THE MUSCULAR SYSTEM

MUSCULAR TISSUE AND HOMEOSTASIS Functions of muscular tissue

Muscular tissue contributes to homeostasis by  Producing body movements

 Stabilizing body positions
producing body movements, moving substances
 Storing and moving substances within the body
through the body, and producing heat to maintain  Generating heat

normal body temperature. Properties of Muscular Tissue

1. Electrical excitability , a property of both muscle and

nerve cells, is the ability to respond to certain stimuli by
Types of Muscular Tissue
producing electrical signals called action potentials
Skeletal muscle tissue is so named because most (impulses).

skeletal muscles move bones of the skeleton. (A few 2. Contractility is the ability of muscular tissue to
contract forcefully when stimulated by an action
skeletal muscles attach to and move the skin or
potential. When a skeletal muscle contracts, it
other skeletal muscles.) Skeletal muscle tissue is generates tension (force of contraction) while pulling on
its attachment points.
striated: Alternating light and dark protein bands
3. Extensibility is the ability of muscular tissue to
(striations) are seen when the tissue is examined
stretch, within limits, without being damaged. The
with a microscope. connective tissue within the muscle limits the range of
extensibility and keeps it within the contractile range of
Smooth muscle tissue is located in the walls of
the muscle cells.
hollow internal structures, such as blood vessels,
4. Elasticity is the ability of muscular tissue to return to
airways, and most organs in the abdominopelvic its original length and shape after contraction or
extension.
cavity. It is also found in the skin, attached to hair
SKELETAL MUSCLE TISSUE
follicles.
Each of your skeletal muscles is a separate organ
Cardiac muscle consists of striated muscle fibers
composed of hundreds to thousands of cells, which
that branch and unite with each other. It forms the
are called muscle fibers because of their elongated
myocardium of the heart. Its fibers tend to be
shapes. Thus, muscle cell and muscle fiber are two
arranged in whorls and spirals, and they have the
terms for the same structure. Skeletal muscle also
property of spontaneous and rhythmic contraction.
contains connective tissues surrounding muscle
Specialized cardiac muscle fibers form the
fibers and whole muscles, and blood vessels and
conducting system of the heart.
nerves.

Connective Tissue Components

Connective tissue surrounds and protects muscular

tissue. The subcutaneous layer or hypodermis,

which separates muscle from skin, is composed of

areolar connective tissue and adipose tissue. It

provides a pathway for nerves, blood vessels, and

lymphatic vessels to enter and exit muscles.

The adipose tissue of the subcutaneous layer stores

most of the body’s triglycerides, serves as an

insulating layer that reduces heat loss, and protects

muscles from physical trauma. Fascia is a dense

sheet or broad band of irregular connective tissue

Microscopic Anatomy of a Skeletal
that lines the body wall and limbs and supports and
Muscle Fiber
surrounds muscles and other organs of the body.

Sarcolemma, Transverse Tubules, and Sarcoplasm

Three layers of connective tissue extend from the
 The multiple nuclei of a skeletal muscle fiber are
fascia to protect and strengthen skeletal muscle
located just beneath the sarcolemma, the
 The outermost layer of dense, irregular plasma membrane of a muscle cell
connectivetissue, encircling the entire muscle, is  Thousands of tiny invaginations of the
the epimysium sarcolemma, called transverse (T) tubules,
 Perimysium is also a layer of dense, irregular tunnel in from the surface toward the center of
connective tissue, but it surrounds groups of 10 each muscle fiber.
to 100 or more muscle fibers, separating them  Within the sarcolemma is the sarcoplasm, the
into bundles called fascicles cytoplasm of a muscle fiber. Sarcoplasm
 Endomysium penetrates the interior of each includes a substantial amount of glycogen,
fascicle and separates individual muscle fibers which is a large molecule composed of many
from oneanother. The endomysium is mostly glucose molecules.
reticular fibers  Glycogen can be used for synthesis of ATP. In
addition, the sarcoplasm contains a red-colored
All three connective tissue layers may extend
protein called myoglobin. This protein, found
beyond the muscle fibers to form a ropelike tendon only in muscle, binds oxygen molecules that
diffuse into muscle fibers from interstitial fluid.
that attaches a muscle to the periosteum of a bone.
Myofibrils and Sarcoplasmic Reticulum

