinal nerves.

Systemic Nerve
A major nerve of the peripheral system, distal to a nerve plexus.

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 Peripheral Nervous System: Reflexes
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of reflexes and how they protect the body.
Specifically, this lesson will cover:
1. Reflexes

1. Reflexes
A reflex is an automatic motor response to given sensory input. You may recall the homeostatic reflex arc which
describes the way in which sensory signals are generated, processed, and responded to. A reflex does just that
using the nervous system as a way to transport sensory and motor signals.

All reflexes include a sensory (afferent) nerve that transports the incoming sensory signal to the central nervous
system. Once this information is processed, all reflexes have a motor (efferent) nerve which transports the
outgoing motor signal to the effector target. Some reflexes additionally contain an interneuron located in the
CNS between the sensory and motor nerve that provides additional processing of sensory signals and transfers
them to the correct subsequent nerve(s).

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 Reflex Arc - Reflexes provide automatic transport of sensory and motor signals along a preset neuron circuit. Sensory

(afferent) nerves transport sensory signals from the sensor(s) to the CNS. In the CNS, different amounts of processing
occur depending on the complexity of the reflex. Motor (efferent) nerves transport motor signals from the CNS to the

effector(s).

 WATCH

Please watch the following video for more information on this topic.
Reflexes can be classified based on multiple criteria:

Their development
Innate reflexes are ones you are born with.
Acquired reflexes are ones you have learned over time.
The target effector
Somatic reflexes have skeletal muscle tissue as a target effector.
Autonomic reflexes have cardiac muscle, smooth muscle, or glandular tissue as a target effector.
The complexity of the circuit
Monosynaptic reflexes have only one synapse and two neurons.

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 Polysynaptic reflexes have more than one synapse and three or more neurons.
The processing site
Spinal reflexes are processed in the spinal cord.
Cranial reflexes are processed in the brain.

You may be familiar with certain reflexes of the body. Every reflex is unique in its location, processing speed,
and action. The patellar tendon (knee-jerk) reflex shown below is an innate somatic monosynaptic spinal reflex
performed at the doctor's office to test the health of your lower spinal cord. Using a reflex hammer, the doctor
will strike the patellar ligament on the distal end of the knee which causes the patellar tendon proximal to the
knee to stretch. This sensory signal is sent to the spinal cord, is directly transferred to the motor nerve for the
quadricep muscle group, and is sent back out. The effect is to shorten the stretched patellar tendon.

In contrast, when your hand touches a hot surface, the sensory signal travels to the spinal cord and requires
additional processing compared to the knee-jerk reflex. The hot surface signal is transferred first to an
interneuron and then transferred again to a motor nerve. This setup is a polysynaptic reflex and requires
additional time. You may notice that this type of reflex may not have as fast of a reaction compared to a
monosynaptic reflex.

Spinal Reflexes - Spinal reflexes are automatic motor responses to a given sensory input that is programmed to
protect the body. (A) The patellar tendon (knee-jerk) reflex is an innate somatic monosynaptic spinal reflex that

protects the quadricep from being overstretched. (B) An innate somatic polysynaptic spinal reflex protects your hand

when it touches hot surfaces.

 TERMS TO KNOW

Reflex
An automatic motor response to a given sensory signal.

Innate Reflex
A reflex that you are born with.

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 Acquired Reflex
A reflex that is learned over time.

Somatic Reflex
A reflex of the somatic nervous system involving a skeletal muscle effector.

Autonomic Reflex
A reflex of the autonomic nervous system involving cardiac muscle, smooth muscle, or glandular tissue
effectors

Monosynaptic Reflex
A reflex that has two neurons and one synapse.

Polysynaptic Reflex
A reflex that has three or more neurons and two or more synapses.

Spinal Reflex
A reflex that is processed in the spinal cord.

Cranial Reflex
A reflex that is processed in the brain.

 SUMMARY

In this lesson, you learned about reflexes. You learned how sensory and motor neurons of the central
nervous system coordinate functions to protect the body without the processing power of the brain.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Acquired Reflex
A reflex that is learned over time.

Autonomic Reflex
A reflex of the autonomic nervous system involving cardiac muscle, smooth muscle, or glandular tissue
effectors.

Cranial Reflex
A reflex that is processed in the brain.

Innate Reflex
A reflex that you are born with.

Monosynaptic Reflex

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 A reflex that has two neurons and one synapse.

Polysynaptic Reflex
A reflex that has three or more neurons and two or more synapses.

Reflex
An automatic motor response to a given sensory signal.

Somatic Reflex
A reflex of the somatic nervous system involving a skeletal muscle effector.

Spinal Reflex
A reflex that is processed in the spinal cord.

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 Peripheral Nervous System: Cranial Nerves
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of the peripheral nervous system
including cranial nerves. Specifically, this lesson will cover:
1. Cranial Nerves

1. Cranial Nerves
The nerves attached to the brain are the cranial nerves, which are primarily responsible for the sensory and
motor functions of the head and neck. There are twelve cranial nerves (CN), which are designated CN I through
CN XII for “Cranial Nerve,” using Roman numerals for 1 through 12.

 TRY IT

For our purposes, there are three Roman numerals that we need to be aware of in order to recognize all of
the 12 cranial nerve numbers. A single vertical line is the mark for 1, a letter V is the mark for 5, and a letter
X is the mark for 10.

I=1
V=5
X = 10

When marks follow one another, so long as the larger value comes first, they are additive, such as III is 3, VI
is six, and XVI is 16. However, when a smaller number precedes a larger, it is subtracted from it, such as IV is
4 (5 - 1 = 4) and IX is 9 (10 - 1 =9). With that in mind, below are the Arabic and Roman numerals for you to
review.

1=I
2 = II
3 = III
4 = IV
5=V
6 = VI

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 7 = VII
8 = VIII
9 = IX
10 =X
11 = XI
12 = XII

Cranial nerves can be classified based on the type of signal that they transport - sensory, motor, or both -
meaning that the axons in these nerves originate out of sensory ganglia external to the cranium or motor nuclei
within the brainstem. Sensory axons enter the brain to synapse in a nucleus. Motor axons connect to the
skeletal muscles of the head or neck. Three of the nerves are solely composed of sensory fibers; five are strictly
motor; and the remaining four are mixed nerves.

Below is a diagram of the location of each cranial nerve as well as a table that summarizes the general
information. Each nerve is listed below with a brief explanation:

The olfactory nerve (CN I) and optic nerve (CN II) are responsible for the sense of smell and vision,
respectively.
The oculomotor nerve (CN III) is responsible for eye movements by controlling four of the extraocular
muscles. It is also responsible for lifting the upper eyelid when the eyes point up, and for pupillary
constriction.
The trochlear nerve (CN IV) and the abducens nerve (CN VI) are both responsible for eye movement, but
do so by controlling different extraocular muscles.
The trigeminal nerve (CN V) is responsible for cutaneous sensations of the face and controlling the
muscles of mastication.
The facial nerve (CN VII) is responsible for the muscles involved in facial expressions, as well as part of the
sense of taste and the production of saliva.
The vestibulocochlear nerve (CN VIII) is responsible for the senses of hearing and balance.
The glossopharyngeal nerve (CN IX) is responsible for controlling muscles in the oral cavity and upper
throat, as well as part of the sense of taste and the production of saliva.
The vagus nerve (CN X) is responsible for contributing to homeostatic control of the organs of the thoracic
and upper abdominal cavities.
The spinal accessory nerve (CN XI) is responsible for controlling the muscles of the neck, along with
cervical spinal nerves.
The hypoglossal nerve (CN XII) is responsible for controlling the muscles of the lower throat and tongue.

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 The Cranial Nerves - The anatomical arrangement of the roots of the cranial nerves observed from an inferior view of
the brain.

 DID YOU KNOW

An exercise to help learn this sort of information is to generate a mnemonic using words that have personal
significance. A mnemonic is a sentence in which the initial letter of each word corresponds to the initial
letter in the name of each nerve. Below are two common mnemonics for the cranial nerve names:

“On Old Olympus’ Towering Tops, A Finn And German Viewed Some Hops.”

“On Occassion Our Trusty Truck Acts Funny. Very Good Vehicle Any How.”

Another important aspect of the cranial nerves that lends itself to a mnemonic is the functional role each
nerve plays. Recall that cranial nerves can transport sensory (S), motor (M), or both (B) signals.

“Some Say Marry Money, But My Brother Says Big Brains Matter More.”

Cranial Nerves

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 Peripheral Connection
Name of Nerve Function (S/M/B) Central Connection (nuclei)
(ganglion or muscle)

I Olfactory Smell (S) Olfactory bulb Olfactory epithelium

Retina (retinal ganglion

II Optic Vision (S) Hypothalamus/thalamus/midbrain
cells)

Extraocular muscles
(other 4), levator
Eye movements
III Oculomotor Oculomotor nucleus palpebrae superioris,
(M)
ciliary ganglion
(autonomic)

Eye movements Superior oblique

IV Trochlear Trochlear nucleus
(M) muscle

Sensory/motor – Trigeminal nuclei in the midbrain,

V Trigeminal Trigeminal
face (B) pons, and medulla

Eye movements
VI Abducens Abducens nucleus Lateral rectus muscle
(M)

Facial muscles,
Taste (B) Facial nucleus, solitary Geniculate ganglion,
VII Facial Motor – face,
nucleus, superior salivatory nucleus Pterygopalatine
ganglion (autonomic)

Spiral ganglion
Auditory Hearing/balance Cochlear nucleus, Vestibular
VIII (hearing), Vestibular
(Vestibulocochlear) (S) nucleus/cerebellum
ganglion (balance)

Pharyngeal muscles,
Motor – throat Solitary nucleus, inferior salivatory Geniculate ganglion,
IX Glossopharyngeal
Taste (B) nucleus, nucleus ambiguus Otic ganglion
(autonomic)

Terminal ganglia serving

Motor/sensory – thoracic and upper
X Vagus viscera Medulla abdominal organs
(autonomic) (B) (heart and small
intestines)

Accessory/Spinal Motor – head and

XI Spinal accessory nucleus Neck muscles
Accessory neck (M)

Motor – lower Muscles of the larynx

XII Hypoglossal Hypoglossal nucleus
throat (M) and lower pharynx

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 IN CONTEXT
Aging and the Nervous System

Anosmia is the loss of the sense of smell. It is often the result of the olfactory nerve being severed,
usually because of blunt force trauma to the head. The sensory neurons of the olfactory epithelium
have a limited lifespan of approximately one to four months, and new ones are made on a regular
basis. The new neurons extend their axons into the CNS by growing along the existing fibers of the
olfactory nerve. The ability of these neurons to be replaced is lost with age. Age-related anosmia is
not the result of impact trauma to the head but rather a slow loss of the sensory neurons with no new
neurons born to replace them.

Smell is an important sense, especially for the enjoyment of food. There are only five tastes sensed by
the tongue, and two of them are generally thought of as unpleasant tastes (sour and bitter). The rich
sensory experience of food is the result of odor molecules associated with the food, both as food is
moved into the mouth and therefore passes under the nose and as it is chewed and molecules are
released to move up the pharynx into the posterior nasal cavity. Anosmia results in a loss of the
enjoyment of food.

As the replacement of olfactory neurons declines with age, anosmia can set in. Without the sense of
smell, many sufferers complain of food tasting bland. Often, the only way to enjoy food is to add
seasoning that can be sensed on the tongue, which usually means adding table salt. The problem with
this solution, however, is that this increases sodium intake, which can lead to cardiovascular problems
through water retention and the associated increase in blood pressure.

 TERMS TO KNOW

Cranial Nerve
One of twelve sets of nerves transporting sensory and motor information into and out of the brain.

Olfactory Nerve
A sensory cranial nerve responsible for the sense of smell; CN I.

Optic Nerve
A sensory cranial nerve responsible for the sense of vision; CN II.

Oculomotor Nerve
A motor cranial nerve responsible for the movement of the eye and pupil; CN III.

Trochlear Nerve
A motor cranial nerve responsible for the movement of the eye; CN IV.

Trigeminal Nerve
A sensory and motor cranial nerve responsible for cutaneous sensations of the face and controlling the
muscles of mastication.

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 Abducens Nerve
A motor cranial nerve responsible for the movement of the eye; CN VI.

Facial Nerve
A sensory and motor cranial nerve responsible for the muscles involved in facial expressions as well as
part of the sense of taste.

Vestibulocochlear Nerve
A sensory cranial nerve responsible for the sense of hearing and equilibrium.

Glossopharyngeal Nerve
A sensory and motor cranial nerve responsible for controlling muscles in the oral cavity and upper
throat as well as part of the sense of taste.

Vagus Nerve
A sensory and motor cranial nerve responsible for contributing to homeostatic control of the organs of
the thoracic and upper abdominal cavities.

Accessory Nerve
A motor cranial nerve responsible for controlling the muscles of the neck, along with cervical spinal
nerves (also called Spinal Accessory Nerve).

Hypoglossal Nerve
A motor cranial nerve responsible for controlling the muscles of the lower throat and tongue.

 SUMMARY

In this lesson, you learned the twelve cranial nerves. You learned how they are numbered, their
identity, and their function—whether sensory, motor, or both.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Abducens Nerve
A motor cranial nerve responsible for the movement of the eye; CN VI.

Accessory Nerve
A motor cranial nerve responsible for controlling the muscles of the neck, along with cervical spinal
nerves (also called Spinal Accessory Nerve).

Cranial Nerve

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 One of twelve sets of nerves transporting sensory and motor information into and out of the brain.

Facial Nerve
A sensory and motor cranial nerve responsible for the muscles involved in facial expressions as well as
part of the sense of taste.

Glossopharyngeal Nerve
A sensory and motor cranial nerve responsible for controlling muscles in the oral cavity and upper throat
as well as part of the sense of taste.

Hypoglossal Nerve
A motor cranial nerve responsible for controlling the muscles of the lower throat and tongue.

Oculomotor Nerve
A motor cranial nerve responsible for the movement of the eye and pupil; CN III.

Olfactory Nerve
A sensory cranial nerve responsible for the sense of smell; CN I.

Optic Nerve
A sensory cranial nerve responsible for the sense of vision; CN II.

Trigeminal Nerve
A sensory and motor cranial nerve responsible for cutaneous sensations of the face and controlling the
muscles of mastication.

Trochlear Nerve
A motor cranial nerve responsible for the movement of the eye; CN IV.

Vagus Nerve
A sensory and motor cranial nerve responsible for contributing to homeostatic control of the organs of the
thoracic and upper abdominal cavities.

Vestibulocochlear Nerve
A sensory cranial nerve responsible for the sense of hearing and equilibrium.

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 Sensory Perception
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the types of sensory receptors. Specifically, this lesson will cover:
1. Sensation vs Perception
2. Sensory Receptors
2a. Structural Receptor Types
2b. Functional Receptor Types

1. Sensation vs Perception
 KEY CONCEPT

A major role of sensory receptors is to help us learn about the environment around us or about the state of
our internal environment. Stimuli from varying sources, and of different types, are received and changed
into the electrochemical signals of the nervous system. This occurs when a stimulus changes the cell
membrane potential of a sensory neuron. The stimulus causes the sensory cell to produce an action
potential that is relayed into the central nervous system (CNS), where it is integrated with other sensory
information—or sometimes higher cognitive functions—to become a conscious perception of that stimulus.
The central integration may then lead to a motor response.
Describing sensory function with the term sensation or perception is a deliberate distinction. Sensation is the
activation of sensory receptor cells at the level of the stimulus. Perception is the central processing of sensory
stimuli into a meaningful pattern. Perception is dependent on sensation but not all sensations are perceived.
Receptors are the cells or structures that detect sensations. A receptor cell is changed directly by a stimulus. A
transmembrane protein receptor is a cell membrane protein that mediates a physiological change in a neuron.
Transmembrane receptors are activated by chemicals called ligands. For example, a molecule in food can serve
as a ligand for taste receptors. Other transmembrane proteins are sensitive to mechanical or thermal changes.
Physical changes in these proteins increase ion flow across the membrane and can generate an action
potential or a graded potential in the sensory neurons.

 TERMS TO KNOW

Sensation
The activation of a sensory receptor cell.

Perception

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 The central processing of sensory stimuli into a meaningful pattern.

2. Sensory Receptors
Stimuli in the environment activate specialized receptor cells in the peripheral nervous system. Different types
of stimuli are sensed by different types of receptor cells. Receptor cells can be classified into types on the basis
of three different criteria:

Cell Type
Position
Function

Receptors can be classified structurally on the basis of cell type and their position in relation to the stimuli they
sense. They can also be classified functionally on the basis of the transduction of stimuli, or how the mechanical
stimulus, light, or chemical changed the cell membrane potential.

2a. Structural Receptor Types

As shown in the image below, the cells that interpret information about the environment can be either:

a. A neuron that has a free nerve ending, or non-encapsulated dendrites, embedded in tissue that would
receive a sensation.
b. A neuron that has an encapsulated ending in which the sensory nerve endings are encapsulated in
connective tissue that enhances their sensitivity.
c. A specialized receptor cell, which has distinct structural components that interpret a specific type of
stimulus.

The pain and temperature receptors in the dermis of the skin are examples of neurons that have free nerve
endings. Also located in the dermis of the skin are lamellated corpuscles, neurons with encapsulated nerve
endings that respond to pressure and touch. The cells in the retina that respond to light stimuli are an example
of a specialized receptor, a photoreceptor.

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 Receptor Classification by Cell Type - Receptor cell types can be classified on the basis of their structure. Sensory
neurons can have either (a) free nerve endings or (b) encapsulated endings. Photoreceptors in the eyes, such as rod

cells, are examples of (c) specialized receptor cells.

Another way that receptors can be classified is based on their location relative to the stimuli. An exteroceptor is
a receptor that is located near a stimulus in the external environment, such as the somatosensory receptors that
are located in the skin. An interoceptor is one that interprets stimuli from internal organs and tissues, such as
the receptors that sense the increase in blood pressure in the aorta or carotid sinus. A proprioceptor is an
interoceptor located near a moving part of the body, such as a muscle, that interprets the positions of the
tissues as they move.

2b. Functional Receptor Types

Receptor cells can be further categorized on the basis of the type of stimuli they transduce. Chemical stimuli
can be interpreted by a chemoreceptor that interprets chemical stimuli, such as an object’s taste or smell.
Osmoreceptors respond to solute concentrations of body fluids. Additionally, pain is primarily a chemical sense
that interprets the presence of chemicals from tissue damage, or similar intense stimuli, through a nociceptor.
Thermoreceptors are either sensitive to temperatures above (heat) or below (cold) normal body temperature.
Physical stimuli are interpreted through a mechanoreceptor. There are three types of mechanoreceptors—
proprioceptors, baroreceptors, and tactile receptors. Recall that proprioceptors detect the position of the body
in space. Baroreceptors detect changes in pressure such as blood pressure as well as others in the digestive,
respiratory, and urinary systems. Tactile receptors detect touch, pressure, and vibration.

 TERMS TO KNOW

Encapsulated Ending
A sensory nerve ending encapsulated in connective tissue.

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 Receptor Cell
A unique sensory cell with structural components that interpret a specific type of stimulus.

Exteroceptor
A receptor that is located near a stimulus in the external environment.

Interoceptor
A receptor that interprets stimuli from internal organs and tissues.

Proprioceptor
An interoceptor located near a moving part of the body that interprets the positions of the tissues as
they move.

Chemoreceptor
A sensory receptor that detects chemical stimuli.

Osmoreceptor
A sensory receptor that detects changes in solute concentration.

Nociceptor
A sensory receptor that detects pain stimuli.

Thermoreceptor
A sensory receptor that detects changes in temperature.

Mechanoreceptor
A sensory receptor that detects physical stimuli.

Baroreceptor
A sensory receptor that detects changes in barometric pressure.

Tactile Receptor
A sensory receptor that detects touch, pressure, and vibration.

 SUMMARY

In this lesson, you learned about sensations. First, you learned to identify the difference between a
sensation and perception. Then, you learned the types of sensory receptors, categorized as structural
receptor types or functional receptor types.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Baroreceptor
A sensory receptor that detects changes in barometric pressure.

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 Chemoreceptor
A sensory receptor that detects chemical stimuli.

Encapsulated Ending
A sensory nerve ending encapsulated in connective tissue.

Exteroceptor
A receptor that is located near a stimulus in the external environment.

Interoceptor
A receptor that interprets stimuli from internal organs and tissues.

Mechanoreceptor
A sensory receptor that detects physical stimuli.

Nociceptor
A sensory receptor that detects pain stimuli.

Osmoreceptor
A sensory receptor that detects changes in solute concentration.

Perception
The central processing of sensory stimuli into a meaningful pattern.

Proprioceptor
An interoceptor located near a moving part of the body that interprets the positions of the tissues as they
move.

Receptor Cell
A unique sensory cell with structural components that interpret a specific type of stimulus.

Sensation
The activation of a sensory receptor cell.

Tactile Receptor
A sensory receptor that detects touch, pressure, and vibration.

Thermoreceptor
A sensory receptor that detects changes in temperature.

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 General Senses
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the types of sensory modalities and general senses. Specifically,
this lesson will cover:
1. Sensory Modalities
2. Somatosensation (Touch)

1. Sensory Modalities
 THINK ABOUT IT

Ask anyone what the senses are and they are likely to list the five major senses—taste, smell, touch,
hearing, and sight. However, these are not all of the senses. The most obvious omission from this list is
balance. Other overlooked senses include temperature perception by thermoreceptors and pain perception
by nociceptors.
Within the realm of physiology, senses can be classified as either general or specific:

A general sense is one that is distributed throughout the body and has receptor cells within the structures
of other organs. Mechanoreceptors in the skin, muscles, or the walls of blood vessels are examples of this
type. General senses often contribute to the sense of touch, as described above, or to proprioception (body
movement) and kinesthesia (body movement), or to a visceral sense, which is most important to autonomic
functions.
A special sense is one that has a specific organ devoted to it, namely the eye, inner ear, tongue, or nose.

Each of the senses is referred to as a sensory modality. Modality refers to the way that information is encoded,
which is similar to the idea of transduction. The main sensory modalities can be described on the basis of how
each is transduced. The chemical senses are taste and smell. The general sense that is usually referred to as
touch includes chemical sensation in the form of nociception, or pain. Pressure, vibration, muscle stretch, and
the movement of hair by an external stimulus, are all sensed by mechanoreceptors. Mechanoreceptors also
sense hearing and balance. Finally, vision involves the activation of photoreceptors.

