NERVOUS TISSUE here.

They send messages in the form of neurotransmitters to other

cells through synapses. But that’s the platonic model of a neuron. In
reality, neurons are one of the most diverse cell types in the body.
Nervous Tissue Histology Explained for Beginners Some axons are thin, bare cables, while some have a squishy layer
around them that helps them transmit signals faster.
It’s called a myelin sheath, so we say that those neurons are
With cells named after stars and neurons that branch out like
myelinated. The neurons we saw on longitudinal view are really
bare trees in the winter. The nervous system is beautiful under a
myelin with axons inside, and in this cross section view, you can see
microscope, but sometimes it looks like a cluttered mess.
the tiny axon with the marshmallowy myelin all around it, and
In this video, I’ll teach you how to look at the histology of the
endoneurium around that. Length and diameters can change too. Like
different types of nervous system cells so you can appreciate what
the myelinated type 1A fibers are anywhere from 4-20 micrometers
you see under a microscope.
wide, Type B fibers are 1 to 4 micrometers wide, while the
If you’re new to the channel, welcome, my name is Patrick and
unmyelinated Type C fibers are only 0.2 to 1.5 micrometers wide.
this channel is all about anatomy and how we learn about it. As
And the wider and more myelinated the neuron, the faster it transmits
always, I have the accompanying notes for this video linked in the
electrical impulses. So those type 1As send signals at 70 to 120
description if you want to check those out. Otherwise, let’s get
meters a second while type C conducts at half to two and half meters
started.
per second. Pretty big difference in size and speed. Not only can
Our biggest challenge in learning nervous system histology is
axons vary, but the branching pattern can vary too.
figuring out big picture anatomy from microscopic anatomy, but we
The most common type of neuron is a multipolar neuron — it
can use some clues to help us out.
has one axon and a cell body with a bunch of branching dendrites.
We can split the nervous system into the central nervous
You’ll usually spot these on the brain and spinal cord. Meanwhile,
system, which includes the brain and spinal cord, and the peripheral
bipolar neurons have a long axon and a single dendritic tree poking
nervous system, pretty much all the nerves that branch out from that.
out the other end and you only see these in certain sensory systems
We’ll see some different structures and cell types depending on
like the nose and retina since they only send afferent, or sensory
where we look, but the overall purpose of the nervous system is to
information. Finally, unipolar neurons are what they sound like, they
send and receive electrical signals. And that helps us deduce the
have a cell body and a single axon— no dendrites. But neurons
anatomy of interest. Like the power lines that send electricity through
aren’t the only type of cell in the nervous system. We also have glial
a city, each nerve is made of clusters of smaller neuron cells. Each of
cells, essentially supportive cells. For instance, astrocytes, or star
these little circles is part of individual neuron cells.
shaped cells support and protect our neurons by regulating the blood
If we took a transverse cross section of a nerve, we’d get a slide
brain barrier, helping form synapses, and clearing excess
like this. Just like this electrical cable. And when we slice a nerve
neurotransmitters. They’re kind of hard to see with traditional light
long ways for a longitudinal view and see the long axons running the
microscopes, so unless you have an electron microscope you
length of the nerve.
probably won’t get quizzed on it.
So while a picture like this nerve cross section seems overly
Oligodendrocytes are another fun one — they help make the
busy at first, see it for what it is, neurons and the tissue that wraps
myelin sheath around neurons in the brain and spinal cord, while
them into little bundles. And if you’re already familiar with the
Schwann cells make the myelin in the peripheral nerves.
connective tissue around muscle bundles, then the naming
Quick summary, this all started with our bundles of neurons
conventions are gonna come easy for nerves.
organized into peripheral nerves like electrical wires in a cable. But
Remember how for muscles you have perimysium, epimysium
we still have some big deal nervous tissue to tackle — the central
and endomysium?
nervous system, including the brain and spinal cord. Luckily for us,
Well in nerves, you keep the same prefixes, but instead of
we can get our bearings with the spinal cord similarly to how we did
-mysium for muscle, you have -nerium for nerves.