When the connective tissue elements extend as a  At high magnification, the sarcoplasm appears
stuffed with little threads. These small
broad, flat sheet, it is called an aponeurosis structures are the myofibrils, the contractile
organelles of skeletal muscle. Myofibrils are
about 2 um in diameter and extend the entire
length of a muscle fiber.
through the center of each I band. A narrow H
zone in the center of each A band contains thick
 A fluid-filled system of membranous sacs called
but not thin filaments.
the sarcoplasmic reticulum or SR encircles each
 A mnemonic that will help you to remember the
myofibril. This elaborate system is similar to
composition of the I and H bands is as follows:
smooth endoplasmic reticulum in nonmuscular
the letter I is thin (contains thin filaments),
cells. Dilated end sacs of the sarcoplasmic
while the letter H is thick (contains thick
reticulum called terminal cisterns
filaments). Supporting proteins that hold the
thick filaments together at the center of the H
zone form the M line, so named because it is at
the middle of the sarcomere

Filaments and the Sarcomere

 Within myofibrils are smaller protein structures

called filaments or myofilaments
 Thin filaments are 8 nm in diameter and 1–2 um
long† and composed mostly of the protein
actin, while thick filaments are 16 nm in
diameter and 1–2 um long and composed
mostly of the protein myosin.
 The filaments inside a myofibril do not extend
the entire length of a muscle fiber. Instead, they
are arranged in compartments called
sarcomeres
 Narrow, plate-shaped regions of dense protein
material called Z discs separate one sarcomere
from the next. Thus, a sarcomerenextends from
one Z disc to the next Z disc
 The darker middle part of the sarcomere is the
A band, which extends the entire length of the
thick filaments
 Toward each end of the A band is a zone of
overlap, where the thick and thin filaments lie
side by side. The I band is a lighter, less dense
area that contains the rest of the thin filaments
but no thick filaments and a Z disc passes
 At the NMJ, the end of the motor neuron, called
the axon terminal, divides into a cluster of
Muscle Proteins synaptic end bulbs the neural part of the NMJ.
 Myofibrils are built from three kinds of proteins: Suspended in the cytosol within each synaptic
(1) contractile proteins, which generate force end bulb are hundreds of membrane-enclosed
during contraction; (2) regulatory proteins, sacs called synaptic vesicles. Inside each
which help switch the contraction process on synaptic vesicle are thousands of molecules of
and off; and (3) structural proteins, which keep acetylcholine, abbreviated ACh, the
the thick and thin filaments in the proper neurotransmitter released at the NMJ.
alignment, give the myofibril elasticity and
extensibility, and link the myofibrils to the
sarcolemma and extracellular matrix.  The region of the sarcolemma opposite the
 The two contractile proteins in muscle are synaptic end bulbs, called the motor end plate,
myosin and actin, components of thick and thin is the muscle fiber part of the NMJ. Within each
filaments, respectively. Myosin is the main motor end plate are 30 million to 40 million
component of thick filaments and functions as a acetylcholine receptors, integral
motor protein in all three types of muscle transmembrane proteins to which ACh
tissue. specifically binds. These receptors are abundant
 Smaller amounts of two regulatory proteins— in junctional folds, deep grooves in the motor
tropomyosin and troponin—are also part of the end plate that provide a large surface area for
thin filament. In relaxed muscle, myosin is ACh.
blocked from binding to actin because strands
of tropomyosin cover the myosin-binding sites
on actin.