Listing all the different sensory modalities, which can number as many as 17, involves separating the five major
senses into more specific categories, or submodalities, of the larger sense. An individual sensory modality
represents the sensation of a specific type of stimulus.

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 EXAMPLE The general sense of touch, which is known as somatosensation, can be separated into light
pressure, deep pressure, vibration, itch, pain, temperature, or hair movement.

 TERMS TO KNOW

General Sense
A sensory system that is widely distributed throughout the body and in the organs of the body.

Special Sense
A sensory system that has a specific organ devoted to it.

2. Somatosensation (Touch)
Somatosensation is the group of sensory modalities that are associated with touch, proprioception, and
interoception. These modalities are listed below:

Pressure
Vibration
Light touch
Tickle
Itch
Temperature
Pain
Proprioception
Kinesthesia

Somatosensation is considered a general sense, meaning that its receptors are not associated with a
specialized organ but are instead spread throughout the body in a variety of organs. Many of the
somatosensory receptors are located in the skin, but receptors are also found in muscles, tendons, joint
capsules, ligaments, and in the walls of visceral organs.

Two types of somatosensory signals that are transduced by free nerve endings are pain and temperature.
These two modalities use thermoreceptors and nociceptors to transduce temperature and pain stimuli,
respectively. Temperature receptors are stimulated when local temperatures differ from body temperature.
Some thermoreceptors are sensitive to just cold and others to just heat. Nociception is the sensation of
potentially damaging stimuli. Mechanical, chemical, or thermal stimuli beyond a set threshold will elicit painful
sensations. Stressed or damaged tissues release chemicals that activate receptor proteins in the nociceptors.

EXAMPLE The sensation of heat associated with spicy foods involves capsaicin, the active molecule in
hot peppers.
Capsaicin molecules bind to a transmembrane ion channel in nociceptors that is sensitive to temperatures
above 37°C. The dynamics of capsaicin binding with this transmembrane ion channel is unusual in that the
molecule remains bound for a long time. Because of this, it will decrease the ability of other stimuli to elicit pain

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 sensations through the activated nociceptor. For this reason, capsaicin can be used as a topical analgesic, such
as in products such as Icy Hot™.

The skin (integument) contains a variety of sensory receptors specific to the detection of pressure, vibration,
and light touch. For the benefit of review, each of these receptors is summarized again below. If you drag your
finger across a textured surface, the skin of your finger will vibrate. Such low-frequency vibrations are sensed
by mechanoreceptors called tactile discs, also known as Merkel discs or type I cutaneous mechanoreceptors.
Merkel cells are located in the epidermis's stratum basale (deepest layer). Deep pressure and vibration are
transduced by lamellar corpuscles, also known as Pacinian corpuscles, which are receptors with encapsulated
endings found deep in the dermis, or subcutaneous tissue. A light touch is transduced by the encapsulated
endings known as tactile corpuscles, also known as Meissner corpuscles. Follicles are also wrapped in a plexus
of nerve endings known as the hair follicle plexus. These nerve endings detect the movement of hair at the
surface of the skin, such as when an insect may be walking along the skin. Stretching of the skin is transduced
by stretch receptors known as bulbous corpuscles, also known as Ruffini corpuscles or type II cutaneous
mechanoreceptors.

Other somatosensory receptors are found in the joints and muscles. Stretch receptors monitor the stretching of
tendons, muscles, and the components of joints in order to determine the position of the body in space
(proprioception) and the current movement of the body (kinesthesia). These sensory signals help provide
information about how the body is currently moving (kinesthesia), where the body is in space (proprioception),
and if the muscle or joint is reaching the end of its range of motion, hoping to avoid overstretching it. In skeletal
muscle tissue, these stretch receptors are called muscle spindles. Golgi tendon organs similarly transduce the
stretch levels of tendons. Bulbous corpuscles are also present in joint capsules, where they measure stretch in
the components of the skeletal system within the joint. The types of nerve endings, their locations, and the
stimuli they transduce are presented in the table below.

Mechanoreceptors of Somatosensation

Historical
Name Location(s) Stimuli
(eponymous) name

Free nerve Dermis, cornea, tongue, joint Pain, temperature, mechanical

n/a
endings capsules, visceral organs deformation

Epidermal-dermal junction,
Tactile discs Merkel’s discs Low frequency vibration (5–15 Hz)
mucosal membranes

Bulbous
Ruffini’s corpuscle Dermis, joint capsules Stretch
corpuscle

Tactile Papillary dermis, especially in the Light touch, vibrations below 50

Meissner’s corpuscle
corpuscle fingertips and lips Hz

Lamellar Deep dermis, subcutaneous Deep pressure, high-frequency

Pacinian corpuscle
corpuscle tissue vibration (around 250 Hz)

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 Hair follicle Wrapped around hair follicles in
n/a Movement of hair
plexus the dermis

Muscle spindle n/a In line with skeletal muscle fibers Muscle contraction and stretch

Tendon stretch
Golgi tendon organ In line with tendons Stretch of tendons
organ

 TERMS TO KNOW

Capsaicin
The active molecule in hot peppers which creates the sensation of heat from spicy foods.

Muscle Spindles
A mechanoreceptor in skeletal muscles that detects stretch.

Golgi Tendon Organ

A mechanoreceptor in tendons that detects stretch.

 SUMMARY

In this lesson, you first learned how to categorize sensory modalities in the body. Then, you learned the
details of the general sense somatosensation (touch) is detected in the body.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Capsaicin
The active molecule in hot peppers which creates the sensation of heat from spicy foods.

General Sense
A sensory system that is widely distributed throughout the body and in the organs of the body.

Golgi Tendon Organ

A mechanoreceptor in tendons that detects stretch.

Muscle Spindles
A mechanoreceptor in skeletal muscles that detects stretch.

Special Sense
A sensory system that has a specific organ devoted to it.

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 Special Senses: Taste and Smell
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the special senses, taste and smell. Specifically, this lesson will cover:
1. Olfaction (Smell)
2. Gustation (Taste)
2a. Gustatory Structures
2b. Tastes
2c. Gustatory and Olfactory Integration

1. Olfaction (Smell)
Olfaction is the special sense of smell and makes use of the nose to turn chemical stimuli into neural signals.
The olfactory receptor neurons are located in a small region within the superior nasal cavity. The nasal cavity is
covered by olfactory epithelium which contains bipolar sensory neurons. Each olfactory sensory neuron has
dendrites that extend from the apical surface of the epithelium into the mucus lining the cavity. As airborne
molecules called odorants are inhaled through the nose; meaning, they pass over the olfactory epithelial region
and dissolve into the mucus. Odorants bind to proteins that keep them dissolved in the mucus and help
transport them to the olfactory dendrites. The odorant-protein complex binds to a receptor on the dendrite of
an olfactory sensory neuron and produces a graded membrane potential.

The axon of an olfactory neuron extends from the basal surface of the epithelium, through a hole in the
cribriform plate of the ethmoid bone, and into the brain to the olfactory bulb, a collection of secondary olfactory
sensory neurons. From the olfactory bulb, a group of axons called the olfactory tract connect to several brain
regions. Some travel to the cerebrum, specifically to the primary olfactory cortex which is located in the inferior
and medial areas of the temporal lobe. Others project to structures within the limbic system and hypothalamus,
where smells become associated with long-term memory and emotional responses.

 DID YOU KNOW

This is how certain smells trigger emotional memories, such as the smell of food associated with one’s
birthplace. Smell is the one sensory modality that does not synapse in the thalamus before connecting to
the cerebral cortex. This intimate connection between the olfactory system and the cerebral cortex is one
reason why smell can be a particularly strong trigger of memories and emotions.

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 The nasal epithelium, including the olfactory cells, can be harmed by airborne toxic chemicals. Therefore, the
olfactory neurons are regularly replaced within the nasal epithelium, after which the axons of the new neurons
must find their appropriate connections in the olfactory bulb. These new axons grow along the axons that are
already in place in the cranial nerve (olfactory nerve, CN I).

 REFLECT

Think about the last time you had a stuffy nose. Maybe you had a cold or similar upper respiratory infection.
Did you notice your sense of smell diminish during your illness?

Recall those olfactory sensory neurons only extend into the top third of the nasal cavity. When illnesses
cause an increase in mucus production, it becomes increasingly difficult for odorants to be transported to
the dendrites. This causes your sense of smell to diminish temporarily until the mucus can be shed or
thinned.

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 The Olfactory System - (a) The olfactory system begins in the peripheral structures of the nasal cavity. (b) The olfactory

receptor neurons are within the olfactory epithelium. (c) Axons of the olfactory receptor neurons project through the
cribriform plate of the ethmoid bone and synapse with the neurons of the olfactory bulb (tissue source: simian). LM ×

812.

IN CONTEXT
Disorders of the Olfactory System: Anosmia
Blunt force trauma to the face, such as that common in many car accidents, can lead to the loss of the

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 olfactory nerve and, subsequently, loss of the sense of smell. This condition is known as anosmia.
When the frontal lobe of the brain moves relative to the ethmoid bone, the olfactory tract axons may
be sheared apart. Professional fighters often experience anosmia because of repeated trauma to the
face and head. In addition, certain pharmaceuticals, such as antibiotics, can cause anosmia by killing
all the olfactory neurons at once. If no axons are in place within the olfactory nerve, then the axons
from newly formed olfactory neurons have no guide to lead them to their connections within the
olfactory bulb. There are temporary causes of anosmia as well, such as those caused by inflammatory
responses related to respiratory infections or allergies.

Loss of the sense of smell can result in food tasting bland. A person with an impaired sense of smell
may require additional spice and seasoning levels for food to be tasted. Anosmia may also be related
to some presentations of mild depression because the loss of enjoyment of food may lead to a
general sense of despair.

The ability of olfactory neurons to replace themselves decreases with age, leading to age-related
anosmia. This explains why some elderly people salt their food more than younger people do.
However, this increased sodium intake can increase blood volume and blood pressure, increasing the
risk of cardiovascular diseases in the elderly.

 TERMS TO KNOW

Olfaction
The sense of smell.

Olfactory Epithelium
The tissue covering the nasal cavity.

Olfactory Sensory Neuron

Afferent sensory neurons in the nose that are sensitive to odorants; part of the olfactory nerve.

Odorant
Chemical molecules able to bind and activate olfactory sensory neurons.

Olfactory Bulb
An enlargement of the olfactory nerve.

Olfactory Tract
The portion of the olfactory nerve that extends from the olfactory bulb to various regions of the brain.

2. Gustation (Taste)
2a. Gustatory Structures

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 Gustation is the special sense of taste that makes use of the tongue to create signals to chemical stimuli. The
surface of the tongue, along with the rest of the oral cavity, is lined by a stratified squamous epithelium. On the
tongue are raised bumps called gustatory papillae (singular, papilla) that are formed by folds in the surface of
the tongue. There are four types of papillae, based on their appearance:

Circumvallate
Foliate
Filiform
Fungiform

Within each papilla are taste buds located next to exposed surfaces or invaginations. The cells within the taste
bud are able to provide taste sensation. Gustatory receptor cells are specialized receptor cells that are
sensitive to the presence of taste stimuli. These cells project microvilli called taste hairs into a small opening in
the tongue epithelium called a taste pore. Here, taste receptors can bind their specific stimulus if it is present.
Gustatory receptor cells live for only approximately 10 days and are then replaced by stem cells known as basal
cells. Immature gustatory receptor cells are called transition cells.

2b. Tastes
Until recently, only four tastes were recognized: sweet, salty, sour, and bitter. Research at the turn of the 20th
century led to recognition of the fifth taste, umami, during the mid-1980s. Umami is a Japanese word that is
often translated to mean savory. Additional research has suggested that there may also be more tastes for fats
(lipids), water, calcium, and more.

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 The Tongue - The tongue is covered with small bumps, called papillae, which contain taste buds that are sensitive to

chemicals in ingested food or drink. Different types of papillae are found in different regions of the tongue. The taste

buds contain specialized gustatory receptor cells that respond to chemical stimuli dissolved in the saliva. These

receptor cells activate sensory neurons that are part of the facial and glossopharyngeal nerves. LM × 1600.

Salty taste is simply the perception of sodium ions (Na⁺) in the saliva. When you eat something salty, the salt
crystals dissociate into the component ions Na⁺ and Cl⁻, which dissolve into the saliva in your mouth. The Na⁺
concentration becomes high outside the gustatory cells, creating a strong concentration gradient that drives the
diffusion of the ion into the cells through open sodium channels. The entry of Na⁺ into these cells results in the
depolarization of the cell membrane.

Sour taste is the perception of H⁺ concentration. Just as with sodium ions in salty flavors, these hydrogen ions
enter the cell and trigger depolarization. Sour flavors are, essentially, the perception of acids in our food.
Increasing hydrogen ion concentrations in the saliva (lowering saliva pH) triggers progressively stronger graded
potentials in the gustatory cells, causing the influx of H⁺ through hydrogen channels and depolarization.

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 EXAMPLE Orange juice—which contains citric acid—will taste sour because it has a pH value of
approximately 3. Of course, it is often sweetened so that the sour taste is masked.
Sweet, bitter, and umami tastes result from food molecules binding to an external receptor which triggers
depolarization of the gustatory cell.

The sweet taste is the sensitivity of gustatory cells to the presence of glucose dissolved in the saliva. Other
monosaccharides such as fructose, or artificial sweeteners such as aspartame (NutraSweet™), saccharine, or
sucralose (Splenda™) also activate the sweet receptors. The affinity for each of these molecules varies and some
will taste sweeter than glucose because they bind to the G protein–coupled receptor differently.

Table: Comparison of Sweetness Intensity for USDA Approved Sweeteners (Edited table from U.S. Food &
Drug Administration)

Compound Example of Brand Name Sweetness Intensity Compared to Table Sugar (Sucrose)

Acesulfame Potassium Sweet One ® 200 X

Aspartame Nutrasweet ®, Equal ® 200 X

Steviol Glycosides Truvia ® 200–400 X

Sucralose Splenda ® 600 X

Saccharin Sweet and Low ® 200–700 X

Neotame Newtame ® 7,000–13,000 X

Advantame N/A 20,000 X

Bitter taste is similar to sweet in that food molecules bind to receptors. However, there are a number of different
ways in which this can happen because there is a large diversity of bitter-tasting molecules. Some bitter
molecules depolarize gustatory cells whereas others hyperpolarize gustatory cells. Likewise, some bitter
molecules increase receptor activation within the gustatory cells whereas other bitter molecules decrease
receptor activation. The specific response depends on which molecule is binding to the receptor.

One major group of bitter-tasting molecules is alkaloids. Alkaloids are nitrogen-containing molecules that are
commonly found in bitter-tasting plant products, such as coffee, hops (in beer), tannins (in wine), tea, and
aspirin. By containing toxic alkaloids, the plant is less susceptible to microbial infection and less attractive to
herbivores. Therefore, the function of bitter taste may primarily be related to stimulating the gag reflex to avoid
ingesting poisons. Because of this, many bitter foods that are normally ingested are often combined with a
sweet component to make them more palatable (cream and sugar in coffee, for example).

The taste known as umami is often referred to as the savory taste. Like sweet and bitter, it is based on the
activation of receptors by a specific molecule. The molecule that activates this receptor is an amino acid, L-
glutamate. Therefore, the umami flavor is often perceived while eating protein-rich foods. Not surprisingly,
dishes that contain meat are often described as savory.

Once the gustatory receptor cells are activated by the taste molecules, they release neurotransmitters onto the
dendrites of sensory neurons. These neurons are part of the facial and glossopharyngeal cranial nerves, as well

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 as a component within the vagus nerve dedicated to the gag reflex. The facial nerve connects to taste buds in
the anterior third of the tongue. The glossopharyngeal nerve connects to taste buds in the posterior two-thirds
of the tongue. The vagus nerve connects to taste buds in the extreme posterior of the tongue, verging on the
pharynx, which are more sensitive to noxious stimuli such as bitterness.

2c. Gustatory and Olfactory Integration

 KEY CONCEPT

When you say you taste something, that perception is generated based on a number of sensations, not just
from the tongue. Your brain relies heavily on both the gustatory and olfactory senses to determine taste. If
either one is temporarily blocked because of respiratory illness (loss of smell) or burned tongue (loss of
taste), food takes on a distinctly different flavor.
This integration is part of the way the brain is wired because it is part of the ritual of eating. When you place
food or drink in your mouth, it must go past your nose, which allows the brain to interpret an olfactory sensation,
before it begins to process gustatory sensations. Furthermore, even with a plugged nose, the back of your
throat is connected to both the nose and mouth which allows for what is called retronasal olfaction—odorants
can travel backward into the nose to create olfactory sensations. Furthermore, your sense of taste also relies on
your hearing (hearing the food), vision (seeing the food), and somatosensation (touching the food) when
determining gustatory perceptions and is particularly weak at making accurate perceptions based on the
sensations from the tongue alone.

 TRY IT

Test your taste discrimination skills the next time you eat.

1. Plug your nose.

2. Place a piece of food (or drink) in your mouth and chew or swish.
3. Try to mentally describe the flavor to yourself.
4. Unplug your nose.
5. Try to mentally describe the flavor to yourself.

Do you notice any difference between the flavor you perceived with the gustatory sense alone versus with
both gustatory and olfactory senses activated?

 TERMS TO KNOW

Gustation
The sense of taste.

Gustatory Papilla
A raised projection on the surface of the tongue that contains taste buds.

Taste Bud
Structures within a gustatory papilla that contain gustatory receptor cells.

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 Gustatory Receptor Cell
Sensory cells able to detect taste stimuli.

Taste Hair
Microvilli of the gustatory receptor cell.

Taste Pore
A small opening in the surface of the tongue epithelium.

Umami
A savory taste based on the presence of an amino acid.

 SUMMARY

In this lesson, you learned about the special senses of olfaction and gustation. You first learned the
anatomy and physiology of olfaction, the sense of smell. Then, learned the gustatory structures and
range of tastes that play a part in gustation, the sense of taste. Lastly, you explored how gustatory and
olfactory integration combines to create the perception of a taste.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

REFERENCES

U.S. Food & Drug Administration. (n.d.) Additional Information about High-Intensity Sweeteners Permitted for
Use in Food in the United States. Retrieved on January 18, 2023 from www.fda.gov/food/food-additives-
petitions/additional-information-about-high-intensity-sweeteners-permitted-use-food-united-states

 TERMS TO KNOW

Gustation
The sense of taste.

Gustatory Papilla
A raised projection on the surface of the tongue that contains taste buds.

Gustatory Receptor Cell

Sensory cells able to detect taste stimuli.

Odorant
Chemical molecules able to bind and activate olfactory sensory neurons.

Olfaction

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 The sense of smell.

Olfactory Bulb
An enlargement of the olfactory nerve.

Olfactory Epithelium
The tissue covering the nasal cavity.

Olfactory Sensory Neuron

Afferent sensory neurons in the nose that are sensitive to odorants; part of the olfactory nerve.

Olfactory Tract
The portion of the olfactory nerve that extends from the olfactory bulb to various regions of the brain.

Taste Bud
Structures within a gustatory papilla that contain gustatory receptor cells.

Taste Hair
Microvilli of the gustatory receptor cell.

Taste Pore
A small opening in the surface of the tongue epithelium.

Umami
A savory taste based on the presence of an amino acid.

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 Special Senses: Hearing and Equilibrium
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the special senses of hearing and equilibrium. Specifically, this
lesson will cover:
1. Audition (Hearing)
2. Vestibulation (Equilibrium, Balance)

1. Audition (Hearing)
Audition is the special sense of hearing that makes use of the ear to convert sound waves into neural signals.
The large, fleshy structure on the lateral aspect of the head is known as the auricle. The C-shaped curves of the
auricle direct sound waves toward the auditory canal. The canal enters the skull through the external auditory
meatus of the temporal bone. At the end of the auditory canal is the tympanic membrane, or ear drum, which
vibrates after it is struck by sound waves. The auricle, ear canal, and tympanic membrane are often referred to
as the external ear. The middle ear consists of a space spanned by three small bones called the ossicles. The
three ossicles are the malleus, incus, and stapes, which are Latin names that roughly translate to hammer, anvil,
and stirrup. The malleus is attached to the tympanic membrane and articulates with the incus. The incus, in turn,
articulates with the stapes. The stapes is then attached to the inner ear, where the sound waves will be
transduced into a neural signal. The middle ear is connected to the pharynx through the auditory tube, also
known as the Eustachian tube or pharyngotympanic tube, which helps equilibrate air pressure across the
tympanic membrane. The tube is normally closed but will pop open when the muscles of the pharynx contract
during swallowing or yawning.

 REFLECT

When you travel up or down in elevation, you might notice your ears can be irritated. This irritation is
caused by the change in atmospheric pressure.

Under normal conditions, the pressure on both sides of the tympanic membrane (ear drum) are equal,
allowing sound waves (i.e., small shifts in pressure) to cause the ear drum to move when they strike it.
However, when atmospheric pressure changes by changing elevation or your position in water (sinking to
the bottom of a pool or rising up to the surface while scuba diving), the pressure is no longer equal. This
can cause irritation and possible damage.

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 Swallowing or yawning can manipulate the auditory tube connecting the middle ear to the mouth, allowing
for air to enter or escape this cavity, and equalizing the pressure across the ear drum.

Structures of the Ear - The external ear contains the auricle, ear canal, and tympanic membrane. The middle ear

contains the ossicles and is connected to the pharynx by the Eustachian tube. The inner ear contains the cochlea and

vestibule, which are responsible for audition and equilibrium, respectively.

The inner ear is often described as a bony labyrinth, as it is composed of a series of bony canals embedded
within the temporal bone. It has two separate regions, the cochlea and the vestibule, which are responsible for
hearing and balance, respectively.

You can see that the cochlea looks like a snail's shell, a spiral moving inwards. On the lateral portion of the
cochlea is the oval window, an oval-shaped hole in the bone where the stapes sits. Inside the bony labyrinth of
the inner ear is a fluid-filled tube that spirals along with the bone to the center. Just inferior to the oval window
is the round window, a round-shaped hole in the cochlea covered by a membrane. As sound waves strike the
tympanic membrane and move it inwards, the ossicle bones shift. When the stapes moves, it presses into the
oval window causing a fluid wave within the cochlea. The round window will bulge out to accommodate the
wave and pucker in when the wave recedes. The frequency of the fluid waves will match the frequency of the
sound waves.