the peripheral nerves. The longitudinal section looks familiar, but
So the outermost layer is epineurium, a layer of dense irregular
different and the transverse cross section is super unique. This cross
connective tissue. Then each bundle, or fascicle, is wrapped in a
section is this diagram, or what I call the butterfly pancake.
thinner connective tissue called the perineurium while each neuron
So at the tissue level, let’s see what we’re working with. There
cell, and all its accessories are wrapped in endoneurium.
are two different colors to work with — which come from myelin
In this slide, you can clearly see the dense tissue wrapping up
status. Since those myelin sheaths are so fatty and fluffy, think of
the neuron bundle here— that’s the perineurium, while each light
myelinated fibers like marshmallows that make up white matter,
colored neuron has a dark ring around it — that’s the endoneurium.
while those dense, slow, unmyelinated fibers are the metallic skewers
The cell that lives inside that connective tissue is called a
that poke through them, making up the gray matter Look, I know that
neuron and we’re only looking at a small section of it on a cross
sounds backwards, the darker color should be gray matter, but I don’t
section.
make the rules. Take it up with management. Since the gray matter
I’m guessing you’ve seen a picture like this before — the
falls into this shape, we label these segments horns. And we have
cliche illustrated neuron with all the tidy pieces, right?
anterior, lateral, and dorsal horns.
We’re only looking at this small section. That’s because actual
But there’s another big component to the central nervous
neuron cells can be really long, and impossible to fit under a
system, the brain. Let’s look at these two different colors, since their
microscope slide.
tissue level anatomy is different.
But depending on where we’re looking, we can identify
Before we get to neurons, we have a few layers of connective
different pieces of them. This big boy is the cell body, or soma,
tissue called the meninges. If you’ve heard of the diseases meningitis
which has a nucleus inside. We need to remember that, as cool and
it’s inflammation of these layers. The most superficial layer is the
specialized as these cells are, neurons are still cells with DNA and
dura mater, a layer of dense connective tissue that sticks to the skull.
organelles.
Deeper than that is the arachnoid layer which is thin and looks like
Branching out from there are any number of dendrites,
spider webs, hence the name, and connects to the delicate thin pia
branches that collect electrical impulses from other cells.
mater underneath. And aside from some connective tissue around
They sum up here at the axon hillock where an impulse will
blood vessels, all the other structures of the brain can be classified as
travel down the axon, this long piece here. The axon can be over
nervous tissue.
95% of the volume of the neuron cell — and they can be long. Like
But like I said, layers. The outermost layer of the cerebrum is
over a meter long. These axons are what we just cut open on the
the cerebral cortex, and deeper than that, the sub cortical white
cross section and most of what we see on longitudinal sections.
matter. The cerebral cortex has 6 layers of its own and only a couple
Finally, the neuron ends at the axon terminals, these tiny branches
of cell types to differentiate between. The most common of which are
pyramidal neurons, named because sure, they look like pyramids I
guess. But they’re also easy to spot because they stain a dark blue
color and have real big nuclei. 75% of cells in the cerebral cortex will
be pyramidal cells.
These other dots are called stellate cells, that look kind of like
stars too, but they’re really tiny so they also get called granular cells.
The deepest layer of the cerebral cortex features fusiform cells,
which gives away their shape. Fusiform is the name for a shape —
just like fusiform muscles, these cells are spindle shaped. And
remember that this is gray matter, they’re unmyelinated neurons.
When we get to the subcortical white matter, we’ll see more
oligodendroglial cells, some of the cells that actually make the
myelin. This is really just the tip of the iceberg with neuroanatomy,
but I didn’t want tomake this video overwhelming.
If you need more help with histology in general though, I’ve
made a bunch of them and you can find a playlist of them here. As
always, thanks to my patrons over on Patreon. And if you haven’t I’d
appreciate it if you subscribed and hit the bell so you get notified
when I post a new video. Have fun, be good.