The Neuromuscular Junction

 The neurons that stimulate skeletal muscle

fibers to contract are called somatic motor
neurons. Each somatic motor neuron has a
threadlike axon that extends from the brain or
spinal cord to a group of skeletal muscle fibers.
A muscle fiber contracts in response to one or
more action potentials propagating along its
sarcolemma and through its system of T
tubules. Muscle action potentials arise at the
neuromuscular junction (NMJ)
 A synapse is a region where communication
occurs between two neurons, or between a
neuron and a target cell At most synapses a
small gap, called the synaptic cleft, separates
the two cells. Because the cells do not physically
touch, the action potential cannot “jump the
gap” from one cell to another. Instead, the first
cell communicates with the second by releasing
a chemical called a neurotransmitter.
 In smooth muscle fibers, the thin filaments
attach to structures called dense bodies, which
are functionally similar to Z discs in striated
muscle fibers.

SMOOTH MUSCLE TISSUE

 Smooth muscle tissue is usually activated

involuntarily. Of the two types of smooth
muscle tissue, the more common type is
visceral (single-unit) smooth muscle tissue.
 The second type of smooth muscle tissue,
multiunit smooth muscle tissue, consists of
individual fibers, each with its own motor
neuron terminals and with few gap junctions
between neighboring fibers. Stimulation of one
visceral muscle fiber causes contraction of many
adjacent fibers, but stimulation of one multiunit
fiber causes contraction of that fiber only

Microscopic Anatomy of Smooth Muscle

 A single relaxed smooth muscle fiber is 30–200

um long. It is thickest in the middle (3–8 um)
and tapers at each end. Within each fiber is a
single, oval, centrally located nucleus. The
sarcoplasm of smooth muscle fibers contains
both thick filaments and thin filaments, in ratios
between 1:10 and 1:15, but they are not
arranged in orderly sarcomeres as in striated
muscle. Smooth muscle fibers also contain
intermediate filaments.
THE NERVOUS SYSTEM
2. Ganglia (swelling or knot; singular is ganglion) are
small masses of nervous tissue, consisting primarily of
NERVOUS TISSUE AND HOMEOSTASIS
neuron cell bodies, that are located outside of the brain
The excitable characteristic of nervous tissue allows for and spinal cord. Ganglia are closely associated with
the generation of nerve impulses (action potentials) cranial and spinal nerves.
that provide communication with and regulation of 3. Enteric plexuses are extensive networks of neurons
most body tissues. located in the walls of organs of the gastrointestinal
tract.

Organization of the Nervous System 4. The term sensory receptor refers to a structure of the
nervous system that monitors changes in the external
or internal environment.
With a mass of only 2 kg (4.5 lb), about 3% of the total
body weight, the nervous system is one of the smallest
and yet the most complex of the 11 body systems. This
intricate network of billions of neurons and even more
neuroglia is organized into two main subdivisions: the
central nervous system and the peripheral nervous
system.

Central Nervous System

The central nervous system (CNS) consists of the brain

and spinal cord. The brain is the part of the CNS that is
located in the skull and contains about 100 billion
neurons. The spinal cord is connected to the brain
through the foramen magnum of the occipital bone and
is encircled by the bones of the vertebral column. The
spinal cord contains about 100 million neurons. Division of the PNS

Peripheral Nervous System

The peripheral nervous system (PNS) consists of all a. Somatic nervous system (SNS)- The SNS consists of
nervous tissue outside the CNS. (1) sensory neurons that convey information from
somatic receptors in the head, body wall, and limbs and
1. A nerve is a bundle of hundreds to thousands of from receptors for the special senses of vision, hearing,
axons plus associated connective tissue and blood taste, and smell to the CNS and (2) motor neurons that
vessels that lies outside the brain and spinal cord. conduct impulses from the CNS to skeletal muscles only.
 Twelve pairs of cranial nerves emerge from the Because these motor responses can be consciously
brain controlled, the action of this part of the PNS is
 thirty-one pairs of spinal nerves emerge from voluntary.
the spinal cord. b. autonomic nervous system (ANS)- consists of (1)
sensory neurons that convey information from
autonomic sensory receptors, located primarily in
visceral organs such as the stomach and lungs, to the Parts of a Neuron
CNS and (2) motor neurons that conduct nerve impulses
Most neurons have three parts:
from the CNS to smooth muscle, cardiac muscle, and
glands. The motor part of the ANS consists of two 1. a cell body
branches, the sympathetic division and the 2. dendrites
parasympathetic division. 3. an axon