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 Transmission of Sound Waves to Cochlea - A sound wave causes the tympanic membrane to vibrate. This vibration is
amplified as it moves across the malleus, incus, and stapes. The amplified vibration is picked up by the oval window

causing pressure waves in the fluid of the scala vestibuli and scala tympani. The complexity of the pressure waves is

determined by the changes in amplitude and frequency of the sound waves entering the ear.

Along the center of the winding cochlea is a cavity called the cochlear duct, which contains auditory sensory
organs called organs of Corti. As the fluid waves move through the cochlea, the cochlear duct moves at a
specific spot, depending on the frequency of the waves. Higher frequency waves move the region of the duct
that is close to the base of the cochlea. Lower frequency waves move the region that is near the tip of the
cochlea.

The organs of Corti contain sensory receptor cells called hair cells, which are named for the hair-like stereocilia
extending from the cell’s apical surfaces. The stereocilia are an array of microvilli-like structures arranged from
tallest to shortest. Protein fibers tether adjacent hairs together within each array, such that the array will bend in
response to movements of the underlying tissue. The stereocilia extend up from the hair cells to the overlying
tectorial membrane, which is attached medially to the organ of Corti. When the pressure waves move the organ
of Corti, the tectorial membrane slides across the stereocilia. This bends the stereocilia either toward or away
from the tallest member of each array. When the stereocilia bend toward the tallest member of their array,

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 tension in the protein tethers opens ion channels in the hair cell membrane. This will depolarize the hair cell
membrane. When the stereocilia bend toward the shortest member of their array, the tension on the tethers
slackens and the ion channels close. When no sound is present and the stereocilia are standing straight, a small
amount of tension still exists on the tethers, keeping the membrane potential of the hair cell slightly
depolarized. Depolarized hair cells send signals to the auditory sensory neuron. All auditory sensory neurons
leave the cochlea as part of the cochlear nerve, which is the auditory portion of the vestibulocochlear nerve
(CN VIII).

Hair Cell - The hair cell is a mechanoreceptor with an array of stereocilia emerging from its apical surface. The

stereocilia are tethered together by proteins that open ion channels when the array is bent toward the tallest member

of their array and closed when the array is bent toward the shortest member of their array.

As stated above, a given region of the cochlear duct will only move if the incoming sound is at a specific
frequency. Because the tectorial membrane only moves where the cochlear duct moves, the hair cells in this
region will also only respond to sounds of this specific frequency. Therefore, as the frequency of a sound
changes, different hair cells are activated all along the basilar membrane. The cochlea encodes auditory stimuli
for frequencies between 20 and 20,000 Hz, which is the range of sound that human ears can detect. The unit
of Hertz measures the frequency of sound waves in terms of cycles produced per second. Frequencies as low
as 20 Hz are detected by hair cells at the apex, or tip, of the cochlea. Frequencies in the higher ranges of 20
KHz are encoded by hair cells at the base of the cochlea, close to the round and oval windows. Most auditory
stimuli contain a mixture of sounds at a variety of frequencies and intensities (represented by the amplitude of
the sound wave). The hair cells along the length of the cochlear duct, which are each sensitive to a particular
frequency, allow the cochlea to separate auditory stimuli by frequency just as a prism separates visible light into
its component colors.

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 Frequency Coding in the Cochlea - The standing sound wave generated in the cochlea by the movement of the oval

window deflects the basilar membrane on the basis of the frequency of sound. Therefore, hair cells at the base of the
cochlea are activated only by high frequencies; whereas, those at the apex of the cochlea are activated only by low

frequencies.

 TERMS TO KNOW

Audition
The special sense of hearing.

Auricle
The fleshy portion of the external ear.

Tympanic Membrane
The membrane at the end of the external acoustic meatus; eardrum.

External Ear
A portion of the auditory anatomy including the auricle, ear canal, and tympanic membrane.

Middle Ear
A portion of the auditory anatomy between the tympanic membrane and bony labyrinth, including the
ossicles.

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 Ossicles
Three small bones located in the middle ear.

Malleus
The ossicle attached to the tympanic membrane; the hammer.

Incus
The ossicle that connects the malleus and stapes; the anvil.

Stapes
The ossicle that connects to the bony labyrinth of the inner ear; the stirrup.

Inner Ear
The portion of the auditory and vestibular anatomy including the cochlea and vestibule.

Cochlea
The auditory portion of the inner ear.

Vestibule
The vestibular portion of the inner ear.

Oval Window
A membrane-covered hole in the cochlea where the stapes is attached.

Round Window
A membrane-covered hole in the cochlea inferior to the oval window.

Cochlear Duct
A cavity within the cochlea that contains the organs of Corti.

Organ of Corti
A structure within the cochlear duct that contains hair cells and can send auditory sensory signals.

Hair Cell
Sensory receptor cells of the inner ear that detect changes in the position of their stereocilia.

Stereocilia
Microvilli-like extension of the hair cell membrane that attaches to the tectorial membrane.

Tectorial Membrane
A structure of the organ of Corti that stereocilia are connected to.

Auditory Sensory Neuron

Afferent sensory neurons in the ear that are sensitive to hair cell signals; part of the cochlear nerve.

Cochlear Nerve
The auditory branch of the vestibulocochlear nerve.

2. Vestibulation (Equilibrium, Balance)

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 Along with audition, the inner ear is responsible for encoding information about vestibulation, also known as
the sense of equilibrium or balance. Recall that the inner ear is a bony labyrinth composed of the cochlea and
vestibule. The cochlea contains auditory structures while the vestibule contains vestibular structures. Within the
vestibule are three semicircular canals that detect rotational movement as well as the utricle and saccule which
detect linear movement and position.

The utricle and saccule both contain hair cells embedded in them. The stereocilia of the hair cells extend into a
viscous gel called the otolithic membrane. On top of the otolithic membrane is a layer of calcium carbonate
crystals called otoliths, also known as ear stones. The otoliths essentially make the otolithic membrane top-
heavy. The otolithic membrane moves separately from the underlying tissue in response to head movements.
Tilting the head causes the otolithic membrane to slide in the direction of gravity. The moving otolithic
membrane, in turn, bends the stereocilia, causing some hair cells to depolarize as others hyperpolarize. The
exact position of the head is interpreted by the brain based on the pattern of hair-cell depolarization. The utricle
and saccule can also detect changes in linear movement. The utricle is positioned in the horizontal plane and
can detect horizontal acceleration or deceleration such as your car picking up speed or slowing down. The
saccule is positioned vertically and can detect vertical acceleration or deceleration such as an elevator.

Linear Acceleration Coding - The utricle and saccule are specialized for sensing linear acceleration, such as when

gravity acts on the tilting head or if the head starts moving in a straight line. The difference in inertia between the hair

cell stereocilia and the otolithic membrane in which they are embedded leads to a shearing force that causes the

stereocilia to bend in the direction of that linear acceleration.

The semicircular canals are three ring-like extensions of the vestibule. One is oriented in the horizontal plane,
whereas the other two are oriented in the vertical plane. The anterior and posterior vertical canals are oriented
at approximately 45 degrees relative to the sagittal plane. The base of each semicircular canal, where it meets

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 with the vestibule, connects to an enlarged region known as the ampulla of the inner ear which contains hair
cells. These hair cells have stereocilia embedded into the cupula, a membrane that attaches to the top of the
ampulla.

When the head moves in a rotational movement (i.e., spinning in circles, doing a somersault, or doing a
cartwheel), fluid inside one or more of the semicircular canals shifts. As this fluid moves along the ampulla, it
bends the cupula and in turn the stereocilia. This depolarizes the hair cells and sends a signal to a connected
ampullary nerve which extends as the vestibular nerve. This nerve combines with the cochlear nerve to
become the vestibulocochlear nerve. By comparing the signals from all ampullae, the vestibular system can
detect the direction of most head movements within three-dimensional (3-D) space.

Rotational Coding by Semicircular Canals - Rotational movement of the head is encoded by the hair cells in the base

of the semicircular canals. As one of the canals moves in an arc with the head, the internal fluid moves in the opposite
direction, causing the cupula and stereocilia to bend. The movement of two canals within a plane results in

information about the direction in which the head is moving, and activation of all six canals can give a very precise

indication of head movement in three dimensions.

 TERMS TO KNOW

Vestibulation
The sense of equilibrium or balance.

Utricle
A structure of the inner ear able to detect head position and horizontal acceleration and deceleration.

Saccule
A structure of the inner ear able to detect head position and vertical acceleration and deceleration.

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 Otolithic Membrane
A viscous gelatinous substance in the utricle and saccule of the inner ear that stereocilia of hair cells
are embedded in.

Otolith
Calcium carbonate crystals located on top of the otolithic membrane.

Semicircular Canal
Structures of the inner ear that detect rotational movement of the head.

Ampulla of the Inner Ear

The enlarged region at the base of a semicircular canal containing hair cells.

Cupula
A membrane located in the ampulla of the semicircular canals where the stereocilia of hair cells are
embedded.

Vestibular Nerve
The vestibular branch of the vestibulocochlear nerve.

 SUMMARY

In this lesson, you learned about the special senses of audition and vestibulation. You learned the
anatomy of the ear and how it physiologically detects sound waves to produce audition and body
position as well as body movement to detect vestibulation.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Ampulla of the Inner Ear

The enlarged region at the base of a semicircular canal containing hair cells.

Audition
The special sense of hearing.

Auditory Sensory Neuron

Afferent sensory neurons in the ear that are sensitive to hair cell signals; part of the cochlear nerve.

Auricle
The fleshy portion of the external ear.

Cochlea
The auditory portion of the inner ear.

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 Cochlear Duct
A cavity within the cochlea that contains the organs of Corti.

Cochlear Nerve
The auditory branch of the vestibulocochlear nerve.

Cupula
A membrane located in the ampulla of the semicircular canals where the stereocilia of hair cells are
embedded.

External Ear
A portion of the auditory anatomy including the auricle, ear canal, and tympanic membrane.

Hair Cell
Sensory receptor cells of the inner ear that detect changes in the position of their stereocilia.

Incus
The ossicle that connects the malleus and stapes; the anvil.

Inner Ear
The portion of the auditory and vestibular anatomy including the cochlea and vestibule.

Malleus
The ossicle attached to the tympanic membrane; the hammer.

Middle Ear
A portion of the auditory anatomy between the tympanic membrane and bony labyrinth, including the
ossicles.

Organ of Corti
A structure within the cochlear duct that contains hair cells and can send auditory sensory signals.

Ossicles
Three small bones located in the middle ear.

Otolith
Calcium carbonate crystals located on top of the otolithic membrane.

Otolithic Membrane
A viscous gelatinous substance in the utricle and saccule of the inner ear that stereocilia of hair cells are
embedded in.

Oval Window
A membrane-covered hole in the cochlea where the stapes is attached.

Round Window
A membrane-covered hole in the cochlea inferior to the oval window.

Saccule

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 A structure of the inner ear able to detect head position and vertical acceleration and deceleration.

Semicircular Canal
Structures of the inner ear that detect rotational movement of the head.

Stapes
The ossicle that connects to the bony labyrinth of the inner ear; the stirrup.

Stereocilia
Microvilli-like extension of the hair cell membrane that attaches to the tectorial membrane.

Tectorial Membrane
A structure of the organ of Corti that stereocilia are connected to.

Tympanic Membrane
The membrane at the end of the external acoustic meatus; ear drum.

Utricle
A structure of the inner ear able to detect head position and horizontal acceleration and deceleration.

Vestibular Nerve
The vestibular branch of the vestibulocochlear nerve.

Vestibulation
The sense of equilibrium or balance.

Vestibule
The vestibular portion of the inner ear.

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 Special Senses: Vision
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the special sense of vision. Specifically, this lesson will cover:
1. Vision

1. Vision
Vision is the special sense of sight that uses the eye to detect light waves and convert them into neural signals.
The eyes are located within either orbit in the skull. The bony orbits surround the eyeballs, protecting them and
anchoring the soft tissues of the eye. The eyelids, with lashes at their leading edges, help to protect the eye
from abrasions by blocking particles that may land on the surface of the eye. Tears are produced by the lacrimal
gland, located just inside the orbit, superior and lateral to the eyeball. Tears exit the lacrimal gland and flow
over the surface of the eye, washing away foreign particles, exiting the eye through the nasolacrimal duct at
the medial corner of the eye. This tube connects and drains tears into the nasal cavity.

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 The Eye in the Orbit - The eye is located within the orbit and surrounded by soft tissues that protect and support its

function. The orbit is surrounded by cranial bones of the skull

Movement of the eye within the orbit is accomplished by the contraction of six extraocular muscles that
originate from the bones of the orbit and insert into the surface of the eyeball. For more detailed information,
please refer to the supplemental  Axial Muscles.pdf. These muscles are innervated by three cranial nerves,
the oculomotor (CN III), trochlear (CN IV), and abducens nerves (CN VI).

The eye itself is a hollow sphere composed of three layers of tissue called tunics (tunic, coat)—the fibrous tunic,
vascular tunic, and neural tunic. The outermost layer is the fibrous tunic, which includes the white sclera and
clear cornea. The sclera accounts for five-sixths of the surface of the eye, most of which is not visible, though
humans are unique compared with many other species in having so much of the “white of the eye” visible. The
transparent cornea acts like a curved window, covering the anterior tip of the eye and allowing light to enter the
eye.

The middle layer of the eye is the vascular tunic, which is mostly composed of the choroid, ciliary body, and iris.
The choroid is a layer of highly vascularized connective tissue that provides a blood supply to the eyeball. The
choroid is posterior to the ciliary body, a muscular structure that is attached to the lens by suspensory
ligaments, or zonule fibers. These two structures bend the lens, allowing it to focus light on the back of the eye.
Overlaying the ciliary body, and visible in the anterior eye, is the iris—the colored part of the eye. The iris is a
smooth muscle that opens or closes the pupil, which is the hole at the center of the eye that allows light to

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 enter. The iris constricts the pupil in response to bright light and dilates the pupil in response to dim light. The
innermost layer of the eye is the retina, or neural tunic, which contains the nervous tissue responsible for
photoreception.

The eye is also divided into two cavities:

The anterior cavity

The posterior cavity

The anterior cavity is the space between the cornea and lens, including the iris and ciliary body. It is filled with a
watery fluid called aqueous humor.

The posterior cavity is the space behind the lens that extends to the posterior side of the interior eyeball,
where the retina is located. The posterior cavity is filled with a more viscous fluid called the vitreous humor.

In the posterior cavity, the retina is composed of several layers and contains specialized cells for the initial
processing of visual stimuli. The photoreceptors are specialized receptor cells that change their membrane
potential when stimulated by light energy. The change in membrane potential alters the amount of
neurotransmitter that the photoreceptor cells release onto bipolar cells in the outer synaptic layer. It is the
bipolar cell in the retina that connects a photoreceptor to a retinal ganglion cell (RGC) in the inner synaptic
layer. The axons of RGCs, which lie at the innermost layer of the retina, all converge at one point in the eye and
exit the eye collectively as the optic nerve (CN II). The point at which they exit the eye is called the optic disc,
also known as the blind spot. Because these axons pass through the retina, there are no photoreceptors in that
space. This creates a “blind spot” in the retina and a corresponding blind spot in our visual field.

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 Structure of the Eye - The sphere of the eye can be divided into anterior and posterior chambers. The wall of the eye is

composed of three layers: the fibrous tunic, vascular tunic, and neural tunic. Within the neural tunic is the retina, with

three layers of cells and two synaptic layers in between. The center of the retina has a small indentation known as the

fovea.

Note that the photoreceptors in the retina are located behind the axons, RGCs, bipolar cells, and retinal blood
vessels. A significant amount of light is absorbed by these structures before the light reaches the photoreceptor
cells. However, at the exact center of the retina is a small area known as the fovea centralis, or fovea for short,
that lacks the supporting cells and blood vessels and only contains photoreceptors. Because this is where the
least amount of light is absorbed by other retinal structures, it is where visual acuity, or the sharpness of vision,
is greatest. As one moves in either direction from this central point of the retina, visual acuity drops significantly.
In addition, each photoreceptor cell of the fovea is connected to a single RGC. Therefore, this RGC does not
have to integrate inputs from multiple photoreceptors, which reduces the accuracy of visual transduction.
Toward the edges of the retina, several photoreceptors converge on RGCs (through the bipolar cells) up to a
ratio of 50 to 1.

 WATCH

Please watch the following video for more information on this topic.

 TRY IT

The difference in visual acuity between the fovea and peripheral retina is easily evidenced by looking
directly at a word in the middle of this paragraph. The visual stimulus in the middle of the field of view falls

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 on the fovea and is in the sharpest focus. Without moving your eyes off that word, notice that words at the
beginning or end of the paragraph are not in focus. The images in your peripheral vision are focused by the
peripheral retina and have vague, blurry edges and words that are not as clearly identified. As a result, a
large part of the neural function of the eyes is concerned with moving the eyes and head so that important
visual stimuli are centered on the fovea.
Light falling on the retina causes chemical changes to pigment molecules in the photoreceptors, ultimately
leading to a change in the activity of the RGCs. Photoreceptor cells have two parts, the inner segment and the
outer segment. The inner segment contains the nucleus and other common organelles of a cell, whereas the
outer segment is a specialized region in which photoreception takes place. There are two types of
photoreceptors—rods and cones—which differ in their shape and light sensitivity. Rods have a rod-shaped outer
segment and contain the photosensitive pigment rhodopsin. These cells are sensitive to light intensity but do
not detect colors. Rod photoreceptors only generate black-and-white images. Cones have a cone-shaped outer
segment and contain one of three photosensitive opsin pigments, each sensitive to a particular wavelength
(color) of light. The pigments in human eyes are specialized in perceiving three different primary colors: red,
green, and blue.

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© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 156
 Photoreceptor - (a) All photoreceptors have inner segments containing the nucleus and other important organelles

and outer segments with membrane arrays containing the photosensitive opsin molecules. Rod outer segments are

long columnar shapes with stacks of membrane-bound discs that contain the rhodopsin pigment. Cone outer
segments are short, tapered shapes with folds of the membrane in place of the discs in the rods. (b) Tissue of the

retina shows a dense layer of nuclei of the rods and cones. LM × 800.

 WATCH

Please watch the following video for more information on this topic.

At the molecular level, visual stimuli cause changes in the photopigment molecule that lead to changes in
membrane potential of the photoreceptor cell. A single unit of light is called a photon. The energy of a photon is
represented by its wavelength, with each wavelength of visible light corresponding to a particular color. Visible
light has a wavelength between 380 and 720 nm. Light with a wavelength of 380 nm is blue whereas light with
a wavelength of 720 nm is dark red. All other colors fall between red and blue at various points along the
wavelength scale.

Wavelengths of Visible Light - Visible light spans wavelengths from 380 to 780 nm along the electromagnetic

spectrum.

Opsin pigments are transmembrane proteins that contain a cofactor known as retinal. Retinal is a molecule
related to vitamin A. When light hits retinal, it changes shape in a process known as photoisomerization. The
shape change of retinal in the photoreceptor membrane initiates a change in its membrane potential. This
causes the photoreceptor cell to decrease neurotransmitter release. This means that photoreceptor cells and
the optic nerve decrease activity in the presence of light stimuli and increase in the absence of light stimuli.

Additionally, until the retinal molecule is changed back to its original shape by another enzymatic reaction, the
opsin cannot respond to light energy, which is called bleaching. When a large group of photopigments is
bleached, the retina will send information as if opposing (negative) visual information is being perceived.

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 EXAMPLE If you look at a bright light and then look away, your vision will have an afterimage of a dark
spot, also known as flash blindness.
The same general process also works for colors. If you look at a relatively light red image and then look away,
your vision will see the same afterimage but in green (the opposite color to red).

All visual pigments are sensitive to limited wavelengths of light. Rhodopsin, the photopigment in rods, is most
sensitive to light at a wavelength of 498 nm. The three color opsins have peak sensitivities of 564 nm, 534 nm,
and 420 nm corresponding roughly to the primary colors of red, green, and blue. The absorbance of rhodopsin
in the rods is much more sensitive than in the cone opsins; specifically, rods are sensitive to vision in low light
conditions, and cones are sensitive to brighter conditions. In normal sunlight, rhodopsin will be constantly
bleached while the cones are active. In a darkened room, there is not enough light to activate cone opsins, and
vision is entirely dependent on rods. Rods are so sensitive to light that a single photon can result in an action
potential from a rod’s corresponding RGC.

The three types of cone opsins, being sensitive to different wavelengths of light, provide us with color vision. By
comparing the activity of the three different cones, the brain can extract color information from visual stimuli.

EXAMPLE A bright blue light that has a wavelength of approximately 450 nm would activate the “red”
cones minimally, the “green” cones marginally, and the “blue” cones predominantly.
The relative activation of the three different cones is calculated by the brain, which perceives the color as blue.
However, cones cannot react to low-intensity light, and rods do not sense the color of light. Therefore, our low-
light vision is—in essence—in grayscale. In other words, in a dark room, everything appears as a shade of gray.
If you think that you can see colors in the dark, it is most likely because your brain knows what color something
is and is relying on that memory.

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 Comparison of Color Sensitivity of Photopigments - Comparing the peak sensitivity and absorbance spectra of the

four photopigments suggests that they are most sensitive to particular wavelengths.

 TERMS TO KNOW

Vision
The special sense of sight that uses the eye to detect light waves and convert them into neural signals.

Lacrimal Gland
A gland superior to the lateral angle of the eye that produces and secretes tears.

Nasolacrimal Duct
A tube that connects the medial angle of the eye to the nasal cavity to drain tears.

Fibrous Tunic
The outer layer of the eye containing the sclera and cornea.

Sclera
The white of the eye.

Cornea
The transparent covering of the anterior tip of the eye that allows light to enter.

Vascular Tunic

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 The middle layer of the eye containing the choroid, ciliary body, and iris.

Choroid
The vascular tissue of the eye.

Ciliary Body
A smooth muscle structure on the interior of the iris that controls the shape of the lens.

Lens
A component of the eye that focuses light on the retina.

Zonule Fiber
Fibrous connection between the ciliary body and lens.

Iris
The colored ring of the eye.

Pupil
The black hole at the center of the iris that allows light to pass into the posterior chamber.

Retina (aka Neural Tunic)

The inner layer of the eye containing sensory receptors and neurons, the retina.

Anterior Cavity
The space between the cornea and lens.

Aqueous Humor
The fluid located in the anterior cavity.

Posterior Cavity
The space behind the lens.

Vitreous Humor
The viscous fluid located in the posterior cavity.