Types of Tissue Part 4: Nervous Tissue
It’s Professor Dave, let’s talk about the nervous system. The
human body is a finely tuned machine, and perhaps its most
impressive feature is the way that we can use sentience to command
this machine at will. If you decide to get up out of your chair and
walk to the fridge, the body obeys. But how does this work? Just as
with everything else, it’s not magic.
It is the result of a staggering amount of chemistry happening
on the molecular level, and the system that carries signals from the
brain to the rest of the body is called the nervous system. The cells
that comprise the nervous system qualify as the fourth type of tissue
that epithelial tissue we will examine in this series. We looked at
epithelial, connective, and muscle tissues, so now let’s learn about
nervous tissue.
The main type of cell that makes up the nervous system is
called a neuron. These look very different from the cells we are
familiar with thus far, in that there is an incredible amount of
branching within each neuron, where cytoplasmic extensions project
in many directions, allowing a neuron to respond to stimuli and
transmit electrical impulses over very long distances.
These are the very same signals that stimulate the muscle
contraction we learned about earlier in the series. Neurons are
amitotic, meaning they do not divide, and they have the potential to
live neurons do not divide (amitotic) a hundred years or more, so the
neurons we are born with last a lifetime.
Beyond neurons there are also supporting cells in the nervous
system called neuroglia or glial cells, which are not neurons, but
rather wrap around delicate parts of neurons for protection.
Let’s get a closer look at a neuron now. First, the central part is
called the cell body. This contains a nucleus and a nucleolus,
surrounded by all the typical organelles we learned about in the
biology series. Projecting from the cell body we can see many
dendrites, which produce a tremendous surface area for receiving
signals.
We also see a single axon. This initiates from an axon hillock,
and extends for some distance. For many neurons, the axon accounts
for the majority of the length of the cell, sometimes extending
macroscopic distances, at which point we can call them nerve fibers.
At the terminus of the axon there will be many terminal branches,
sometimes thousands, which are the axon terminals that interface
with other neurons, or with muscles to form the neuromuscular
junctions we discussed earlier in the series. The axon is the
conducting region of the neuron, in that it generates electrical signals
for communicating with other neurons, starting at the axon hillock
and traveling through to the axon terminals, also called the secretory
region.
When the impulse gets here, neurotransmitters are released
which will then either excite or inhibit nearby nerve cells. Many
axons are protected by a myelin sheath, which also increases the
speed at which impulses are transmitted, and nerve fibers covered in
this manner are called myelinated fibers.
Particularly in the peripheral nervous system, the myelin sheath
is often made of individual units called Schwann cells. These first
envelop the nerve fiber, and then wrap themselves around it many
times, which results in numerous layers of plasma membrane to
protect the nerve. These are membranes which lack most of the
channel proteins found in other types of cells, making them great
electrical insulators, and allowing them to increase the speed of
electrical conduction down an axon. There are tiny gaps in between
each Schwann cell where a bit of the fiber is exposed, and these are
called nodes of Ranvier, and sometimes axon collaterals branch out
at these gaps.
Now we want to be aware that there are several types of
neurons, and just like some other anatomical features, we can classify
them either by structure or by function. Going by structure first,
neurons can be unipolar, bipolar, or multipolar. These terms refer to
the number of processes extending from the cell body.
A unipolar neuron has a single axon that quickly divides into
proximal and distal branches, so this still technically qualifies as a
single axon. One of these is a central process that goes towards the
central nervous system, while the other is a peripheral process that
acts as a sensory receptor. Then there are bipolar neurons, which
have two processes, one axon and one dendrite, extending from
opposite sides of the cell body. These are not very common but they
are found in certain places like specific parts of the eye and nose.
Then, multipolar neurons have three or more processes, precisely one
of which is an axon, and the rest of which are dendrites. Around
ninety-nine percent of our neurons are of this type.
Then we can describe function. A sensory, or afferent neuron,
transmits information from sensory receptors towards the central
nervous system. These are typically unipolar. Motor, or efferent
neurons, transmit information from the central nervous system to
muscles and glands. These are always multipolar. Then there are
interneurons, which sit between the other two types and help shuttle
signals around the system. These are typically multipolar neurons,
found entirely within the central nervous system, and most of our
neurons are of this type. For any neuron, regardless of type, we can
describe some basic common features.