c. Enteric nervous system (ENS)- The operation of

the ENS, the “brain of the gut,” is involuntary

Functions of the Nervous System

a. Sensory function. Sensory receptors detect internal

stimuli, such as an increase in blood pressure, or
external stimuli (for example, a raindrop landing on
your arm). This sensory information is then carried into
the brain and spinal cord through cranial and spinal
nerves.
Structural Diversity in Neurons
b. Integrative function. The nervous system processes
sensory information by analyzing it and making
decisions for appropriate responses—an activity known
Neurons display great diversity in size and shape.
as integration.
For example, their cell bodies range in diameter
c. Motor function. Once sensory information is
integrated, the nervous system may elicit an from 5 micrometers (slightly smaller than a red
appropriate motor response by activating effectors blood cell) up to 135 um (barely large enough to see
(muscles and glands) through cranial and spinal nerves.
with the unaided eye). The pattern of dendritic
Stimulation of the effectors causes muscles to contract
and glands to secrete. branching is varied and distinctive for neurons in
different parts of the nervous system. A few small
neurons lack an axon, and many others have very
HISTOLOGY OF NERVOUS TISSUE
short axons. As we have already discussed, the
longest axons are almost as long as a person is tall,
Nervous tissue comprises two types of cells—neurons
extending from the toes to the lowest part of the
and neuroglia
brain.

Neurons - Like muscle cells, neurons (nerve cells) Classification of Neurons

possess
electrical excitability, the ability to respond to a Both structural and functional features are used to
stimulus and convert it into an action potential. classify the various neurons in the body.
STRUCTURAL CLASSIFICATION Structurally, or appearance; examples include Purkinje cells in the
cerebellum and pyramidal cells, found in the cerebral
neurons are classified according to the number of
cortex of the brain, which have pyramid-shaped cell
processes extending from the cell body
bodies.

1. Multipolar neurons usually have several dendrites

FUNCTIONAL CLASSIFICATION Functionally,
and one axon. Most neurons in the brain and spinal
neurons are classified according to the direction in
cord are of this type, as well as all motor neurons.
which the nerve impulse (action potential) is conveyed
with respect to the CNS.
2. Bipolar neurons have one main dendrite and one
axon. They are found in the retina of the eye, the inner
1. Sensory or afferent neurons either contain sensory
ear, and the olfactory area of the brain.
receptors at their distal ends (dendrites) or are located
just after sensory receptors that are separate cells.
3. Unipolar neurons have dendrites and one axon that
Once
are fused together to form a continuous process that
an appropriate stimulus activates a sensory receptor,
emerges from the cell body. These neurons are more
the sensory neuron forms an action potential in its axon
appropriately called pseudounipolar neurons because
and the action potential is conveyed into the CNS
they begin in the embryo as bipolar neurons. During
through cranial or spinal nerves. Most sensory neurons
development, the dendrites and axon fuse together and
are unipolar in structure.
become a single process.

2. Motor or efferent neurons convey action potentials

away from the CNS to effectors (muscles and glands)
in the periphery (PNS) through cranial or spinal nerves.
Motor neurons are multipolar in structure.

3. Interneurons or association neurons are mainly

located within the CNS between sensory and motor
neurons. Interneurons integrate (process) incoming
sensory information from sensory neurons and then
elicit a motor response by activating the appropriate
motor neurons. Most interneurons are multipolar in
In addition to the structural classification scheme just
structure.
described, some neurons are named for the histologist
Neuroglia
who first described them or for an aspect of their shape
Neuroglia or glia make up about half the volume of Constrictions in this expanded tube soon appear,
the CNS. Their name derives from the idea of early creating three regions called primary brain vesicles:
histologists that they were the “glue” that held nervous prosencephalon, mesencephalon, and
tissue together. We now know that neuroglia are not rhombencephalon. Both the prosencephalon and
merely passive bystanders but rather actively rhombencephalon subdivide further, forming
participate in the activities of nervous tissue. secondary brain vesicles. The prosencephalon, or
Generally, neuroglia are smaller than neurons, and they forebrain, gives rise to the telencephalon and
are 5 to 25 times more numerous. diencephalon, and the rhombencephalon, or hindbrain,
develops into the metencephalon and myelencephalon.
Neuroglia of the CNS