Photoreceptor
Specialized receptor cell in the neural layer of the eye that is sensitive to light stimuli.

Bipolar Cell
A cell that connects the photoreceptor to the retinal ganglion cell.

Retinal Ganglion Cell

A neuron of the retina which transports a signal along the optic nerve.

Optic Disc
The point at which all retinal ganglion cell axons exit the eye as the optic nerve; blind spot.

Fovea Centralis
The central point in the eye with the greatest visual acuity.

Rod
Photoreceptor that is specialized for low-light vision.

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 Rhodopsin
A photosensitive pigment found in rod photoreceptors.

Cone
One of three photoreceptors that is specialized for color vision.

Photon
A single unit of light.

Photoisomerization
A change in the chemical structure of a photosensitive pigment.

Bleaching
A process by which an opposite visual after-image is perceived after exposure to a bright light source.

 SUMMARY

In this lesson, you learned about the special sense of vision. You learned the anatomy of the eye and
how the light performs photoreception.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Anterior Cavity
The space between the cornea and lens.

Aqueous Humor
The fluid located in the anterior cavity.

Bipolar Cell
A cell that connects the photoreceptor to the retinal ganglion cell.

Bleaching
A process by which an opposite visual after-image is perceived after exposure to a bright light source.

Choroid
The vascular tissue of the eye.

Ciliary Body
A smooth muscle structure on the interior of the iris that controls the shape of the lens.

Cone Photoreceptor
One of three photoreceptors that is specialized for color vision.

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 Cornea
The transparent covering of the anterior tip of the eye that allows light to enter.

Fibrous Tunic
The outer layer of the eye containing the sclera and cornea.

Fovea Centralis
The central point in the eye with the greatest visual acuity.

Iris
The colored ring of the eye.

Lacrimal Gland
A gland superior to the lateral angle of the eye that produces and secretes tears.

Lens
A component of the eye that focuses light on the retina.

Nasolacrimal Duct
A tube that connects the medial angle of the eye to the nasal cavity to drain tears.

Neural Tunic (aka Retina)

The inner layer of the eye containing sensory receptors and neurons, the retina.

Optic Disc
The point at which all retinal ganglion cell axons exit the eye as the optic nerve; blind spot.

Photoisomerization
A change in the chemical structure of a photosensitive pigment.

Photon
A single unit of light.

Photoreceptor
Specialized receptor cell in the neural layer of the eye that is sensitive to light stimuli.

Posterior Cavity
The space behind the lens.

Pupil
The black hole at the center of the iris that allows light to pass into the posterior chamber.

Retinal Ganglion Cell

A neuron of the retina which transports a signal along the optic nerve.

Rhodopsin
A photosensitive pigment found in rod photoreceptors.

Rod Photoreceptor

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 Photoreceptor that is specialized for low-light vision.

Sclera
The white of the eye.

Vascular Tunic
The middle layer of the eye containing the choroid, ciliary body, and iris.

Vision
The special sense of sight that uses the eye to detect light waves and convert them into neural signals.

Vitreous Humor
The viscous fluid located in the posterior cavity.

Zonule Fiber
Fibrous connection between the ciliary body and lens.

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 Central Processing: Spinal Cord and Brainstem
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about how sensory signals are transported to the brain. Specifically, this
lesson will cover:
1. Sensory Pathways
1a. Spinal Cord and Brainstem
1b. Diencephalon

1. Sensory Pathways
Specific regions of the central nervous system (CNS) coordinate different somatic processes using sensory
inputs and motor outputs of peripheral nerves. A simple case is a spinal reflex caused by a single synapse
transferring information between a dorsal sensory neuron and a motor neuron in the anterior horn. More
complex arrangements with multiple synapses are possible to integrate peripheral sensory information with
higher processes. The important regions of the CNS that play a role in somatic processes are the spinal cord
brainstem, diencephalon, cerebral cortex, and subcortical structures.

1a. Spinal Cord and Brainstem

A sensory pathway that carries peripheral sensations to the brain is referred to as an ascending pathway, or
ascending tract. The various sensory modalities each follow specific pathways through the CNS. Tactile and
other somatosensory stimuli activate receptors in the skin, muscles, tendons, and joints throughout the entire
body. Somatosensory stimuli from the head and neck travel through the cranial nerves, (specifically, the
trigeminal system which is not covered in this course) whereas somatosensory stimuli from below the neck pass
along the sensory pathways of the spinal cord. Two major pathways that transport sensory information through
the spinal cord to the brain are the posterior column system and spinothalamic tract.

The posterior column system transports sensations of light touch, pressure, vibration, and proprioception from
various parts of the body below the neck to the brain via the spinal cord. To do this, information is passed along
three neurons. The first, referred to as the first-order neuron, is the sensory neuron that enters the posterior
horn of the spinal cord via the posterior root. This neuron enters the posterior column and ascends to the
medulla in the brainstem, where it synapses with the second-order neuron. The second-order neuron crosses
over the medulla (it switches from left to right side or right to left side) and ascends to the thalamus in the
diencephalon. Here, it synapses with the third-order neuron which will project to the postcentral gyrus of the

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 cerebral cortex. Recall that this is the primary somatosensory cortex where somatosensory signals are
processed to generate perceptions (conscious awareness) of the stimuli.

The spinothalamic tract transports crude touch and pressure as well as pain and temperature signals from
below the neck to the brain via the spinal cord. This pathway also includes three neurons. The first-order
neuron, much like the posterior column, begins by entering the posterior horn of the spinal cord via the
posterior root. However, it synapses with the second-order neuron there. The second-order neuron then
crosses over the spinal cord, enters the anterior or lateral column, and ascends to the thalamus to synapse with
the third-order neuron. As with the posterior column pathway, the third-order neuron will project to the
appropriate region within the somatosensory cortex to allow for processing of the signal.

Table: Ascending Pathways

Neuron Attribute Posterior Column Pathway Spinothalamic Tract

Cell Body Posterior Root Ganglion Posterior Root Ganglion

Receives signal at… Sensory receptor Sensory receptor

First-Order Neuron
Projects axon to… Medulla Posterior horn of spinal cord

Crosses Over? No No

Cell Body Medulla Posterior horn of spinal cord

Receives signal at… Medulla Posterior horn of spinal cord

Second-Order Neuron
Projects axon to… Thalamus (opposite side) Thalamus (opposite side)

Crosses Over? Yes Yes

Cell Body Thalamus Thalamus

Receives signal at… Thalamus Thalamus

Third-Order Neuron
Projects axon to… Somatosensory cortex Somatosensory cortex

Crosses Over? No No

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 Anterior View of Ascending Sensory Pathways of the Spinal Cord - The dorsal column system and spinothalamic tract

are the major ascending pathways that connect the periphery with the brain.

Sound localization is a feature of central processing in the brainstem. Sound localization is achieved by the
brain calculating the interaural time difference and the interaural intensity difference. A sound originating from
a specific location will arrive at each ear at different times unless the sound is directly in front of the listener. If
the sound source is slightly to the left of the listener, the sound will arrive at the left ear microseconds before it
arrives at the right ear. This time difference is an example of an interaural time difference. Also, the sound will

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 be slightly louder in the left ear than in the right ear because some of the sound waves reaching the opposite
ear are blocked by the head. This is an example of an interaural intensity difference.

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 Auditory Brainstem Mechanisms of Sound Localization - Localizing sound in the horizontal plane is achieved by

processing in the medullary nuclei of the auditory system. Connections between neurons on either side are able to
compare very slight differences in sound stimuli that arrive at either ear and represent interaural time and intensity

differences.

Auditory processing continues on to the inferior colliculus in the midbrain of the brainstem. Axons project from
the inferior colliculus to the thalamus and superior colliculus. From the thalamus, the information is sent to the
primary auditory cortex in the temporal lobe of the cerebrum. The superior colliculus uses auditory as well as
visual and somatosensory input to turn the head and neck in the direction of the auditory stimulus.

Balance is coordinated through the vestibular system, the nerves of which are composed of axons from the
vestibular nerve that carries information from the utricle, saccule, and semicircular canals. The system
contributes to controlling head and neck movements in response to vestibular signals. An important function of

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 the vestibular system is coordinating eye and head movements to maintain visual attention. Most of the axons
terminate in the medulla. Some axons from the vestibular nerve project directly to the cerebellum, with no
intervening synapse. The cerebellum is primarily responsible for initiating movements on the basis of
equilibrium information.

The connections of the optic nerve are more complicated than those of other cranial nerves. Instead of the
connections being between each eye and the brain, visual information is segregated between the left and right
sides of the visual field. In addition, some of the information from one side of the visual field projects to the
opposite side of the brain. Within each eye, the axons projecting from the medial side of the retina cross over at
the optic chiasm. For example, the axons from the medial retina of the left eye cross over to the right side of the
brain at the optic chiasm. However, within each eye, the axons projecting from the lateral side of the retina do
not decussate. For example, the axons from the lateral retina of the right eye project back to the right side of
the brain. Therefore the left field of view of each eye is processed on the right side of the brain, whereas the
right field of view of each eye is processed on the left side of the brain. Extending from the optic chiasm, the
axons of the visual system are referred to as the optic tract instead of the optic nerve. The majority of the
connections of the optic tract are to the thalamus—specifically, the lateral geniculate nucleus. Axons from this
nucleus then project to the visual cortex of the cerebrum, located in the occipital lobe. Another target of the
optic tract is the superior colliculus which can move the head and neck in the direction of a visual stimuli.

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 Segregation of Visual Field Information at the Optic Chiasm - Opposite side visual field information from the lateral

retina projects to the same side of the brain, whereas the same side visual field information has to cross over at the

optic chiasm to reach the opposite side of the brain. (Note: This is an inferior view.)

In addition, a very small number of axons project from the optic chiasm to the suprachiasmatic nucleus of the
hypothalamus. These cells are photosensitive, in that they respond to the presence or absence of light. Unlike
the photoreceptors, however, these photosensitive cells cannot be used to perceive images. By simply
responding to the absence or presence of light, these cells can send information about day length. The

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 perceived proportion of sunlight to darkness establishes the circadian rhythm of our bodies, allowing certain
physiological events to occur at approximately the same time every day.

 DID YOU KNOW

Damage to any portion of the optic nerve or its projecting fibers leads to unique visual field impairments
which can indicate the location of the damage. In the image below, the red lines indicate sites of damage
along the visual pathway and the corresponding visual field changes. Notice that each visual pattern is
different. This can help to pinpoint the location of the damage.

Visual Field Damage - Damage to any of the axons along the visual pathway creates a unique pattern of visual

loss.

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 1b. Diencephalon
Recall that the diencephalon is deep in the cerebrum and includes the thalamus, hypothalamus, and
epithalamus. Sensory input to the thalamus comes from most of the special senses and ascending
somatosensory tracts. Each sensory system is relayed through a particular nucleus in the thalamus. The
thalamus is a required transfer point for most sensory tracts that reach the cerebral cortex, where conscious
sensory perception begins. The one exception to this rule is the olfactory system. The olfactory tract axons from
the olfactory bulb project directly to the cerebral cortex, along with the limbic system and hypothalamus.

 TERMS TO KNOW

Ascending Pathway
A sensory pathway that carries peripheral sensations to the brain.

Posterior Column System

An ascending tract of the spinal cord associated with fine touch and proprioception.

Spinothalamic Tract
An ascending tract of the spinal cord associated with crude touch and pressure as well as pain and
temperature.

Interaural Time Difference

A cue used to help sound localization that compares the arrival time of an auditory stimuli at each ear.

Interaural Intensity Difference

A cue used to help sound localization that compares the intensity (volume) of an auditory stimuli at each
ear.

Optic Chiasm
The crossover point of the visual pathway where select optic nerve fibers cross to the other side

Optic Tract
The group of axons that project posterior from the optic chiasm.

 SUMMARY

In this lesson, you learned to identify major sensory pathways, their location, and the types of
information they transport. You began by looking at several major systems that transport signals
through the spinal cord and brainstem. Then, you covered the overall function of the diencephalon as
a relay station for sensory pathways.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

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 Ascending Pathway
A sensory pathway that carries peripheral sensations to the brain.

Interaural Intensity Difference

A cue used to help sound localization that compares the intensity (volume) of an auditory stimuli at each
ear.

Interaural Time Difference

A cue used to help sound localization that compares the arrival time of an auditory stimuli at each ear.

Optic Chiasm
The crossover point of the visual pathway where select optic nerve fibers cross to the other side.

Optic Tract
The group of axons that project posterior from the optic chiasm.

Posterior Column System

An ascending tract of the spinal cord associated with fine touch and proprioception.

Spinothalamic Tract
An ascending tract of the spinal cord associated with crude touch and pressure as well as pain and
temperature.

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 Central Processing: Sensory Cortex
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about how sensory signals are processed in the brain to generate
perceptions. Specifically, this lesson will cover:
1. Cortical Processing

1. Cortical Processing
As described earlier, many of the sensory axons are positioned in the same way as their corresponding receptor
cells in the body. This allows identification of the position of a stimulus on the basis of which receptor cells are
sending information. The cerebral cortex also maintains this sensory topography in the particular areas of the
cortex that correspond to the position of the receptor cells in the body. The somatosensory cortex provides an
example in which, in essence, the locations of the somatosensory receptors in the body are mapped onto the
somatosensory cortex. This mapping is often depicted using a sensory homunculus shown in the image below.

The term homunculus comes from the Latin word for “little man” and refers to a map of the human body that is
laid across a portion of the cerebral cortex. In the somatosensory cortex, the external genitals, feet, and lower
legs are represented on the medial face of the gyrus within the longitudinal fissure. As the gyrus curves out of
the fissure and along the surface of the parietal lobe, the body map continues through the thighs, hips, trunk,
shoulders, arms, and hands. The head and face are just lateral to the fingers as the gyrus approaches the lateral
sulcus. The representation of the body in this topographical map is medial to lateral from the lower to the upper
body. Note that this correspondence does not result in a perfectly miniature scaled version of the body but
rather exaggerates the more sensitive areas of the body where more receptor cells exist, such as the fingers
and lower face. Less sensitive areas of the body where fewer receptor cells exist, such as the shoulders and
back, are mapped to smaller areas of the cortex.

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 The Sensory Homunculus - A cartoon representation of the sensory homunculus arranged on top of the

somatosensory cortex in which the processing takes place.

Likewise, the topographic relationship between the retina and the visual cortex is maintained throughout the
visual pathway. The visual field is projected onto the two retinae, as described above, with sorting at the optic
chiasm. The right peripheral visual field falls on the medial portion of the right retina and the lateral portion of
the left retina. The right medial retina then projects across the midline through the optic chiasm. This results in
the right visual field being processed in the left visual cortex. Likewise, the left visual field is processed in the
right visual cortex. Though the chiasm is helping to sort right and left visual information, superior and inferior
visual information is maintained topographically in the visual pathway. Light from the superior visual field falls on

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 the inferior retina, and light from the inferior visual field falls on the superior retina. This topography is
maintained such that the superior region of the visual cortex processes the inferior visual field and vice versa.
Therefore, the visual field information is inverted and reversed as it enters the visual cortex—up is down, and
left is right. However, the cortex processes the visual information such that the final conscious perception of the
visual field is correct. The topographic relationship is evident in that information from the foveal region of the
retina is processed in the center of the primary visual cortex. Similar to the exaggerations in the sensory
homunculus of the somatosensory cortex, the foveal-processing area of the visual cortex is disproportionately
larger than the areas processing peripheral vision.

 DID YOU KNOW

In an experiment performed in the 1960s, subjects wore prism glasses so that the visual field was inverted
before reaching the eye. On the first day of the experiment, subjects would duck when walking up to a
table, thinking it was suspended from the ceiling. However, after a few days of acclimation, the subjects
behaved as if everything were represented correctly. Therefore, the visual cortex is somewhat flexible in
adapting to the information it receives from our eyes.

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 Topographic Mapping of the Retina onto the Visual Cortex - The visual field projects onto the retina through the lenses

and falls on the retinae as an inverted, reversed image. The topography of this image is maintained as the visual

information travels through the visual pathway to the cortex.

The cortex has been described as having specific regions that are responsible for processing specific
information; there is the visual cortex, somatosensory cortex, gustatory cortex, and others. However, our
experience of these senses is not divided. Instead, we experience what can be referred to as a seamless
percept. Our perceptions of the various sensory modalities—though distinct in their content—are integrated by
the brain so that we experience the world as a continuous whole.

In the cerebral cortex, sensory processing begins at the primary sensory cortex, then proceeds to an
association area, and finally into a multimodal integration area.

EXAMPLE The visual pathway projects from the retinae through the thalamus to the primary visual
cortex in the occipital lobe. In this area, the brain is able to determine the light intensity (brightness) and
color (wavelength) of each incoming signal. This is like determining each individual pixel of a larger image.
From the visual cortex, this information is sent to the visual association area where the brain takes the
individual pieces of information and compares them to previous sets of data such as shapes, colors, and
similar attributes. Finally, this information is passed onto an integration area where information from other
cortices and association areas (such as somatosensory, auditory, or olfactory) are mixed and again
compared to previous sets of data. Here, you are able to recognize that what you’re looking at is a pizza
because it is shaped like pizzas you’ve seen before (visual information), smells like the combination of
dough and cheese (olfactory information), and you can hear the sounds the crust on the pizza box as
people grab a slice (auditory information).
Your brain uses many ways to process information, always checking it against previous experience and double-
checking with other senses.

IN CONTEXT
Everyday Connection - Depth Perception, 3-D Movies, and Optical Illusions
The visual field is projected onto the retinal surface, where photoreceptors transduce light energy into
neural signals for the brain to interpret. The retina is a two-dimensional surface, so it does not encode
three-dimensional information. However, we can perceive depth. How is that accomplished?

Two ways in which we can extract depth information from the two-dimensional retinal signal are based

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 on:

Monocular cues and binocular cues, respectively. Monocular depth cues are those that are the result
of information within the two-dimensional visual field. One object that overlaps another object has to
be in front. Relative size differences are also a cue.

EXAMPLE If a basketball appears larger than the basket, then the basket must be further away.

On the basis of experience, we can estimate how far away the basket is. Binocular depth cues
compare information represented in the two retinae because they do not see the visual field exactly
the same.

The centers of the two eyes are separated by a small distance, which is approximately 6 to 6.5 cm in
most people. Because of this offset, visual stimuli do not fall on exactly the same spot on both retinae
unless we are fixated directly on them and they fall on the fovea of each retina. All other objects in the
visual field, either closer or farther away than the fixated object, will fall on different spots on the
retina. When vision is fixed on an object in space, closer objects will fall on the lateral retina of each
eye and more distant objects will fall on the medial retina of either eye. This is easily observed by
holding a finger up in front of your face as you look at a more distant object. You will see two images
of your finger that represent the two disparate images that are falling on either retina.

These depth cues, both monocular and binocular, can be exploited to make the brain think there are
three dimensions in two-dimensional information. This is the basis of 3-D movies. The projected image
on the screen is two-dimensional, but it has disparate information embedded in it. The 3-D glasses
that are available at the theater filter the information so that only one eye sees one version of what is
on the screen, and the other eye sees the other version. If you take the glasses off, the image on the
screen will have varying amounts of blur because both eyes are seeing both layers of information, and
the third dimension will not be evident. Some optical illusions can take advantage of depth cues as
well; though, those are more often using monocular cues to fool the brain into seeing different parts of

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 the scene as being at different depths.

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 Retinal Disparity - Because of the interocular distance, which results in objects of different distances falling

on different spots of the two retinae, the brain can extract depth perception from the two-dimensional

information of the visual field.

IN CONTEXT
Disorders of the Brain: Prosopagnosia
The failures of sensory perception can be unusual and debilitating. A particular sensory deficit that
inhibits an important social function of humans is prosopagnosia, or face blindness. The word comes
from the Greek words prosopa, which means “faces,” and agnosia, which means “not knowing.” Some
people may feel that they cannot recognize people easily by their faces. However, a person with
prosopagnosia cannot recognize the most recognizable people in their respective cultures. They
would not recognize the face of a celebrity, an important historical figure, or even a family member like
their parent. They may not even recognize their own face.

Prosopagnosia can be caused by trauma to the brain, or it can be present from birth. The exact cause
of prosopagnosia and the reason that it happens to some people is unclear. A study of the brains of
people born with the deficit found that a specific region of the brain, the anterior fusiform gyrus of the
temporal lobe, is often underdeveloped. This region of the brain is concerned with the recognition of
visual stimuli and its possible association with memories. Though the evidence is not yet definitive,
this region is likely to be where facial recognition occurs.

Though this can be a devastating condition, people who suffer from it can get by—often by using
other cues to recognize the people they see. Often, the sound of a person’s voice or the presence of
unique cues such as distinct facial features (a mole, for example) or hair color can help the sufferer
recognize a familiar person. In the video on prosopagnosia provided in this section, a woman is shown
having trouble recognizing celebrities, family members, and herself. In some situations, she can use
other cues to help her recognize faces.

 TERMS TO KNOW

Sensory Homunculus
A map of the human body representing the receptor locations within the somatosensory cortex.

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 Primary Sensory Cortex
A region of the cerebral cortex that receives sensory information and processes it to generate an initial
sensory perception.

Association Area
A region of the cerebral cortex connected to the primary sensory cortex that performs additional
processing for more complex sensory perceptions.

Integration Area
A region of the cerebral cortex where information from more than one sensory modality is processed
together for higher level perceptions and functions.

 SUMMARY

In this lesson, you learned how cortical processing of sensory information is done. You first learned that
processing sites in the cerebral cortex are mapped relative to the position of their associated receptor
cells in the body. Then, you learned about the multiple levels of processing that sensory information
undergoes to generate a sensory perception.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Association Area
A region of the cerebral cortex connected to the primary sensory cortex that performs additional
processing for more complex sensory perceptions.

Integration Area
A region of the cerebral cortex where information from more than one sensory modality is processed
together for higher level perceptions and functions.

Primary Sensory Cortex

A region of the cerebral cortex that receives sensory information and processes it to generate an initial
sensory perception.

Sensory Homunculus
A map of the human body representing the receptor locations within the somatosensory cortex.

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 Central Processing: Motor Responses
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about how the brain responds to sensory stimuli by producing motor
stimuli. Specifically, this lesson will cover:
1. Cortical Responses
1a. Secondary Motor Cortices
1b. Primary Motor Cortex
2. Descending Pathways
2a. Appendicular Control
2b. Axial Control
3. Extrapyramidal Controls
4. Anterior Horn Output

 BEFORE YOU START

The defining characteristic of the somatic nervous system is that it controls skeletal muscles. Somatic
senses inform the nervous system about the external environment, but the response to that is through
voluntary muscle movement. The term “voluntary” suggests that there is a conscious decision to make a
movement. However, some aspects of the somatic system use voluntary muscles without conscious control.
One example is the ability of our breathing to switch to unconscious control while we are focused on
another task. However, the muscles that are responsible for the basic process of breathing are also utilized
for speech, which is entirely voluntary.

1. Cortical Responses
Let’s start with sensory stimuli that have been registered through receptor cells and the information relayed to
the CNS along ascending pathways. In the cerebral cortex, the initial processing of sensory perception
progresses to associative processing and then integration areas of the cortex. These levels of processing can
lead to the incorporation of sensory perceptions into memory, but more importantly, they lead to a response.
The completion of cortical processing through the primary, associative, and integrative sensory areas initiates a
similar progression of motor processing, usually in different cortical areas. Whereas the sensory cortical areas
are located in the occipital, temporal, and parietal lobes, motor functions are largely controlled by the frontal
lobe. The most anterior regions of the frontal lobe—the prefrontal areas—are important for executive functions,
which are those cognitive functions that lead to goal-directed behaviors. These higher cognitive processes

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 include working memory, which has been called a “mental scratch pad,” that can help organize and represent
information that is not in the immediate environment. The prefrontal lobe is responsible for aspects of attention,
such as inhibiting distracting thoughts and actions so that a person can focus on a goal and direct behavior
toward achieving that goal.

The functions of the prefrontal cortex are integral to the personality of an individual because it is largely
responsible for what a person intends to do and how they accomplish those plans.

IN CONTEXT
Everyday Connection - Phineas Gage
A famous case of damage to the prefrontal cortex is that of Phineas Gage, dating back to 1848. He
was a railroad worker who had a metal spike impale his prefrontal cortex. He survived the accident,
but according to second-hand accounts, his personality changed drastically. Friends described him as
no longer acting like himself. Whereas he was a hardworking, amiable man before the accident, he
turned into an irritable, temperamental, and lazy man after the accident. Many of the accounts of his
change may have been inflated in the retelling, and some behavior was likely attributable to alcohol
used as a pain medication. However, the accounts suggest that some aspects of his personality did
change. Also, there is new evidence that though his life changed dramatically, he was able to become
a functioning stagecoach driver, suggesting that the brain has the ability to recover even from major

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 trauma such as this.

Phineas Gage - The victim of an accident while working on a railroad in 1848, Phineas Gage had a large iron

rod impaled through the prefrontal cortex of his frontal lobe. After the accident, his personality appeared to

change, but he eventually learned to cope with the trauma and lived as a coach driver even after such a

traumatic event.

1a. Secondary Motor Cortices

In generating motor responses, the executive functions of the prefrontal cortex will need to initiate actual
movements. One way to define the prefrontal area is any region of the frontal lobe that does not elicit
movement when electrically stimulated. These are primarily in the anterior part of the frontal lobe. The regions
of the frontal lobe that remain are the regions of the cortex that produce movement. The prefrontal areas
project into the secondary motor cortices, which include the premotor cortex and the supplemental motor area.

Two important regions that assist in planning and coordinating movements are located adjacent to the primary
motor cortex. The premotor cortex is more lateral, whereas the supplemental motor area is more medial and
superior. The premotor cortex aids in controlling movements of the core muscles to maintain posture during
movement, whereas the supplemental motor area is hypothesized to be responsible for planning and

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 coordinating movement. The supplemental motor area also manages sequential movements that are based on
prior experience (that is, learned movements). Neurons in these areas are most active leading up to the
initiation of movement.

EXAMPLE These areas might prepare the body for the movements necessary to drive a car in
anticipation of a traffic light changing.

1b. Primary Motor Cortex

Recall that the primary motor cortex is located in the precentral gyrus of the frontal lobe. A neurosurgeon,
Walter Penfield, described much of the basic understanding of the primary motor cortex by electrically
stimulating the surface of the cerebrum. Penfield would probe the surface of the cortex while the patient was
only under local anesthesia so that he could observe responses to the stimulation. This led to the belief that the
precentral gyrus directly stimulated muscle movement. We now know that the primary motor cortex receives
input from several areas that aid in planning movement, and its principal output stimulates spinal cord neurons
to stimulate skeletal muscle contraction.

The primary motor cortex is arranged in a similar fashion to the primary somatosensory cortex, in that it has a
topographical map of the body, creating a motor homunculus (see image below). The neurons responsible for
musculature in the feet and lower legs are in the medial wall of the precentral gyrus, with the thighs, trunk, and
shoulder at the crest of the longitudinal fissure. The hand and face are in the lateral face of the gyrus. Also, the
relative space allotted for the different regions is exaggerated in muscles that have greater innervation. The
greatest amount of cortical space is given to muscles that perform fine, agile movements, such as the muscles
of the fingers and the lower face. The “power muscles” that perform coarser movements, such as the buttock
and back muscles, occupy much less space on the motor cortex.

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 Sensory and Motor Homunculi - The primary motor cortex in the precentral gyrus and primary somatosensory cortex

in the postcentral gyrus are both mapped in the form of a homunculus.

 TERMS TO KNOW

Executive Functions
Cognitive functions that lead to goal-directed behaviors.

Working Memory
An executive function that can help organize and represent information that is not in the immediate
environment.

Premotor Cortex
A region of the frontal lobe anterior to the primary motor cortex that aids in controlling movements of
the core muscles to maintain posture during movement.

Supplemental Motor Area

A region of the frontal lobe anterior to the primary motor cortex that is responsible for planning and
coordinating movement.

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 2. Descending Pathways
The motor output from the cortex descends into the brainstem and to the spinal cord to control the musculature
through motor neurons. Neurons located in the primary motor cortex are large cortical neurons referred to as
upper motor neurons that synapse with lower motor neurons in the brainstem or in the spinal cord that go on
to innervate the target skeletal muscle. The two descending pathways traveled by the neurons are the
corticobulbar tract and the corticospinal tract. Both tracts are named for their origin in the cortex and their
targets—either the brainstem (the term “bulbar” refers to the brainstem as the bulb, or enlargement, at the top
of the spinal cord) or the spinal cord.

These two descending pathways are responsible for the conscious or voluntary movements of skeletal muscles.
Any motor command from the primary motor cortex is sent down to upper motor neurons to activate lower
motor neurons in either the cranial motor nuclei of the brainstem or in the anterior horn of the spinal cord. The
axons of the corticobulbar tract project from the cortex to the ipsilateral (same side) motor nucleus. Conversely,
the axons of the corticospinal tract largely cross the midline of the brainstem or spinal cord and synapse on the
contralateral (opposite) side. Therefore, the right motor cortex of the cerebrum controls muscles on the left side
of the body and vice versa.

The corticospinal tract descends from the cortex through the deep white matter of the cerebrum, midbrain, and
pons. Upon entering the medulla, the tracts make up the large white matter tract referred to as the pyramids.
The defining landmark of the medullary-spinal border is the pyramidal decussation, which is where most of the
fibers in the corticospinal tract cross over (decussate) to the opposite side of the brain. At this point, the tract
separates into two parts, which have control over different domains of the musculature.

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© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 191
 Corticospinal Tract - The major descending tract that controls skeletal muscle movements is the corticospinal tract. It

is composed of two neurons, the upper motor neuron and the lower motor neuron. The upper motor neuron has its

cell body in the primary motor cortex of the frontal lobe and synapses on the lower motor neuron, which is in the
anterior horn of the spinal cord and projects to the skeletal muscle in the periphery.

2a. Appendicular Control

The lateral corticospinal tract is composed of the fibers that cross the midline at the pyramidal decussation.
The axons cross over from the anterior position of the pyramids in the medulla to the lateral column of the
spinal cord. These axons are responsible for controlling appendicular muscles and, therefore, the movement of
the arms and legs.

The anterior horn in both the lower cervical spinal cord and the lumbar spinal cord both have wider anterior
horns, representing the greater number of muscles controlled by these motor neurons. The cervical
enlargement is particularly large because there is greater control over the fine musculature of the upper limbs,
particularly of the fingers. The lumbar enlargement is not as significant in appearance because there is less fine
motor control of the lower limbs.

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 2b. Axial Control
The anterior corticospinal tract is responsible for controlling the muscles of the body trunk. These axons do not
decussate (cross over) in the medulla. Instead, they remain in an anterior position as they descend the
brainstem and enter the spinal cord. These axons then travel to the spinal cord level at which they synapse with
a lower motor neuron. Upon reaching the appropriate level, the axons decussate, entering the anterior horn on
the opposite side of the spinal cord from which they entered. In the anterior horn, these axons synapse with
their corresponding lower motor neurons.

Because movements of the body trunk involve both sides of the body, the anterior corticospinal tract is not
entirely contralateral. Some collateral branches of the tract will project into the ipsilateral anterior horn to control
synergistic muscles on that side of the body or to inhibit antagonistic muscles through interneurons within the
anterior horn. Through the influence of both sides of the body, the anterior corticospinal tract can coordinate
postural muscles in broad movements of the body. These coordinating axons in the anterior corticospinal tract
are often considered bilateral, as they are both ipsilateral and contralateral.

 TERMS TO KNOW

Upper Motor Neuron

A neuron in the primary motor cortex that projects to the lower motor neuron in the brainstem or spinal
cord.

Lower Motor Neuron

A neuron in the brainstem or spinal cord that receives a signal from the upper motor neuron and
projects to the target skeletal muscle.

Corticobulbar Tract
A descending pathway that connects the primary motor cortex to the brainstem.

Corticospinal Tract
A descending pathway that connects the primary motor cortex to the spinal cord.

Pyramids
The large white matter region of the corticospinal tract located in the medulla.

Pyramidal Decussation
The location at the medullary-spinal border where most of the fibers in the corticospinal tract cross over
to the opposite side of the brain.

Lateral Corticospinal Tract

A portion of the corticospinal tract that controls appendicular muscles.

Cervical Enlargement
A region in the cervical spinal cord with an increased number of motor neurons in the anterior horn for
control of the movement of the upper limb muscles.

Lumbar Enlargement

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 A region in the lumbar spinal cord with an increased number of motor neurons in the anterior horn for
control of the movement of the lower limb muscles.

Anterior Corticospinal Tract

A portion of the corticospinal tract that controls axial muscles.

3. Extrapyramidal Controls
Other descending connections between the brain and the spinal cord are called the extrapyramidal system.
The name comes from the fact that this system is outside the corticospinal pathway, which includes the
pyramids in the medulla.

The pathways of the extrapyramidal system are influenced by subcortical structures. For example, connections
between the secondary motor cortices and the extrapyramidal system modulate spine and cranium movements.
The basal nuclei, which are important for regulating movement initiated by the CNS, influence the
extrapyramidal system as well as its thalamic feedback to the motor cortex.

 KEY CONCEPT

The conscious movement of our muscles is more complicated than simply sending a single command from
the precentral gyrus down to the proper motor neurons. During the movement of any body part, our
muscles relay information back to the brain, and the brain is constantly sending “revised” instructions back
to the muscles. The cerebellum is important in contributing to the motor system because it compares
cerebral motor commands with proprioceptive feedback. The corticospinal fibers that project to the anterior
horn of the spinal cord have branches that also synapse in the pons, which project to the cerebellum. Also,
the proprioceptive sensations of the dorsal column system have a collateral projection to the medulla that
projects to the cerebellum. These two streams of information are compared in the cerebellar cortex.
Conflicts between the motor commands sent by the cerebrum and body position information provided by
the proprioceptors cause the cerebellum to stimulate the midbrain which then sends corrective commands
to the spinal cord.

EXAMPLE A good example of how the cerebellum corrects cerebral motor commands can be
illustrated by walking in water. An original motor command from the cerebrum to walk will result in a highly
coordinated set of learned movements. However, in water, the body cannot actually perform a typical
walking movement as instructed. The cerebellum can alter motor command, stimulating the leg muscles to
take larger steps to overcome water resistance. The cerebellum can make the necessary changes through
the midbrain. Modulating the basic command to walk also relies on spinal reflexes, but the cerebellum is
responsible for calculating the appropriate response. When the cerebellum does not work properly,
coordination and balance are severely affected. The most dramatic example of this is during the
overconsumption of alcohol. Alcohol inhibits the ability of the cerebellum to interpret proprioceptive
feedback, making it more difficult to coordinate body movements, such as walking in a straight line or
guiding the movement of the hand to touch the tip of the nose.

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  TERM TO KNOW

Extrapyramidal System
A descending pathway that connects the brain and spinal cord outside the corticospinal pathway.

4. Anterior Horn Output

The somatic nervous system provides output strictly to skeletal muscles. The soma (cell body) of the lower
motor neurons, which are responsible for the contraction of these muscles, are found in the anterior horn of the
spinal cord. These large, multipolar neurons have a corona (crown) of dendrites surrounding the cell body and
an axon that extends out of the anterior horn. This axon travels through the anterior nerve root to join the
emerging spinal nerve. The axon is relatively long because it needs to reach muscles in the periphery of the
body.

 DID YOU KNOW

The diameters of cell bodies may be on the order of hundreds of micrometers to support the long axon;
some axons are a meter in length, such as the lumbar motor neurons that innervate muscles in the first
digits of the feet.
The axons will also branch to innervate multiple muscle fibers. Recall that a motor neuron and all the muscle
fibers that it controls make up a motor unit.

 THINK ABOUT IT

Motor units vary in size. Some may contain up to 1000 muscle fibers, such as in the quadriceps, or they may
only have 10 fibers, such as in an extraocular muscle. The number of muscle fibers that are part of a motor
unit corresponds to the precision of control of that muscle. The greater the number and smaller the motor
units in a muscle, the more fine motor control the nervous system has of it. Also, muscles that have finer
motor control have more motor units connecting to them, and this requires a larger topographical field in
the primary motor cortex (larger representation in the homunculus).
Motor neuron axons connect to muscle fibers at a neuromuscular junction. This is a specialized synaptic
structure at which multiple axon terminals synapse with the muscle fiber sarcolemma. The synaptic end bulbs of
the motor neurons secrete the neurotransmitter acetylcholine, which binds to receptors on the sarcolemma.
The binding of acetylcholine opens chemical-gated ion channels, increasing the movement of cations across
the sarcolemma (cell membrane of the muscle cell). This depolarizes the sarcolemma, initiating muscle
contraction. Whereas other synapses result in graded potentials that must reach a threshold in the postsynaptic
target, activity at the neuromuscular junction reliably leads to muscle fiber contraction with every nerve impulse
received from a motor neuron. However, the strength of contraction and the number of fibers that contract can
be affected by the frequency of the motor neuron impulses.

 SUMMARY

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 In this lesson, you learned about cortical responses, also known as motor signals, created by the brain
in response to sensory stimuli. You learned about and reviewed the structures of the brain that control
somatic motor responses, the secondary motor cortices and the primary motor cortex. Then, you
learned the descending pathways that these motor signals use to maintain appendicular control as
well as axial control of skeletal muscle. Then you learned about the additional pathways that exist
called the extrapyramidal controls. Lastly, you reviewed the anterior horn output from the spinal cord
which summarized information about the spinal cord and muscular system to understand the nervous
system’s control over somatic muscle movements.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Anterior Corticospinal Tract

A portion of the corticospinal tract that controls axial muscles.

Cervical Enlargement
A region in the cervical spinal cord with an increased number of motor neurons in the anterior horn for
control of the movement of the upper limb muscles.

Corticobulbar Tract
A descending pathway that connects the primary motor cortex to the brainstem.

Corticospinal Tract
A descending pathway that connects the primary motor cortex to the spinal cord.

Executive Functions
Cognitive functions that lead to goal-directed behaviors.

Extrapyramidal System
A descending pathway that connects the brain and spinal cord outside the corticospinal pathway.

Lateral Corticospinal Tract

A portion of the corticospinal tract that controls appendicular muscles.

Lower Motor Neuron

A neuron in the brainstem or spinal cord that receives a signal from the upper motor neuron and projects
to the target skeletal muscle.

Lumbar Enlargement
A region in the lumbar spinal cord with an increased number of motor neurons in the anterior horn for
control of the movement of the lower limb muscles.

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 Premotor Cortex
A region of the frontal lobe anterior to the primary motor cortex that aids in controlling movements of the
core muscles to maintain posture during movement.

Pryamidal Decussation
The location at the medullary-spinal border where most of the fibers in the corticospinal tract cross over to
the opposite side of the brain.

Pyramids
The large white matter region of the corticospinal tract located in the medulla.

Supplemental Motor Area

A region of the frontal lobe anterior to the primary motor cortex that is responsible for planning and
coordinating movement.

Upper Motor Neuron

A neuron in the primary motor cortex that projects to the lower motor neuron in the brainstem or spinal
cord.

Working Memory
An executive function that can help organize and represent information that is not in the immediate
environment.

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 Divisions of the Autonomic Nervous System
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about the structure and function of the autonomic nervous system.
Specifically, this lesson will cover:
1. Autonomic Nervous System
2. Sympathetic Division
3. Parasympathetic Division
4. Chemical Signaling in the Autonomic Nervous System

 WATCH

Please watch the following video for more information on this topic:

1. Autonomic Nervous System

 BIG IDEA

The nervous system can be divided into multiple ways. Anatomically, it can be divided into central and
peripheral nervous systems. Physiologically, it can be divided by the direction of sensory or motor signals. It
can also be divided by what parts of the body it controls, namely the somatic and autonomic nervous
systems. You’ve already seen that the somatic nervous system receives sensory information to which the
body responds with motor responses that control voluntary muscles (skeletal muscle tissue). The autonomic
nervous system also receives sensory information to which it responds with motor responses. However, the
autonomic nervous system controls involuntary muscle (cardiac and smooth muscle) as well as glandular
tissue. The somatic nervous system is associated with voluntary responses (though many can happen
without conscious awareness, like breathing), and the autonomic nervous system is associated with
involuntary responses, such as those related to homeostasis.
The autonomic nervous system regulates many of the internal organs through a balance of two aspects, or
divisions. In addition to the endocrine system which produces and releases hormones, the autonomic nervous
system is instrumental in homeostatic mechanisms in the body. The two divisions of the autonomic nervous
system are the sympathetic division and the parasympathetic division. The sympathetic division is associated
with the fight, flight, or freeze response, and parasympathetic division is referred to by the epithet of rest and

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 digest response. Homeostasis is the balance between the two systems. At each target effector, one or both
divisions determine activity.

Sympathetic and Parasympathetic Division Effects - Each division of the autonomic nervous system causes its own set

of effects in various target effectors of the body.

Some effectors are controlled by only one division of the autonomic nervous system. However, some are
controlled by both. This dual innervation allows the sympathetic and parasympathetic divisions to work together
to create more fine-tuned control of the activity of that effector.

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 EXAMPLE The heart receives connections from both the sympathetic and parasympathetic divisions.
One causes heart rate to increase, whereas the other causes heart rate to decrease. They work together to
create a wide range of potential heart rates depending on the current conditions.
Recall that the descending pathway for somatic motor signals involves two neurons, the upper and lower motor
neuron. The upper motor neuron projects from the primary motor cortex to the brainstem or spinal cord, while
the lower motor neuron projects from the brainstem or spinal cord to the skeletal muscle effector. The circuitry
for the autonomic nervous system has some similarities and differences.

The autonomic nervous system uses three neurons in the descending pathway for its motor signals.

1. The motor signal originates from its control center in the hypothalamus where the visceral motor neuron
(VMN) projects to the brainstem or spinal cord where it synapses onto a second neuron.
2. From the brainstem or spinal cord, the preganglionic neuron projects to an autonomic ganglion where it
synapses on the third and final neuron.
3. Lastly, the ganglionic neuron projects from the ganglion to the target effector.

Somatic and Autonomic Nervous System Descending Motor Circuits - The somatic nervous system relays motor

signals to their effectors through two neurons, the upper and lower motor neurons. The autonomic nervous system

relays motor signals through three neurons—the visceral motor neuron, preganglionic neuron, and ganglionic neuron.

© 2024 SOPHIA Learning, LLC. SOPHIA is a registered trademark of SOPHIA Learning, LLC. Page 200
 The primary difference between the circuitry in the two divisions of the autonomic nervous system is the length
of the axons and the location of the ganglia. In the sympathetic division, the ganglia are located adjacent to or
near the spinal cord. This causes the axons of the preganglionic neuron, called preganglionic fibers, to be short
and the axons of the ganglionic neuron, called postganglionic fibers to be long. The ganglia of the
parasympathetic division tend to be near or within the target effectors. This causes the preganglionic fibers to
be long while the postganglionic fibers are short.

 TERMS TO KNOW

Sympathetic Division
A division of the autonomic nervous system that is responsible for the fight, flight, or freeze response.

Fight, Flight, or Freeze Response

A set of autonomic responses caused by the activity of the sympathetic nervous system that prepare
the body to stand up to a threat, escape it, or suspend all motor activity.

Parasympathetic Division
A division of the autonomic nervous system that is responsible for the rest and digest response.

Rest and Digest Response

A set of autonomic responses caused by the activity of the parasympathetic nervous system that allows
the body to increase relaxation and digestive functions.

Visceral Motor Neuron

A neuron of the autonomic descending motor pathway which projects from the hypothalamus to the
brainstem or spinal cord.

Preganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from the brainstem or spinal cord
to a ganglion.

Postganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from a ganglion to the target
effector.

Preganglionic Fiber
The axon of a preganglionic neuron.

Postganglionic Fiber
The axon of a ganglionic neuron.

2. Sympathetic Division
When responding to a threat, the body may make preparations to fight, run away, or freeze in place. The
sympathetic division of the autonomic nervous system causes divergent effects as many different effector
organs are activated together for a common purpose.

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  KEY CONCEPT

To coordinate all these responses, the connections in the sympathetic system diverge from a limited region
of the central nervous system (CNS) to a wide array of ganglia that project to the many effector organs
simultaneously. The complex set of structures that compose the output of the sympathetic system makes it
possible for these disparate effectors to come together in a coordinated, systemic change.
The sympathetic division of the autonomic nervous system influences the various organ systems of the body
through connections emerging from the thoracic and upper lumbar spinal cord. The preganglionic neuron in the
lateral horn of any of these spinal regions projects to ganglia adjacent to the vertebral column through the
anterior rami of the spinal nerve. The majority of ganglia of the sympathetic system belong to a network of
sympathetic chain ganglia, also known as paravertebral ganglia, that runs alongside the vertebral column. The
ganglia appear as a series of clusters of neurons linked (or chained) by axonal bridges. There are typically 23
ganglia in the chain on either side of the spinal column:

3 correspond to the cervical region.