Neuroglia of the CNS can be classified on the basis

of size, cytoplasmic processes, and intracellular
organization into four types:

 Astrocytes
 Oligodendrocytes
 Microglia
 ependymal cells

Major Parts of the Brain

Neuroglia of the PNS
The adult brain consists of four major parts:
Neuroglia of the PNS completely surround axons
 brain stem
and cell bodies.  cerebellum,
 diencephalon
The two types of glial cells in the PNS are
 cerebrum
 Schwann cells
 satellite cells
Protective Coverings of the Brain

BRAIN ORGANIZATION,
The cranium and the cranial meninges surround and
PROTECTION, AND BLOOD
protect the brain. The cranial meninges are
SUPPLY
continuous with the spinal meninges, have the same
basic structure, and bear the same names: the outer
The brain and spinal cord develop from the ectodermal
dura mater, the middle arachnoid mater, and the
neural tube. The anterior part of the neural tube
inner pia mater.
expands, along with the associated neural crest tissue.
Three extensions of the dura mater separate parts of Cerebrospinal fluid (CSF) is a clear, colorless liquid
the brain:
composed primarily of water that protects the brain and
spinal cord from chemical and physical injuries. It also
1. The falx cerebri separates the two hemispheres
carries small amounts of oxygen, glucose, and other
(sides) of the cerebrum.
needed chemicals from the blood to neurons and
2. The falx cerebelli separates the two hemispheres
neuroglia. CSF continuously circulates through cavities
of the cerebellum.
in the brain and spinal cord and around the brain and
3. The tentorium cerebelli separates the cerebrum
spinal cord in the subarachnoid space (the space
from the cerebellum
between the arachnoid mater and pia mater). The total
volume of CSF is 80 to 150 mL (3 to 5 oz) in an adult.

The four CSF-filled cavities within the brain, which are

called ventricles. There is one lateral ventricle in
each hemisphere of the cerebrum. (Think of them as
ventricles 1 and 2.) Anteriorly, the lateral ventricles are
separated by a thin membrane, the septum
pellucidum. The third ventricle is a narrow slitlike
cavity along the midline superior to the hypothalamus
and between the right and left halves of the thalamus.
The fourth ventricle lies between the brain stem and
the cerebellum.

Functions of CSF

The CSF has three basic functions:

1. Mechanical protection. CSF serves as a
shock-absorbing medium that protects the delicate
tissues of the brain and spinal cord from jolts that
The blood–brain barrier (BBB) consists mainly of
would otherwise cause them to hit the bony walls of
tight junctions that seal together the endothelial cells of
the cranial cavity and vertebral canal. The fluid also
brain blood capillaries and a thick basement membrane
buoys the brain so that it “floats” in the cranial cavity.
that surrounds the capillaries

2. Homeostatic function. The pH of the CSF affects

CEREBROSPINAL FLUID
pulmonary ventilation and cerebral blood flow, which
is important in maintaining homeostatic controls for Medulla Oblongata
brain tissue. CSF also serves as a transport system for The medulla oblongata, or more simply the

polypeptide hormones secreted by hypothalamic medulla, is continuous with the superior part of the

neurons that act at remote sites in the brain. spinal cord; it forms the inferior part of the brain
stem. The medulla begins at the foramen magnum

3. Circulation. CSF is a medium for minor exchange and extends to the inferior border of the pons, a

of nutrients and waste products between the blood and distance of about 3 cm (1.2 in.).

adjacent nervous tissue.