12 are in the thoracic region.
4 are in the lumbar region.
4 correspond to the sacral region.

The cervical and sacral levels are not connected to the spinal cord directly through the spinal roots but through
ascending or descending connections through the bridges within the chain.

A diagram that shows the connections of the sympathetic system is somewhat like a circuit diagram that shows
the electrical connections between different receptacles and devices. In the image below, the “circuits” of the
sympathetic system are intentionally simplified.

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 Connections of Sympathetic Division of the Autonomic Nervous System - Neurons from the lateral horn of the spinal

cord (preganglionic nerve fibers - solid lines) project to the chain ganglia on either side of the vertebral column or to

collateral (prevertebral) ganglia that are anterior to the vertebral column in the abdominal cavity. Axons from these

ganglionic neurons (postganglionic nerve fibers - dotted lines) then project to target effectors throughout the body.

To continue with the analogy of the circuit diagram, there are three different types of “junctions” that operate
within the sympathetic system. The first type is the most direct: the sympathetic nerve projects to the chain
ganglion at the same level as the target effector (the organ, tissue, or gland to be innervated). The axon from
the preganglionic fiber (shown as a solid line) synapses with the ganglionic neuron (with the postganglionic fiber
shown as a dashed line). This neuron then projects to a target effector.

In some cases, the target effectors are located superior or inferior to the spinal segment at which the
preganglionic fiber emerges. With respect to the “wiring” involved, the synapse with the ganglionic neuron
occurs at the chain ganglia superior or inferior to the location of the central neuron. An example of this is the
spinal nerve T1 which innervates the eye. The spinal nerve tracks up through the chain until it reaches the
superior cervical ganglion, where it synapses with the postganglionic neuron.

Not all axons from the central neurons terminate in the chain ganglia. Collateral ganglia, also called
prevertebral ganglia, are a group of three ganglia situated anterior to the vertebral column. They are associated
with controlling organs in the abdominal cavity and are also considered part of the enteric nervous system.

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 Sympathetic Connections and Chain Ganglia - The axon from a central sympathetic neuron in the spinal cord can

project to the periphery in a number of different ways. (a) The fiber can project out to the ganglion at the same level

and synapse on a ganglionic neuron. (b) A branch can project to a more superior or inferior ganglion in the chain. (c) A

branch can project through the white ramus communicans but not terminate on a ganglionic neuron in the chain.

Instead, it projects through to a collateral ganglion or the adrenal medulla (not pictured).

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 One type of preganglionic sympathetic fiber does not terminate in a ganglion but instead projects directly to the
target effector, the adrenal medulla. The adrenal medulla is the interior portion of the adrenal gland which sits
on top of each kidney. This gland produces hormones (chemical signaling molecules) and a portion of the
adrenal medulla is under the control of the sympathetic nervous system. These axons are still referred to as
preganglionic fibers, but the target is not a ganglion. The adrenal medulla releases signaling molecules into the
bloodstream, rather than using axons to communicate with target structures.

The projections of the sympathetic division of the autonomic nervous system diverge widely, resulting in a
broad influence of the system throughout the body. All of those physiological changes are going to be required
to occur together to run away from the hunting lioness, or the modern equivalent. This divergence is seen in the
branching patterns of preganglionic sympathetic neurons—a single preganglionic sympathetic neuron may
have 10–20 targets. All of these branches mean that one preganglionic neuron can influence different regions
of the sympathetic system very broadly, by acting on widely distributed organs.

 TERMS TO KNOW

Sympathetic Chain Ganglia

A series of ganglia adjacent to the vertebral column that receive input from sympathetic preganglionic
neurons.

Collateral Ganglia
Three ganglia outside of the sympathetic ganglia that receive input from sympathetic preganglionic
neurons.

Adrenal Medulla
The interior portion of the adrenal gland.

3. Parasympathetic Division
The connections, or “circuits,” of the parasympathetic division are similar to the general layout of the
sympathetic division with a few specific differences.

1. First, the preganglionic fibers of the sympathetic division exit the CNS at the level of the thoracic or lumbar
spinal cord, whereas those of the parasympathetic exit from the brainstem and sacral spinal cord.
2. Second, all preganglionic fibers of the sympathetic division exit the CNS as spinal nerves whereas the
preganglionic fibers of the parasympathetic division that exit from the brainstem exit as cranial nerves.
3. Third, the targets of the parasympathetic preganglionic fibers are terminal ganglia, which are located near
—or even within—the target effector. These ganglia are often referred to as intramural ganglia when they
are found within the walls of the target organ. The postganglionic fiber projects from the terminal ganglia a
short distance to the target effector or to the specific target tissue within the organ.
4. Lastly, as discussed above, the parasympathetic preganglionic fibers are long and the postganglionic fibers
are short compared to those in the sympathetic division because the ganglia are close to—and sometimes

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 within—the target effectors.

Table: Differences in Autonomic Nervous System Circuitry

Difference Sympathetic Parasympathetic

Brainstem and sacral spinal

Exit from CNS Thoracic & lumbar spinal cord
cord

Exiting Nerve Spinal nerves Cranial and spinal nerves

Preganglionic Neuron Sympathetic chain ganglia, collateral ganglia, Terminal ganglia, intramural
Targets adrenal medulla ganglia

Preganglionic Fiber
Short Long
Length

Postganglionic Fiber
Long Short
Length
The cranial component of the parasympathetic system is based on particular nuclei of the brain stem. From the
midbrain, the oculomotor nerve (CN III) extends to a terminal ganglion. From here, it innervates the extraocular
muscles to move the eye while parasympathetic fibers of CN III project to the smooth muscle of the iris to
control pupillary size. In the upper medulla, nuclei contain neurons with axons that project through the facial (CN
VII) and glossopharyngeal nerves (CN IX) to ganglia that control salivary glands. Tear production by the lacrimal
gland is influenced by parasympathetic fibers in the facial nerve (CN VII). Additional neurons in the medulla
project through the vagus nerve (CN X) to the terminal ganglia of the thoracic and abdominal cavities.
Parasympathetic preganglionic fibers primarily influence the heart, bronchi, and esophagus in the thoracic
cavity and the stomach, liver, pancreas, gallbladder, and small intestine of the abdominal cavity. The
postganglionic fibers from the ganglia activated by the vagus nerve are often incorporated into the structure of
the organ, such as the nervous tissue network of the digestive tract organs.

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 Connections of Parasympathetic Division of the Autonomic Nervous System - Neurons from brain-stem nuclei, or from

the lateral horn of the sacral spinal cord, project to terminal ganglia near or within the various organs of the body.

Axons from these ganglionic neurons then project the short distance to those target effectors.

 WATCH

Please watch the following video for more information on this topic:

 TERMS TO KNOW

Terminal Ganglia
Ganglia located near or within the target effector that receive input from the parasympathetic
preganglionic neurons.

Intramural Ganglia
Terminal ganglia that are located within the target effector.

4. Chemical Signaling in the Autonomic Nervous

System
 KEY CONCEPT

Where an autonomic neuron connects with a target, there is a synapse. The electrical signal of the action
potential causes the release of a signaling molecule, which will bind to receptor proteins on the target cell.
Synapses of the autonomic system are classified based on whether they are cholinergic or adrenergic.
These terms refer not only to the signaling molecule that is released but also to the class of receptors that
each binds. Recall that a cholinergic synapse releases the signaling molecule acetylcholine which can bind
either nicotinic or muscarinic receptors. An adrenergic synapse releases the signaling molecule
norepinephrine and can bind multiple receptors.
The adrenergic system has two types of receptors, named the alpha (α)-adrenergic receptor and beta (β)-
adrenergic receptor. Unlike cholinergic receptors, these receptor types are not classified by which drugs can
bind to them. There are two types of α-adrenergic receptors, termed α₁, and α₂, and there are three types of β-
adrenergic receptors, termed β₁, β₂ and β₃. An additional aspect of the adrenergic system is that there is a
second signaling molecule called epinephrine. The chemical difference between norepinephrine and
epinephrine is the addition of a methyl group (CH₃) in epinephrine. The prefix “nor-” actually refers to this
chemical difference, in which a methyl group is missing.

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 The term adrenergic should remind you of the word adrenaline, which is associated with the fight-or-flight
response described at the beginning of the lesson. Adrenaline and epinephrine are two names for the same
molecule. The adrenal gland (ad, on top of; renal, kidney) secretes adrenaline. The ending “-ine” refers to the
chemical being derived, or extracted, from the adrenal gland. A similar construction from Greek instead of Latin
results in the word epinephrine (epi, above; nephr, kidney).

 DID YOU KNOW

In scientific usage, the term epinephrine is preferred in the United States, whereas adrenaline is preferred
in Great Britain, because “adrenalin” was once a registered, proprietary drug name in the United States.
Though the drug is no longer sold, the convention of referring to this molecule by the two different names
persists. Similarly, norepinephrine and noradrenaline are two names for the same molecule.
Having understood the cholinergic and adrenergic systems, their role in the autonomic system is relatively
simple to understand. All preganglionic fibers, both sympathetic and parasympathetic, release ACh. All
ganglionic neurons—the targets of these preganglionic fibers—have nicotinic receptors in their cell membranes.
The nicotinic receptor is a ligand-gated cation channel that results in depolarization of the postsynaptic
membrane. The postganglionic parasympathetic fibers also release ACh, but the receptors on their targets are
muscarinic receptors and do not exclusively cause depolarization of the postsynaptic membrane.
Postganglionic sympathetic fibers release norepinephrine, except for fibers that project to sweat glands and to
blood vessels associated with skeletal muscles, which release ACh.

Autonomic System Signaling Molecules

Sympathetic Parasympathetic

Acetylcholine →
Preganglionic Acetylcholine → nicotinic receptor
nicotinic receptor

Norepinephrine → α- or β-adrenergic receptors Acetylcholine →

Acetylcholine →
Postganglionic muscarinic receptor (associated with sweat glands and the blood
muscarinic receptor
vessels associated with skeletal muscles only)

 TERMS TO KNOW

Adrenergic
A synapse that releases of norepinephrine to bind to α- or β-adrenergic receptors.

Norepinephrine
A signaling molecule released from adrenergic synapses.

Alpha (α)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Beta (β)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Epinephrine
A signaling molecule released from adrenergic synapses.

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  SUMMARY

In this lesson, you learned about the structure and function of the autonomic nervous system. You
learned about the two functional divisions of this system, the sympathetic division and
parasympathetic division, including their effects, circuitry. You also learned about the chemical
signaling in the autonomic nervous system and how it is similar and different between its two divisions.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Adrenal Medulla
The interior portion of the adrenal gland.

Adrenergic
A synapse that releases of norepinephrine to bind to α- or β-adrenergic receptors.

Alpha (α)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Beta (β)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Collateral Ganglia
Three ganglia outside of the sympathetic ganglia that receive input from sympathetic preganglionic
neurons.

Epinephrine
A signaling molecule released from adrenergic synapses.

Fight, Flight, or Freeze Response

A set of autonomic responses caused by the activity of the sympathetic nervous system that prepare the
body to stand up to a threat, escape it, or suspend all motor activity.

Intramural Ganglia
Terminal ganglia that are located within the target effector.

Norepinephrine
A signaling molecule released from adrenergic.

Parasympathetic Division
A division of the autonomic nervous system that is responsible for the rest and digest response.

Postganglionic Fiber

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 The axon of a ganglionic neuron.

Postganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from a ganglion to the target
effector.

Preganglionic Fiber
The axon of a preganglionic neuron.

Preganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from the brainstem or spinal cord to
a ganglion.

Rest and Digest Response

A set of autonomic responses caused by the activity of the parasympathetic nervous system that allows
the body to increase relaxation and digestive functions.

Sympathetic Chain Ganglia

A series of ganglia adjacent to the vertebral column that receive input from sympathetic preganglionic
neurons.

Sympathetic Division
A division of the autonomic nervous system that is responsible for the fight, flight, or freeze response.

Terminal Ganglia
Ganglia located near or within the target effector that receive input from the parasympathetic
preganglionic neurons.

Visceral Motor Neuron

A neuron of the autonomic descending motor pathway which projects from the hypothalamus to the
brainstem or spinal cord.

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 Autonomic Reflexes and Homeostasis
by Sophia

 WHAT'S COVERED

In this lesson, you will learn to identify the structures and functions of autonomic reflexes and how they
differ from somatic reflexes. Specifically, this lesson will cover:
1. The Structure of Reflexes
1a. Afferent Branch
1b. Short and Long Reflexes
2. Balance in Competing Autonomic Reflex Arcs
2a. Competing Neurotransmitters
2b. Autonomic Tone

 BEFORE YOU START

A quick recap… The central nervous system can be divided physiologically into the somatic and autonomic
nervous systems. Both of these receive sensory information from many locations in the body. The primary
functional difference between the somatic and autonomic systems is in the effectors that they target.
Somatic responses are solely based on skeletal muscle contraction. The autonomic system, however,
targets cardiac and smooth muscle, as well as glandular tissue. Both systems contain reflexes whereby
specific sensory inputs elicit specific motor responses. You may recall that the basic circuit is a reflex arc, as
was discussed in previous lessons about homeostasis and the spinal cord. However, in the somatic and
autonomic nervous systems, there are differences in the structure of those reflexes for the somatic and
autonomic systems.

1. The Structure of Reflexes

Recall that a reflex arc contains multiple components. A reflex is first initiated by a change in the environment,
called a stimulus. This change is detected by a receptor and an action potential is generated in a sensory
neuron. This action potential, also referred to as a sensation travels along the afferent pathway to the control
center, most often the CNS. This sensation is processed, and if a motor response is warranted, it travels along
the efferent pathway, also known as a motor neuron to cause a change in the effector. Do you remember this
image?

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 Generic homeostatic arc indicating stimulus, sensor, afferent pathway, control center, efferent pathway, effector, and

effect.

For somatic reflexes and autonomic reflexes, also known as visceral reflexes, the afferent pathway is often the
same between the two systems. Sensory neurons receiving input from the periphery—with cell bodies in the
sensory ganglia, either of a cranial nerve or a posterior root ganglion adjacent to the spinal cord—project into
the CNS to initiate the reflex. The Latin root “effere” means “to carry.” Adding the prefix “ef-” suggests the
meaning “to carry away,” whereas adding the prefix “af-” suggests “to carry toward or inward.”

One difference between a somatic and visceral reflex is in the efferent pathway. The output of a somatic reflex
is the lower motor neuron in the ventral horn of the spinal cord that projects directly to a skeletal muscle to
cause its contraction. The output of a visceral reflex is a two-step pathway starting with the preganglionic fiber
emerging from a lateral horn neuron in the spinal cord, or a cranial nucleus neuron in the brain stem, to a
ganglion—followed by the postganglionic fiber projecting to a target effector.

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 Comparison of Somatic and Visceral Reflexes - The afferent inputs to somatic and visceral reflexes are essentially the
same, whereas the efferent branches are different. Somatic reflexes, for instance, involve a direct connection from the

anterior horn of the spinal cord to the skeletal muscle. Visceral reflexes involve a projection from the preganglionic

neuron to a ganglion, followed by a second projection from the ganglion to the target effector.

1a. Afferent Branch

The afferent branch of a reflex arc does differ between somatic and visceral reflexes in some select instances.
Many of the inputs to visceral reflexes are from special or somatic senses, but particular senses are associated
with the viscera that are not part of the conscious perception of the environment through the somatic nervous
system.

EXAMPLE There is a specific type of mechanoreceptor, called a baroreceptor, in the walls of the large
blood vessels and portions of the heart that senses the stretch of those organs when blood volume or
pressure increases. You do not have a conscious perception of having high blood pressure, but that is an
important afferent branch of the cardiovascular reflexes.
Though visceral senses are not primarily a part of conscious perception, those sensations sometimes make it to
conscious awareness. If a visceral sense is strong enough, it will be perceived. The sensory homunculus—the
representation of the body in the primary somatosensory cortex—only has a small region allotted for the
perception of internal stimuli. If you swallow a large bolus of food, for instance, you will probably feel the lump
of that food as it pushes through your esophagus, or even if your stomach is distended after a large meal. If you
inhale especially cold air, you can feel it as it enters your larynx and trachea. These sensations are not the same
as feeling high blood pressure or blood sugar levels.

When particularly strong visceral sensations rise to the level of conscious perception, the sensations are often
felt in unexpected places.

EXAMPLE Strong visceral sensations of the heart will be felt as pain in the left shoulder and left arm.
This irregular pattern of projection of conscious perception of visceral sensations is called referred pain.
Depending on the organ system affected, the referred pain will project to different areas of the body as shown
in the image below. The location of referred pain is not random but a definitive explanation of the mechanism
has not been established.

 BIG IDEA

The most broadly accepted theory for this phenomenon is that the visceral sensory fibers enter the same
level of the spinal cord as the somatosensory fibers of the referred pain location. By this explanation, the
visceral sensory fibers from the mediastinal region, where the heart is located, would enter the spinal cord
at the same level as the spinal nerves from the shoulder and arm so the brain misinterprets the sensations
from the mediastinal region as being from the axillary and brachial regions. Projections from the medial and
inferior divisions of the cervical ganglia do enter the spinal cord at the middle to lower cervical levels, which
is where the somatosensory fibers enter.

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 Referred Pain Chart - Conscious perception of visceral sensations map to specific regions of the body, as shown in this

chart. Some sensations are felt locally, whereas others are perceived as affecting areas that are quite distant from the
involved organ.

IN CONTEXT
Disorders of the Nervous System: Kehr’s Sign
Kehr’s sign is the presentation of pain in the left shoulder, chest, and neck regions following the
rupture of the spleen. The spleen is in the upper-left abdominopelvic quadrant, but the pain is more in
the shoulder and neck. How can this be?

The sympathetic fibers connected to the spleen are from the celiac ganglion, which would be from the
mid-thoracic to lower thoracic region, whereas parasympathetic fibers are found in the vagus nerve,
which connects in the medulla of the brain stem. However, the neck and shoulder would connect to
the spinal cord at the mid-cervical level of the spinal cord. These connections do not fit with the
expected correspondence of visceral and somatosensory fibers entering at the same level of the
spinal cord.

The incorrect assumption would be that the visceral sensations are coming from the spleen directly. In

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 fact, the visceral fibers are coming from the diaphragm. The nerve connecting to the diaphragm takes
a special route. The phrenic nerve is connected to the spinal cord at cervical levels 3 to 5. The motor
fibers that make up this nerve are responsible for the muscle contractions that drive ventilation. These
fibers have left the spinal cord to enter the phrenic nerve, meaning that spinal cord damage below the
mid-cervical level is not fatal by making ventilation impossible. Therefore, the visceral fibers from the
diaphragm enter the spinal cord at the same level as the somatosensory fibers from the neck and
shoulder.

The diaphragm plays a role in Kehr’s sign because the spleen is just inferior to the diaphragm in the
upper-left quadrant of the abdominopelvic cavity. When the spleen ruptures, blood spills into this
region. The accumulating hemorrhage then puts pressure on the diaphragm. The visceral sensation is
actually in the diaphragm, so the referred pain is in a region of the body that corresponds to the
diaphragm, not the spleen.

1b. Short and Long Reflexes

Somatic reflexes involve sensory neurons that connect sensory receptors to the CNS and motor neurons that
project back out to the skeletal muscles. Visceral reflexes that involve the thoracolumbar or craniosacral
systems share similar connections. However, there are visceral reflexes that do not involve any CNS
components. A long reflex has afferent branches that enter the spinal cord or brain and involve the entire
efferent branch, the preganglionic and ganglionic neurons. A short reflex projects sensory input to the
peripheral ganglion instead of the CNS and the motor response is generated by the ganglionic neuron only.

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 Short and Long Reflexes - Sensory input can stimulate either a short or a long reflex. A sensory neuron can project to

the CNS or to an autonomic ganglion. The short reflex involves the direct stimulation of a postganglionic fiber by the

sensory neuron, whereas the long reflex involves integration in the spinal cord or brain.

A division of the nervous system that is related to the autonomic nervous system is the enteric nervous system.
The word enteric refers to the digestive organs, so this represents the nervous tissue that is part of the
digestive system. There are a few myenteric plexuses in which the nervous tissue in the wall of the digestive
tract organs can directly influence digestive function. If stretch receptors in the stomach are activated by the
filling and distension of the stomach, a short reflex will directly activate the smooth muscle fibers of the stomach
wall to increase motility to digest the excessive food in the stomach. No CNS involvement is needed because
the stretch receptor is directly activating a neuron in the wall of the stomach that causes the smooth muscle to
contract. That neuron, connected to the smooth muscle, is a postganglionic parasympathetic neuron that can
be controlled by a fiber found in the vagus nerve.

 TERMS TO KNOW

Visceral Reflex

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 A reflex of the autonomic nervous system involving an internal organ as an effector.

Referred Pain
The conscious perception of a visceral sensation in a different region of the body.

Long Reflex
A visceral reflex involving the central nervous system.

Short Reflex
A visceral reflex not involving the central nervous system.

2. Balance in Competing Autonomic Reflex Arcs

Recall that the sympathetic and parasympathetic divisions of the autonomic nervous system generally have
opposite effects on the body. Certain organs and structures are only innervated by one of these divisions while
others are innervated and controlled by both which is called dual innervation. In this regard, the autonomic
nervous system’s ability to compete against itself (sympathetic versus parasympathetic) allows it to function in
maintaining homeostasis. At the level of the target effector, the signal to which the system is sending the
message is strictly chemical. A signaling molecule binds to a receptor that causes changes in the target cell,
which in turn causes the tissue or organ to respond to the changing conditions of the body.

2a. Competing Neurotransmitters

 KEY CONCEPT

The postganglionic fibers of the sympathetic and parasympathetic divisions both release neurotransmitters
that bind to receptors on their targets. Postganglionic sympathetic fibers release norepinephrine, with a
minor exception, whereas postganglionic parasympathetic fibers release ACh. For any given target, the
difference in which division of the autonomic nervous system is exerting control is just in what chemical
binds to its receptors. The target cells will have adrenergic and muscarinic receptors. If norepinephrine is
released, it will bind to the adrenergic receptors present on the target cell, and if ACh is released, it will bind
to the muscarinic receptors on the target cell.
In the sympathetic system, there are exceptions to this pattern of dual innervation. The postganglionic
sympathetic fibers that contact the blood vessels within skeletal muscle and that contact sweat glands do not
release norepinephrine; they release ACh. This does not create any problems because there is no
parasympathetic input to the sweat glands. Sweat glands have muscarinic receptors and produce and secrete
sweat in response to the presence of ACh.

At most of the other targets of the autonomic system, the effector response is based on which neurotransmitter
is released and what receptor is present.

EXAMPLE Regions of the heart that establish heart rate are contacted by postganglionic fibers from
both systems. If norepinephrine is released onto those cells, it binds to an adrenergic receptor that causes
the cells to depolarize faster, and the heart rate increases. If ACh is released onto those cells, it binds to a

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 muscarinic receptor that causes the cells to hyperpolarize so that they cannot reach the threshold as easily,
and the heart rate slows.
Without this parasympathetic input, the heart would work at a rate of approximately 100 beats per minute (bpm).
The sympathetic system speeds that up, as it would during exercise, to 120–140 bpm, for example. The
parasympathetic system slows it down to the resting heart rate of 60–80 bpm.

Another example is in the control of pupillary size. The afferent branch responds to light hitting the retina.
Photoreceptors are activated, and the signal is transferred to the retinal ganglion cells that send an action
potential along the optic nerve. If light levels are low, the sympathetic system sends a signal out through the
upper thoracic spinal cord to the sympathetic chain ganglia. The postganglionic fiber then projects to the iris,
where it releases norepinephrine onto the iris (a smooth muscle). When those fibers contract, the pupil dilates—
increasing the amount of light hitting the retina. If light levels are too high, the parasympathetic system sends a
signal out from the midbrain through the oculomotor nerve (CN III) to a terminal ganglion. The postganglionic
fiber then projects to the iris, where it releases ACh onto the circular fibers of the iris—another smooth muscle.
When those fibers contract, the pupil constricts to limit the amount of light hitting the retina.

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 Autonomic Control of Pupillary Size - Activation of the pupillary reflex comes from the amount of light activating the

retinal ganglion cells, as sent along the optic nerve. The output of the sympathetic system projects from the cervical

vertebrae, whereas the parasympathetic system originates out of the midbrain and projects through the oculomotor

nerve to the iris. The postganglionic fibers of either division release neurotransmitters onto the smooth muscles of the

iris to cause changes in the pupillary size. Norepinephrine results in dilation and ACh results in constriction.

In this example, the autonomic system is controlling how much light hits the retina. It is a homeostatic reflex
mechanism that keeps the activation of photoreceptors within certain limits. In the context of avoiding a threat
like a lioness on the savannah, the sympathetic response for fight or flight will increase pupillary diameter so

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 that more light hits the retina and more visual information is available for running away. Likewise, the
parasympathetic response of rest reduces the amount of light reaching the retina, allowing the photoreceptors
to cycle through bleaching and be regenerated for further visual perception; this is what the homeostatic
process is attempting to maintain.

2b. Autonomic Tone

Organ systems are balanced between the input from the sympathetic and parasympathetic divisions. When
something upsets that balance, the homeostatic mechanisms strive to return it to its regular state. For each
organ system, there may be more of a sympathetic or parasympathetic tendency to the resting state, which is
known as the autonomic tone of the system.

EXAMPLE The heart rate was described above. Because the resting heart rate is the result of the
parasympathetic system slowing the heart down from its intrinsic rate of 100 bpm, the heart can be said to
be in a parasympathetic tone.
In a similar fashion, another aspect of the cardiovascular system is primarily under sympathetic control. Blood
pressure is partially determined by the contraction of smooth muscle in the walls of blood vessels. These
tissues have adrenergic receptors that respond to the release of norepinephrine from postganglionic
sympathetic fibers by constricting and increasing blood pressure. The hormones released from the adrenal
medulla—epinephrine and norepinephrine—will also bind to these receptors. Those hormones travel through
the bloodstream where they can easily interact with the receptors in the vessel walls. The parasympathetic
system has no significant input to the systemic blood vessels, so the sympathetic system determines their tone.

There are a limited number of blood vessels that respond to sympathetic input in a different fashion. Blood
vessels in skeletal muscle, particularly those in the lower limbs, are more likely to dilate. It does not have an
overall effect on blood pressure to alter the tone of the vessels but rather allows for blood flow to increase for
those skeletal muscles that will be active in the fight-or-flight response. The blood vessels that have a
parasympathetic projection are limited to those in the erectile tissue of the reproductive organs. Acetylcholine
released by these postganglionic parasympathetic fibers causes the vessels to dilate, leading to the
engorgement of the erectile tissue.

IN CONTEXT
Homeostatic Imbalances - Orthostatic Hypotension
Have you ever stood up quickly and felt dizzy for a moment? This is because, for one reason or
another, blood is not getting to your brain so it is briefly deprived of oxygen. When you change
position from sitting or lying down to standing, your cardiovascular system has to adjust for a new
challenge, keeping blood pumping up into the head while gravity is pulling more and more blood
down into the legs.

The reason for this is a sympathetic reflex that maintains the output of the heart in response to
postural change. When a person stands up, proprioceptors indicate that the body is changing position.
A signal goes to the CNS, which then sends a signal to the upper thoracic spinal cord neurons of the
sympathetic division. The sympathetic system then causes the heart to beat faster and the blood
vessels to constrict. Both changes will make it possible for the cardiovascular system to maintain the

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 rate of blood delivery to the brain. Blood is being pumped superiorly through the internal branch of
the carotid arteries into the brain, against the force of gravity. Gravity is not increasing while standing,
but blood is more likely to flow down into the legs as they are extended for standing. This sympathetic
reflex keeps the brain well oxygenated so that cognitive and other neural processes are not
interrupted.

Sometimes this does not work properly. If the sympathetic system cannot increase cardiac output,
then blood pressure in the brain will decrease and a brief neurological loss can be felt. This can be
brief, as a slight “wooziness” when standing up too quickly, or a loss of balance and neurological
impairment for a period of time. The name for this is orthostatic hypotension, which means that blood
pressure goes below the homeostatic set point when standing. It can be the result of standing up
faster than the reflex can occur, which may be referred to as a benign “head rush,” or it may be the
result of an underlying cause.

There are two basic reasons that orthostatic hypotension can occur. First, blood volume is too low and
the sympathetic reflex is not effective. This hypovolemia may be the result of dehydration or
medications that affect fluid balance, such as diuretics or vasodilators. Both of these medications are
meant to lower blood pressure, which may be necessary in the case of systemic hypertension, and
regulation of the medications may alleviate the problem. Sometimes increasing fluid intake or water
retention through salt intake can improve the situation.

The second underlying cause of orthostatic hypotension is autonomic failure. There are several
disorders that result in compromised sympathetic functions. The disorders range from diabetes to
multiple system atrophy (a loss of control over many systems in the body), and addressing the
underlying condition can improve the hypotension. For example, with diabetes, peripheral nerve
damage can occur, which would affect the postganglionic sympathetic fibers. Getting blood glucose
levels under control can improve neurological deficits associated with diabetes.

 MAKE THE CONNECTION

If you're taking the Anatomy & Physiology I Lab course simultaneously with this lecture, it's a good time to
try the Lab Smooth Muscle: Learn how your gut contracts! in Unit 5 of the Lab course. Review the lab-to-
lecture crosswalk if you need more information. Good luck!

 TERMS TO KNOW

Dual Innervation
When a target effector receives signals from both sympathetic and parasympathetic fibers.

Autonomic Tone
The tendency of an organ in its rested state to receive more sympathetic or parasympathetic input.

 SUMMARY

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 In this lesson, you learned about the structure of reflexes in the body. You learned about how somatic
and autonomic reflexes are similar in their afferent branch, differ in their efferent branch, and differ in
the use of short and long reflexes. Then, you learned about how sympathetic and parasympathetic
divisions find a balance in competing autonomic reflex arcs by using separate but competing
neurotransmitters. Lastly, you learned that even at rest, autonomic tone may still favor sympathetic or
parasympathetic control.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Autonomic Tone
The tendency of an organ in its rested state to receive more sympathetic or parasympathetic input.

Dual Innervation
When a target effector receives signals from both sympathetic and parasympathetic fibers.

Long Reflex
A visceral reflex involving the central nervous system.

Referred Pain
The conscious perception of a visceral sensation in a different region of the body.

Short Reflex
A visceral reflex not involving the central nervous system.

Visceral Reflex
A reflex of the autonomic nervous system involving an internal organ as an effector.

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 Central Control
by Sophia

 WHAT'S COVERED

In this lesson, you will learn about higher order structures in the brain that play a role in autonomic
reflexes and how they work. Specifically, this lesson will cover:
1. Cerebral Structures
1a. Pupillary Reflex
1b. Hypothalamus
1c. Amygdala
2. Medulla Oblongata
3. Pons

1. Cerebral Structures
Autonomic control is based on the visceral reflexes, composed of the afferent and efferent branches. These
homeostatic mechanisms are based on the balance between the two divisions of the autonomic system, which
results in tone for various organs that is based on the predominant input from the sympathetic or
parasympathetic systems. Coordinating that balance requires integration that begins with forebrain structures
like the hypothalamus and continues into the brainstem and spinal cord.

1a. Pupillary Reflex

The pupillary light reflex, as shown in the image below, regulates the size of the pupils in response to the
intensity of light detected by the retina. The greater the intensity, the smaller the pupil. The reflex begins when
light hits the retina. The photoreceptors are activated, causing a signal to travel along the optic nerve. As
previously discussed, sensory signals regarding light in the retina will travel to the visual cortex to become a
conscious perception. However, they also project to the midbrain (pretectal nuclei) which leads to a
parasympathetic response through the oculomotor nerve. Postganglionic fibers from the ciliary ganglion then
stimulate the iris to contract and constrict the pupil.

 THINK ABOUT IT

When light hits the retina in one eye, both pupils contract. When that light is removed, both pupils dilate
again back to the resting position. When the stimulus is unilateral (presented to only one eye), the response
is bilateral (both eyes). The same is not true for somatic reflexes. If you touch a hot radiator, you only pull

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 that arm back, not both. Central control of autonomic reflexes is different from somatic reflexes. The
hypothalamus, along with other CNS locations, controls the autonomic system.

Pupillary Reflex Pathways - The pupil is under competing autonomic control in response to light levels hitting the

retina. The sympathetic system will dilate the pupil when the retina is not receiving enough light, and the
parasympathetic system will constrict the pupil when too much light hits the retina.

1b. Hypothalamus
The hypothalamus is the control center for many homeostatic mechanisms. It regulates both autonomic function
and endocrine function. The role it plays in the pupillary reflexes demonstrates the importance of this control
center. The optic nerve projects primarily to the thalamus, which is the necessary relay to the occipital cortex for
conscious visual perception. Another projection of the optic nerve, however, goes to the hypothalamus.

The hypothalamus then uses this visual system input to drive the pupillary reflexes. If the retina is activated by
high levels of light, the hypothalamus stimulates the parasympathetic response. If the optic nerve message
shows that low levels of light are falling on the retina, the hypothalamus activates the sympathetic response.

The hypothalamus also receives input from other areas of the cerebrum including the olfactory cortex and the
amygdala. These structures inform the hypothalamus about the state of the nervous system and can influence
the regulatory processes of homeostasis. A good example of this is found in the amygdala, which is found
beneath the cerebral cortex of the temporal lobe and plays a role in our ability to remember and feel emotions.

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 1c. Amygdala
The amygdala is a group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe. The
limbic lobe is an arched region of the cerebral cortex that includes structures involved in emotional responses,
as well as structures that contribute to memory function. The limbic lobe has strong connections with the
hypothalamus and influences the state of its activity on the basis of emotional state.

EXAMPLE When you are anxious or scared, the amygdala will send signals to the hypothalamus along
the medial forebrain bundle that will stimulate the sympathetic fight, flight, or freeze response. The
hypothalamus will also stimulate the release of stress hormones through its control of the endocrine system
in response to amygdala input.

The Limbic Lobe - Structures arranged around the edge of the cerebrum constitute the limbic lobe, which includes the

amygdala, hippocampus, and cingulate gyrus, and connects to the hypothalamus.

 TERMS TO KNOW

Pupillary Light Reflex

A visceral reflex that controls the iris to regulate the size of the pupil in response to light intensity levels.

Amygdala
A group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe and plays a
role in our ability to remember and feel emotions.

Limbic Lobe

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 An arched region of the cerebral cortex that includes structures involved in emotional responses, as
well as structures that contribute to memory function.

2. Medulla Oblongata
The medulla oblongata contains nuclei that function as autonomic reflex centers. They receive autonomic
sensory input and in response, they activate various effectors. The first is known as the cardiovascular center.
This reflex center responds to changes in blood pH and pressure by regulating heart rate, the strength of
cardiac contractions, and the flow of blood through peripheral tissues. Two other nuclei known as the
respiratory group respond to changes in oxygen and carbon dioxide concentration in the blood as well as blood
pH by regulating the pace and depth of breathing. The dorsal respiratory group (DRG) regulates inhalation
(breathing in) and breathing rate while the ventral respiratory group (VRG) regulates exhalation (breathing out).
The respiratory groups are further regulated by higher-level centers located in the pons. Furthermore, when
descending inputs from the hypothalamus stimulate these centers, the sympathetic system can increase
activity, such as in response to anxiety or stress.

IN CONTEXT
Everyday Connection - Exercise and the Autonomic System
In addition to its association with the fight, flight, or freeze response and rest-and-digest functions, the
autonomic system is responsible for certain everyday functions. For example, it comes into play when
homeostatic mechanisms dynamically change, such as the physiological changes that accompany
exercise. Getting on the treadmill and putting in a good workout will cause the heart rate to increase,
breathing to be stronger and deeper, sweat glands to activate, and the digestive system to suspend
activity. These are the same physiological changes associated with the fight, flight, or freeze response,
but there is nothing chasing you on that treadmill.

This is not a simple homeostatic mechanism at work because “maintaining the internal environment”
would mean getting all those changes back to their set points. Instead, the sympathetic system has
become active during exercise so that your body can cope with what is happening. A homeostatic
mechanism is dealing with the conscious decision to push the body away from a resting state while
the heart is, in fact, moving away from its homeostatic set point. Without any input from the autonomic
system, the heart would beat at approximately 100 bpm and the parasympathetic system slows that
down to the resting rate of approximately 70 bpm. But in the middle of a good workout, you should
see your heart rate at 120–140 bpm. You could say that the body is stressed because of what you are
doing to it. Homeostatic mechanisms are trying to keep blood pH in the normal range, or to keep body
temperature under control, but those are in response to the choice to exercise.

 TERMS TO KNOW

Cardiovascular Center

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 Autonomic nuclei in the medulla oblongata that regulate heart rate, cardiac contraction strength, and
peripheral blood flow.

Dorsal Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory inhalation and rhythm.

Ventral Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory exhalation.

3. Pons
The pons contains multiple nuclei for cranial nerves V, VI, VII, and VIII along with the pontine respiratory group
(PRG), a set of two nuclei known as the apneustic center and pneumotaxic centers. The apneustic center
stimulates the dorsal respiratory group of the medulla, increasing breathing rate and decreasing depth. The
pneumotaxic center inhibits the DRG, decreasing breathing rate and increasing depth.

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 Brainstem Autonomic Control -The pons and medulla oblongata contain autonomic centers, or reflex nuclei, that

control the rate and depth of breathing. The dorsal and ventral respiratory group in the medulla can be controlled and

i fl i i i

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 influenced by the pontine respiratory group in the pons.

 TERMS TO KNOW

Pontine Respiratory Group

Autonomic nuclei in the pons composed of the apneustic and pneumotaxic centers that regulate the
respiratory nuclei of the medulla.

Apneustic Center
A nucleus of the pontine respiratory group that stimulates the dorsal respiratory group.

Pneumotaxic Center
A nucleus of the pontine respiratory group that inhibits the dorsal respiratory group.

 SUMMARY

In this lesson, you learned about how higher order brain structures play a role in autonomic reflexes.
You first reviewed the pupillary reflex for reference and then focused on how cerebral structures such
as the hypothalamus and amygdala affect this reflex. Then you learned about autonomic reflex centers
in the medulla oblongata and pons as well as how they interact with one another.

Source: THIS CONTENT HAS BEEN ADAPTED FROM OPENSTAX "ANATOMY AND PHYSIOLOGY 2E"
AT openstax.org/details/books/anatomy-and-physiology-2e

 TERMS TO KNOW

Amygdala
A group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe and plays a role
in our ability to remember and feel emotions.

Apneustic Center
A nucleus of the pontine respiratory group that stimulates the dorsal respiratory group.

Cardiovascular Center
Autonomic nuclei in the medulla oblongata that regulate heart rate, cardiac contraction strength, and
peripheral blood flow.

Dorsal Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory inhalation and rhythm.

Limbic Lobe
An arched region of the cerebral cortex that includes structures involved in emotional responses, as well
as structures that contribute to memory function.

Pneumotaxic Center

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 A nucleus of the pontine respiratory group that inhibits the dorsal respiratory group.

Pontine Respiratory Group

Autonomic nuclei in the pons composed of the apneustic and pneumotaxic centers that regulate the
respiratory nuclei of the medulla.

Pupillary Light Reflex

A visceral reflex that controls the iris to regulate the size of the pupil in response to light intensity levels.

Ventral Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory exhalation.

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 Terms to Know
Abducens Nerve
A motor cranial nerve responsible for the movement of the eye; CN VI.

Absolute Refractory Period

The time period during an action potential when a new action potential cannot be generated.

Accessory Nerve
A motor cranial nerve responsible for controlling the muscles of the neck, along with cervical
spinal nerves (also called Spinal Accessory Nerve).

Acquired Reflex
A reflex that is learned over time.

Action Potential
The change in the membrane potential of an excitable cell in response to a stimulus.

Activation Gate
A portion of the voltage-gated sodium channel that opens when the membrane voltage
reaches threshold.

Adrenal Medulla
The interior portion of the adrenal gland.

Adrenergic
A synapse that releases of norepinephrine to bind to α- or β-adrenergic receptors.

Alpha (α)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Ampulla of the Inner Ear

The enlarged region at the base of a semicircular canal containing hair cells.

Amygdala

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 A group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe and
plays a role in our ability to remember and feel emotions.

Anterior Cavity
The space between the cornea and lens.

Anterior Column
White matter region of the spinal cord.

Anterior Corticospinal Tract

A portion of the corticospinal tract that controls axial muscles.

Anterior Horn
Anterior gray matter region of the spinal cord.

Anterior Median Fissure

A deep midline groove on the ventral side of the spinal cord.

Anterior Rami
The branch of the spinal nerve which serves the anterior and lateral portions of the trunk as
well as the limbs.

Anterior Root
Motor axons exiting the spinal cord at the anterior or lateral horn.

Apneustic Center
A nucleus of the pontine respiratory group that stimulates the dorsal respiratory group.

Aqueous Humor
The fluid located in the anterior cavity.

Arachnoid Granulations
Extensions of the arachnoid mater into the dural sinus in the cranial cavity.

Arachnoid Mater
The middle, spider-like layer of the meninges.

Arachnoid Trabeculae

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 Extensions of the arachnoid mater into the subarachnoid space that form a spider-like web.

Arbor Vitae
The branching white matter of the cerebellum.

Ascending Pathway
A sensory pathway that carries peripheral sensations to the brain.

Ascending Tracts of the Spinal Cord

Axons of the spinal columns carrying sensory signals.

Association Area
A region of the cerebral cortex connected to the primary sensory cortex that performs
additional processing for more complex sensory perceptions.

Audition
The special sense of hearing.

Auditory Sensory Neuron

Afferent sensory neurons in the ear that are sensitive to hair cell signals; part of the cochlear
nerve.

Auricle
The fleshy portion of the external ear.

Autonomic Nervous System (ANS)

A functional division of the nervous system which is responsible for involuntary perception
and control of the body.

Autonomic Reflex
A reflex of the autonomic nervous system involving cardiac muscle, smooth muscle, or
glandular tissue effectors.

Autonomic Tone
The tendency of an organ in its rested state to receive more sympathetic or parasympathetic
input.

Axolemma

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 The cell membrane of the axon of a neuron.

Axon Hillock
A tapering of the neuron cell body towards the axon.

Axoplasm
The cytoplasm within the axon of a neuron.

Baroreceptor
A sensory receptor that detects changes in barometric pressure.

Basal Nuclei
Subcortical groups of cell bodies that influence cortical motor signaling to maintain body
position and coordination.

Beta (β)-Adrenergic Receptor

One type of receptor that epinephrine and norepinephrine bind.

Bipolar Cell
A cell that connects the photoreceptor to the retinal ganglion cell.

Bipolar Neuron
A neuron with two processes.

Bleaching
A process by which an opposite visual after-image is perceived after exposure to a bright
light source.

Blood Brain Barrier (BBB)

A physiological barrier between the blood and central nervous system that restricts the
movement of substances into or out of the CNS.

Brachial Plexus
A nerve plexus associated with the lower cervical spinal nerves and first thoracic spinal
nerve.

Brain

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 A nervous system organ located in the cranial cavity and primarily composed of nervous
tissue.

Brainstem
The inferior portion of the brain which contains the midbrain, pons, and medulla oblongata.

Broca’s Area
The region of the frontal lobe that is responsible for production of meaningful language.

Capsaicin
The active molecule in hot peppers which creates the sensation of heat from spicy foods.

Cardiovascular Center
Autonomic nuclei in the medulla oblongata that regulate heart rate, cardiac contraction
strength, and peripheral blood flow.

Cauda Equina
A bundle of spinal nerves that resembles a horse’s tail and extends beyond the conus
medullaris in the vertebral cavity.

Central Canal of the Spinal Cord

A hollow tube in the center of the spinal cord which allows for the circulation of cerebrospinal
fluid.

Central Nervous System (CNS)

The anatomical division of the central nervous system that consists of the brain and spinal
cord.

Central Sulcus
The groove which separates the frontal and parietal lobes of the cerebrum.

Cerebellum
The posterior portion of the brain which functions to coordinate motor and sensory signals;
little brain.

Cerebral Aqueduct
An opening between the third and fourth ventricles that allows for the passage of CSF.

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 Cerebral Cortex
The outer gray matter region of the cerebrum.

Cerebral Hemisphere
Right and left halves of the cerebrum.

Cerebrospinal Fluid (CSF)

Fluid produced by ependymal cells that circulates within the central nervous system.

Cerebrum
The superior region of the brain which controls higher neurological functions.