Pons

Formation of CSF in the Ventricles

The pons lies directly superior to the medulla and

The majority of CSF production is from the choroid anterior to the cerebellum and is about 2.5 cm (1 in.)

plexuses, networks of blood capillaries in the walls of long. Like the medulla, the pons consists of both

the ventricles. Ependymal cells joined by tight nuclei and tracts. As its name implies, the pons is a

junctions cover the capillaries of the choroid plexuses. bridge that connects parts of the brain with one

Selected substances (mostly water) from the blood another. These connections are provided by bundles

plasma, which are filtered from the capillaries, are of axons. Some axons of the pons connect the right

secreted by the ependymal cells to produce the and left sides of the cerebellum. Others are part of

cerebrospinal fluid ascending sensory tracts and descending motor

tracts.

This blood–cerebrospinal fluid barrier permits

certain substances to enter the CSF but excludes others, Midbrain

protecting the brain and spinal cord from potentially

harmful blood-borne substances. In contrast to the The midbrain or mesencephalon extends from the

blood–brain barrier, which is formed mainly by tight pons to the diencephalon and is about 2.5 cm (1 in.)

junctions of brain capillary endothelial cells, the long. The aqueduct of the midbrain (cerebral

blood–cerebrospinal fluid barrier is formed by tight aqueduct) passes through the midbrain, connecting

junctions of ependymal cells. the third ventricle above with the fourth ventricle
below. Like the medulla and the pons, the midbrain
contains both nuclei and tracts

THE BRAIN STEM AND

Reticular Formation
RETICULAR FORMATION
In addition to the well-defined nuclei already
described, much of the brain stem consists of small The cerebellum is posterior to the medulla and pons
clusters of neuronal cell bodies (gray matter) and inferior to the posterior portion of the cerebrum. A
interspersed among small bundles of myelinated deep groove known as the transverse fissure, along
axons (white matter). The broad region where white with the tentorium cerebelli, which supports the
matter and gray matter exhibit a netlike arrangement posterior part of the cerebrum, separates the
cerebellum
is known as the reticular formation. It extends
from the cerebrum.
from the superior part of the spinal cord, throughout
the brain stem, and into the inferior part of the
THE DIENCEPHALON
diencephalon. Neurons within the reticular
formation have both ascending (sensory) and
The diencephalon forms a central core of brain tissue
descending (motor) functions.
just superior to the midbrain. It is almost completely
surrounded by the cerebral hemispheres and contains
The ascending portion of the reticular formation is
numerous nuclei involved in a wide variety of sensory
called the reticular activating system (RAS), which
and motor processing between higher and lower brain
consists of sensory axons that project to the cerebral
centers. The diencephalon extends from the brain stem
cortex, both directly and through the thalamus. Many
to the cerebrum and surrounds the third ventricle; it
sensory stimuli can activate the ascending portion of
includes
the RAS. Among these are visual and auditory stimuli;
 Thalamus
mental activities; stimuli from pain, touch, and
 Hypothalamus
pressure receptors; and receptors in our limbs and head  epithalamus.
 Subthalamus
that keep us aware of the position of our body parts.

THE CEREBRUM
THE CEREBELLUM

The cerebrum is the “seat of intelligence.” It provides

The cerebellum, second only to the cerebrum in size,
us with the ability to read, write, and speak; to make
occupies the inferior and posterior aspects of the
calculations and compose music; and to remember the
cranial cavity. Like the cerebrum, the cerebellum has a
past, plan for the future, and imagine things that have
highly folded surface that greatly increases the surface
never existed before. The cerebrum consists of an outer
area of its outer gray matter cortex, allowing for a
cerebral cortex, an internal region of cerebral white
greater number of neurons. The cerebellum accounts
matter, and gray matter nuclei deep within the white
for about a tenth of the brain mass yet contains nearly
matter.
half of the neurons in the brain.
The Spinal Cord

About 100 million neurons and even more neuroglia

compose the spinal cord, the part of the central
nervous system that extends from the brain. The spinal
cord and its associated spinal nerves contain neural
circuits that control some of your most rapid reactions
to environmental changes. If you pick up something
hot, the grasping muscles may relax and you may drop
the hot object even before you are consciously aware
of the extreme heat or pain.