Cervical Enlargement
A region in the cervical spinal cord with an increased number of motor neurons in the
anterior horn for control of the movement of the upper limb muscles.

Cervical Plexus
A nerve plexus associated with the upper cervical spinal nerves.

Chemical Synapse
A cell-to-cell connection in which a neurotransmitter is released from one cell into a synaptic
cleft and interacts with another cell.

Chemoreceptor
A sensory receptor that detects chemical stimuli.

Cholinergic System
A neurotransmitter system in which acetylcholine is the neurotransmitter used at the
chemical synapse.

Choroid
The vascular tissue of the eye.

Choroid Plexus
A tissue containing ependymal cells that filters blood to form CSF in the ventricles.

Ciliary Body

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 A smooth muscle structure on the interior of the iris that controls the shape of the lens.

Cochlea
The auditory portion of the inner ear.

Cochlear Duct
A cavity within the cochlea that contains the organs of Corti.

Cochlear Nerve
The auditory branch of the vestibulocochlear nerve.

Collateral Ganglia
Three ganglia outside of the sympathetic ganglia that receive input from sympathetic
preganglionic neurons.

Cone Photoreceptor
One of three photoreceptors that is specialized for color vision.

Continuous Propagation
Slow conduction of an action potential along an unmyelinated axon.

Conus Medullaris
The cone-shaped inferior end of the spinal cord.

Cornea
The transparent covering of the anterior tip of the eye that allows light to enter.

Corpora Quadrigemina
A group of two pairs of enlargements (colliculi).

Corticobulbar Tract
A descending pathway that connects the primary motor cortex to the brainstem.

Corticospinal Tract
A descending pathway that connects the primary motor cortex to the spinal cord.

Cranial Nerve

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 One of twelve sets of nerves transporting sensory and motor information into and out of the
brain.

Cranial Nerve Ganglion

A sensory ganglion of a cranial nerve.

Cranial Reflex
A reflex that is processed in the brain.

Cupula
A membrane located in the ampulla of the semicircular canals where the stereocilia of hair
cells are embedded.

Depolarize
The reduction of a membrane potential, making the inside less negative.

Descending Tracts of the Spinal Cord

Axons of the spinal column carrying motor signals.

Diencephalon
Region of the brain deep to the cerebrum and superior to the midbrain.

Dorsal Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory inhalation and
rhythm.

Dual Innervation
When a target effector receives signals from both sympathetic and parasympathetic fibers.

Dura Mater
The outer, fibrous layer of the meninges.

Electrical Synapse
A cell-to-cell connection in which there is a direct connection between the two cells so that
ions can pass directly from one cell to the next.

Encapsulated Ending

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 A sensory nerve ending encapsulated in connective tissue.

Endoneurium
A loose connective tissue wrapping around axons in a nerve.

Enteric Nervous System (ENS)

A division of the autonomic nervous system which is responsible for controlling the smooth
muscle and glandular tissue in your digestive system.

Epinephrine
A signaling molecule released from adrenergic synapses.

Epineurium
A loose connective tissue wrapping around a nerve.

Epithalamus
Plays a role in the limbic system and the production of cerebrospinal fluid.

Excitatory Postsynaptic Potential (EPSP)

A depolarizing graded potential in the postsynaptic membrane.

Executive Functions
Cognitive functions that lead to goal-directed behaviors.

External Ear
A portion of the auditory anatomy including the auricle, ear canal, and tympanic membrane.

Exteroceptor
A receptor that is located near a stimulus in the external environment.

Extrapyramidal System
A descending pathway that connects the brain and spinal cord outside the corticospinal
pathway.

Facial Nerve
A sensory and motor cranial nerve responsible for the muscles involved in facial expressions
as well as part of the sense of taste.

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 Fibrous Tunic
The outer layer of the eye containing the sclera and cornea.

Fight, Flight, or Freeze Response

A set of autonomic responses caused by the activity of the sympathetic nervous system that
prepare the body to stand up to a threat, escape it, or suspend all motor activity.

Filum Terminale
A strand of connective tissue that connects the inferior spinal cord to the sacrum.

Fourth Ventricle
A ventricle located in the brainstem.

Fovea Centralis
The central point in the eye with the greatest visual acuity.

Frontal Lobe
The region of the cerebrum located under the frontal bone and responsible for motor and
cognitive functions.

Ganglion
A group of neuron cell bodies in the peripheral nervous system.

General Sense
A sensory system that is widely distributed throughout the body and in the organs of the
body.

Glossopharyngeal Nerve
A sensory and motor cranial nerve responsible for controlling muscles in the oral cavity and
upper throat as well as part of the sense of taste.

Golgi Tendon Organ

A mechanoreceptor in tendons that detects stretch.

Graded Potential
Local changes in the membrane potential.

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 Gray Matter
Regions of nervous tissue with many cell bodies and dendrites.

Gustation
The sense of taste.

Gustatory Papilla
A raised projection on the surface of the tongue that contains taste buds.

Gustatory Receptor Cell

Sensory cells able to detect taste stimuli.

Hair Cell
Sensory receptor cells of the inner ear that detect changes in the position of their stereocilia.

Hyperpolarize
The increase of a membrane potential beyond its resting state.

Hypoglossal Nerve
A motor cranial nerve responsible for controlling the muscles of the lower throat and tongue.

Hypothalamus
Executive region in charge of the subconscious nervous system control (ANS) and the
endocrine system.

Inactivation Gate
A portion of the voltage-gated sodium channel that closes when the membrane potential
reaches +30 mV.

Incus
The ossicle that connects the malleus and stapes; the anvil.

Inferior Colliculi
A pair of enlargements in the midbrain along the auditory pathway.

Inhibitory Postsynaptic Potential (IPSP)

A hyperpolarizing graded potential in the postsynaptic membrane.

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 Initial Segment
The first portion of the axon of a neuron.

Innate Reflex
A reflex that you are born with.

Inner Ear
The portion of the auditory and vestibular anatomy including the cochlea and vestibule.

Insula
The region of the cerebrum located deep to the temporal and frontal lobes and responsible
for gustatory function.

Integration
A nervous system function in which various stimuli are compared and/or integrated to
produce a response.

Integration Area
A region of the cerebral cortex where information from more than one sensory modality is
processed together for higher level perceptions and functions.

Interaural Intensity Difference

A cue used to help sound localization that compares the intensity (volume) of an auditory
stimuli at each ear.

Interaural Time Difference

A cue used to help sound localization that compares the arrival time of an auditory stimuli at
each ear.

Interoceptor
A receptor that interprets stimuli from internal organs and tissues.

Interventricular Foramina
Openings between the lateral ventricles and the third ventricle that allow for the passage of
CSF.

Intramural Ganglia

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 Terminal ganglia that are located within the target effector.

Iris
The colored ring of the eye.

Kinesthesia
The sensation of body movement.

Lacrimal Gland
A gland superior to the lateral angle of the eye that produces and secretes tears.

Lateral Aperture
One of a pair of openings in the fourth ventricle that allows for passage of CSF into the
subarachnoid space.

Lateral Column
White matter region of the spinal cord.

Lateral Corticospinal Tract

A portion of the corticospinal tract that controls appendicular muscles.

Lateral Horn
Gray matter region of the spinal cord only found in the thoracic, upper lumber, and sacral
regions.

Lateral Sulcus
The groove that separates the temporal lobe from the frontal and anterior parietal lobes of
the cerebrum.

Lateral Ventricle
One of two large ventricles in the cerebrum.

Leakage Channel
A membrane channel that is randomly gated, opening and closing randomly.

Lens
A component of the eye that focuses light on the retina.

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 Ligand-Gated Channel
A membrane channel that opens once a specific signaling molecule binds to it.

Limbic Lobe
An arched region of the cerebral cortex that includes structures involved in emotional
responses, as well as structures that contribute to memory function.

Long Reflex
A visceral reflex involving the central nervous system.

Longitudinal Fissure
The sagittal deep groove which separates the cerebrum into right and left hemispheres.

Lower Motor Neuron

A neuron in the brainstem or spinal cord that receives a signal from the upper motor neuron
and projects to the target skeletal muscle.

Lumbar Enlargement
A region in the lumbar spinal cord with an increased number of motor neurons in the anterior
horn for control of the movement of the lower limb muscles.

Lumbar Plexus
A nerve plexus associated with the lumbar spinal nerves.

Lumbar Puncture
A medical procedure that obtains a small amount of cerebrospinal fluid from the body.

Malleus
The ossicle attached to the tympanic membrane; the hammer.

Mechanically-Gated Channel
A membrane channel that opens because of physical distortion of the cell membrane.

Mechanoreceptor
A sensory receptor that detects physical stimuli.

Median Aperture

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 A single midline opening in the fourth ventricle that allows for passage of CSF into the
subarachnoid space.

Medulla Oblongata
The inferior portion of the brainstem.

Meninges
Three connective tissue membranes which surround and protect the CNS.

Midbrain
The superior portion of the brainstem.

Middle Ear
A portion of the auditory anatomy between the tympanic membrane and bony labyrinth,
including the ossicles.

Monosynaptic Reflex
A reflex that has two neurons and one synapse.

Multipolar Neuron
A neuron with more than two processes.

Muscarinic Receptor
An acetylcholine receptor that is also activated by muscarine.

Muscle Spindles
A mechanoreceptor in skeletal muscles that detects stretch.

Myelin Sheath
A lipid-rich layer of insulation produced by oligodendrocytes and Schwann cells that
surrounds an axon.

Nasolacrimal Duct
A tube that connects the medial angle of the eye to the nasal cavity to drain tears.

Nerve
A bundle of axons in the peripheral nervous system.

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 Nerve Plexus
A network of nerves without cell bodies.

Neural Tunic (aka Retina)

The inner layer of the eye containing sensory receptors and neurons, the retina.

Neuron Fascicle
A bundle of axon fibers wrapped collectively in perineurium.

Nicotinic Receptor
An acetylcholine receptor that is also activated by nicotine.

Nociceptor
A sensory receptor that detects pain stimuli.

Node of Ranvier
A gap in the myelin surrounding a neuron where the axon is exposed.

Nonspecific Ion Channel

An ion channel that is specific to charge but not size.

Norepinephrine
A signaling molecule released from adrenergic.

Nucleus (CNS)
A group of neuron cell bodies in the central nervous system.

Occipital Lobe
The region of the cerebrum located under the occipital bone and responsible for visual
function.

Oculomotor Nerve
A motor cranial nerve responsible for the movement of the eye and pupil; CN III.

Odorant
Chemical molecules able to bind and activate olfactory sensory neurons.

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 Olfaction
The sense of smell.

Olfactory Bulb
An enlargement of the olfactory nerve.

Olfactory Epithelium
The tissue covering the nasal cavity.

Olfactory Nerve
A sensory cranial nerve responsible for the sense of smell; CN I.

Olfactory Sensory Neuron

Afferent sensory neurons in the nose that are sensitive to odorants; part of the olfactory
nerve.

Olfactory Tract
The portion of the olfactory nerve that extends from the olfactory bulb to various regions of
the brain.

Optic Chiasm
The crossover point of the visual pathway where select optic nerve fibers cross to the other
side.

Optic Disc
The point at which all retinal ganglion cell axons exit the eye as the optic nerve; blind spot.

Optic Nerve
A sensory cranial nerve responsible for the sense of vision; CN II.

Optic Tract
The group of axons that project posterior from the optic chiasm.

Organ of Corti
A structure within the cochlear duct that contains hair cells and can send auditory sensory
signals.

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 Osmoreceptor
A sensory receptor that detects changes in solute concentration.

Ossicles
Three small bones located in the middle ear.

Otolith
Calcium carbonate crystals located on top of the otolithic membrane.

Otolithic Membrane
A viscous gelatinous substance in the utricle and saccule of the inner ear that stereocilia of
hair cells are embedded in.

Oval Window
A membrane-covered hole in the cochlea where the stapes is attached.

Parasympathetic Division
A division of the autonomic nervous system that is responsible for the rest and digest
response.

Parietal Lobe
The region of the cerebrum located under the parietal bone and responsible for
somatosensory function.

Parieto-Occipital Sulcus
The groove that separates the medial occipital lobe from the parietal lobe of the cerebrum.

Perception
The central processing of sensory stimuli into a meaningful pattern.

Perineurium
A fibrous connective tissue wrapping around a neuron fascicle in a nerve.

Peripheral Nervous System (PNS)

The anatomical division of the central nervous system that consists of structures outside of
the brain and spinal cord.

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 Photoisomerization
A change in the chemical structure of a photosensitive pigment.

Photon
A single unit of light.

Photoreceptor
Specialized receptor cell in the neural layer of the eye that is sensitive to light stimuli.

Pia Mater
The inner, delicate layer of the meninges.

Pineal Gland
Produces and releases melatonin to regulate circadian rhythms (sleep-wake cycles).

Pneumotaxic Center
A nucleus of the pontine respiratory group that inhibits the dorsal respiratory group.

Polysynaptic Reflex
A reflex that has three or more neurons and two or more synapses.

Pons
The middle portion of the brainstem.

Pontine Respiratory Group

Autonomic nuclei in the pons composed of the apneustic and pneumotaxic centers that
regulate the respiratory nuclei of the medulla.

Postcentral Gyrus
The cerebral gyrus located immediately posterior to the central sulcus.

Posterior Cavity
The space behind the lens.

Posterior Column
White matter region of the spinal cord.

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 Posterior Column System
An ascending tract of the spinal cord associated with fine touch and proprioception.

Posterior Horn
Posterior gray matter region of the spinal cord.

Posterior Median Sulcus

A shallow midline groove on the dorsal side of the spinal cord.

Posterior Rami
The branch of the spinal nerve which serves the posterior portions of the trunk.

Posterior Root
Sensory axons entering the spinal cord at the posterior horn.

Posterior Root Ganglion

A collection of sensory neuron cell bodies along the posterior root.

Postganglionic Fiber
The axon of a ganglionic neuron.

Postganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from a ganglion to the
target effector.

Postsynaptic Potential (PSP)

The graded potential in the dendrites of a neuron that is receiving synapses from other cells.

Precentral Gyrus
The cerebral gyrus located immediately anterior to the central sulcus.

Prefrontal Cortex
The anterior region of the frontal lobe that is responsible for cognitive functions.

Preganglionic Fiber
The axon of a preganglionic neuron.

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 Preganglionic Neuron
A neuron of the autonomic descending motor pathway which projects from the brainstem or
spinal cord to a ganglion.

Premotor Cortex
A region of the frontal lobe anterior to the primary motor cortex that aids in controlling
movements of the core muscles to maintain posture during movement.

Primary Auditory Cortex

The region of the temporal lobe that is responsible for processing auditory sensory
information.

Primary Gustatory Cortex

The region of the insula that is responsible for processing gustatory sensory information.

Primary Motor Cortex

The region of the frontal lobe that is responsible for voluntary motor movement in the body,
located in the precentral gyrus.

Primary Sensory Cortex

A region of the cerebral cortex that receives sensory information and processes it to
generate an initial sensory perception.

Primary Somatosensory Cortex

The region of the parietal lobe that is responsible for processing somatosensory sensory
information, located in the postcentral gyrus.

Primary Visual Cortex

The region of the occipital lobe that is responsible for processing visual sensory information.

Proprioception
The sensation of where the body is in space.

Proprioceptor
An interoceptor located near a moving part of the body that interprets the positions of the
tissues as they move.

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 Pryamidal Decussation
The location at the medullary-spinal border where most of the fibers in the corticospinal tract
cross over to the opposite side of the brain.

Pupil
The black hole at the center of the iris that allows light to pass into the posterior chamber.

Pupillary Light Reflex

A visceral reflex that controls the iris to regulate the size of the pupil in response to light
intensity levels.

Pyramids
The large white matter region of the corticospinal tract located in the medulla.

Rami Communicantes
The branch of the spinal nerve which serves the internal organs of the trunk.

Receptor Cell
A unique sensory cell with structural components that interpret a specific type of stimulus.

Referred Pain
The conscious perception of a visceral sensation in a different region of the body.

Reflex
An automatic motor response to a given sensory signal.

Refractory Period
The time period during an action potential when a new action potential cannot be generated.

Relative Refractory Period

The time period during an action potential when a new action potential can only be
generated by a stronger stimulus.

Repolarize
The return of a membrane potential to its negative state.

Response

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 A nervous system function in which an event is caused to occur as a consequence of a
specific stimulus or stimuli.

Rest and Digest Response

A set of autonomic responses caused by the activity of the parasympathetic nervous system
that allows the body to increase relaxation and digestive functions.

Resting Membrane Potential

The difference in charge across a cell membrane during steady state conditions.

Reticular Formation
A region of gray matter throughout the brainstem that regulates sleep and wakefulness.

Retinal Ganglion Cell

A neuron of the retina which transports a signal along the optic nerve.

Rhodopsin
A photosensitive pigment found in rod photoreceptors.

Rod Photoreceptor
Photoreceptor that is specialized for low-light vision.

Round Window
A membrane-covered hole in the cochlea inferior to the oval window.

Saccule
A structure of the inner ear able to detect head position and vertical acceleration and
deceleration.

Sacral Plexus
A nerve plexus associated with the sacral spinal nerves.

Saltatory Propagation
Fast conduction of an action potential along a myelinated axon.

Sclera
The white of the eye.

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 Semicircular Canal
Structures of the inner ear that detect rotational movement of the head.

Sensation
A nervous system function in which sensory information about the environment is detected
translated into electrochemical signals.

Sensory Homunculus
A map of the human body representing the receptor locations within the somatosensory
cortex.

Short Reflex
A visceral reflex not involving the central nervous system.

Somatic Nervous System (SNS)

A functional division of the nervous system which is responsible for conscious perception
and voluntary motor responses.

Somatic Reflex
A reflex of the somatic nervous system involving a skeletal muscle effector.

Spatial Summation
The combination of various inputs to a neuron with each other.

Special Sense
A sensory system that has a specific organ devoted to it.

Spinal Cord
A nervous system organ located in the vertebral cavity and primarily composed of nervous
tissue.

Spinal Nerve
One of thirty-one sets of nerves transporting sensory and motor information into and out of
the spinal cord through the anterior and posterior roots.

Spinal Reflex
A reflex that is processed in the spinal cord.

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 Spinothalamic Tract
An ascending tract of the spinal cord associated with crude touch and pressure as well as
pain and temperature.

Stapes
The ossicle that connects to the bony labyrinth of the inner ear; the stirrup.

Stereocilia
Microvilli-like extension of the hair cell membrane that attaches to the tectorial membrane.

Subarachnoid Space
The space in which the arachnoid trabeculae are located and cerebrospinal fluid circulates
around the CNS.

Summate
Add together.

Superior Colliculi
A pair of enlargements in the midbrain that integrates visual, auditory, and somatosensory
information.

Supplemental Motor Area

A region of the frontal lobe anterior to the primary motor cortex that is responsible for
planning and coordinating movement.

Sympathetic Chain Ganglia

A series of ganglia adjacent to the vertebral column that receive input from sympathetic
preganglionic neurons.

Sympathetic Division
A division of the autonomic nervous system that is responsible for the fight, flight, or freeze
response.

Synapse
A cell-to-cell connection by which information is transferred and received.

Synaptic End Bulb

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 The expansion at the end of the axon where a synapse is formed.

Systemic Nerve
A major nerve of the peripheral system, distal to a nerve plexus.

Tactile Receptor
A sensory receptor that detects touch, pressure, and vibration.

Taste Bud
Structures within a gustatory papilla that contain gustatory receptor cells.

Taste Hair
Microvilli of the gustatory receptor cell.

Taste Pore
A small opening in the surface of the tongue epithelium.

Tectorial Membrane
A structure of the organ of Corti that stereocilia are connected to.

Tectum
The roof of the midbrain.

Tegmentum
The floor of the midbrain.

Temporal Lobe
The region of the cerebrum located under the temporal bone and responsible for auditory
function.

Temporal Summation
The combination of multiple action potentials from a single cell resulting in a significant
change in the membrane potential.

Terminal Ganglia
Ganglia located near or within the target effector that receive input from the parasympathetic
preganglionic neurons.

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 Thalamus
A collection of nuclei that relay information between the cerebral cortex and the periphery,
spinal cord, or brainstem.

Thermoreceptor
A sensory receptor that detects changes in temperature.

Third Ventricle
A ventricle located in the diencephalon.

Tract
A bundle of axons in the central nervous system.

Trigeminal Nerve
A sensory and motor cranial nerve responsible for cutaneous sensations of the face and
controlling the muscles of mastication.

Trochlear Nerve
A motor cranial nerve responsible for the movement of the eye; CN IV.

Tympanic Membrane
The membrane at the end of the external acoustic meatus; ear drum.

Umami
A savory taste based on the presence of an amino acid.

Unipolar Neuron
A neuron with a single process.

Upper Motor Neuron

A neuron in the primary motor cortex that projects to the lower motor neuron in the
brainstem or spinal cord.

Utricle
A structure of the inner ear able to detect head position and horizontal acceleration and
deceleration.

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 Vagus Nerve
A sensory and motor cranial nerve responsible for contributing to homeostatic control of the
organs of the thoracic and upper abdominal cavities.

Vascular Tunic
The middle layer of the eye containing the choroid, ciliary body, and iris.

Ventral Respiratory Group

An autonomic nucleus in the medulla oblongata that regulates respiratory exhalation.

Ventricle
An open space within the brain where CSF is produced and circulates.

Vestibular Nerve
The vestibular branch of the vestibulocochlear nerve.

Vestibulation
The sense of equilibrium or balance.

Vestibule
The vestibular portion of the inner ear.

Vestibulocochlear Nerve
A sensory cranial nerve responsible for the sense of hearing and equilibrium.

Visceral Motor Neuron

A neuron of the autonomic descending motor pathway which projects from the
hypothalamus to the brainstem or spinal cord.

Visceral Reflex
A reflex of the autonomic nervous system involving an internal organ as an effector.

Vision
The special sense of sight that uses the eye to detect light waves and convert them into
neural signals.

Vitreous Humor

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 The viscous fluid located in the posterior cavity.

Voltage-Gated Channel
A membrane channel that responds to changes in the electrical properties of the membrane.

Wernicke’s Area
The region of the temporal lobe that is responsible for the comprehension of written and
spoken language.

White Matter
Regions of nervous tissue with many axons.

Working Memory
An executive function that can help organize and represent information that is not in the
immediate environment.

Zonule Fiber
Fibrous connection between the ciliary body and lens.

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