Common features of neurons. Any neuron will have a receptive
region, where a stimulus is received, typically at dendrites. Then
there is a trigger zone, which initiates a conducting region, where the
electrical signal travels. This leads to a secretory region with axon
terminals that release neurotransmitters. That covers the basics
regarding the structure of a nerve cell. But how exactly does it
generate electrical impulses? This could be discussed for hours upon
hours, but let’s go over a very basic summary here.
First we must understand the concept of membrane potential.
For a more rigorous discussion of electric potential from a physics
standpoint, check out my tutorial on that subject now, otherwise for
our purposes here, we just need to know that opposite charges attract,
so if there exist opposite charges near one another, work must be
done to separate them. If these regions are already separated within a
cell, there is an opportunity to use the potential energy that exists by
virtue of their separation. This charge separation exists within nerve
cells due to the concentrations of specific ions that sit inside and
outside of the plasma membrane, so we say there is a potential
difference across that membrane, and the membrane itself resists the
current flow, as formally charged ions have difficulty traversing the
nonpolar section of the membrane. However, there are ion channels
in the membrane which can allow specific ions through at specific
times. Some of these are non-gated, meaning they remain open.
Others are gated, meaning they are closed, and only open as the
result of a particular signal.
 Chemically-gated channels open when a specific
neurotransmitter binds. Voltage-gated channels respond to changes in
the membrane potential. And mechanically gated channels open
when the receptor becomes physically deformed. When these
channels open, ions can freely diffuse through, obeying the
electrochemical gradient, which seeks to balance charge, so an
electrical current is generated.
A resting neuron has a resting membrane potential, based on
the sodium and potassium ion concentration in and out of the cell,
due to their differing abilities to diffuse across the membrane, and it
is maintained by sodium-potassium pumps that keep the
concentration gradient as it is. But when a signal is received, whether
from sensory input or a neurotransmitter from another neuron, there
will be a change in this potential which can produce a signal. This
signal will be either a graded potential, which operates over short
distances, or an action potential, which operates over long distances,
like the length of an axon.
In the latter case, depolarization must exceed a particular
threshold, meaning sodium channels depolarization must open and a
sufficient number of sodium ions must diffuse into the cell, but if
achieved, a nerve impulse will result, and a current will propagate
along the axon towards the axon terminals.
Repolarization will then occur, where voltage-gated
potassium channels allow potassium ions to exit the cell, which leads
to a brief hyperpolarization, and then everything resets to original
levels and positions. As for the impulse, when it reaches the axon
terminals and neurotransmitters are released, these enter the synaptic
space. A synapse is a junction between neurons or between a neuron
and an effector cell. This is where communication happens. These
synapses can be axodendritic, axosomatic, or axoaxonal, depending
on where the axon terminals from the presynaptic neuron connect to
the postsynaptic neuron. So that covers the basics regarding the
action potential.
We could get much more detailed and quantitative than this
brief description, and perhaps we will dig a little deeper later in the
series. But as we are currently focused on the big picture, let’s be
satisfied with this rudimentary understanding for the moment.
Now that we know a bit about the structure and function of a
neuron, let’s discuss how these are assembled to form the nervous
system. This system receives sensory input through receptors,
integrates this information to decide what to do, and then generates a
motor output, which is a response to stimulus. This is a lot more
complicated than it sounds, so the nervous system has many
components.
First, it is divided into two main parts, the central nervous
system and the peripheral nervous system. The central nervous
system consists of the brain and spinal cord, while the peripheral is
all the rest, mainly the nerves that extend all around the body, which
are bundles of axons. That means the central nervous system is the
control center, where all the sensory information is integrated, after
which a motor output is determined and implemented. The peripheral
nervous system is what allows for communication throughout the
body. The peripheral nervous system is also divided into two parts.
There is the sensory, or afferent division, which sends signals from
receptors to the central nervous system, and the motor, or efferent
division, which sends signals from the central nervous system to all
the muscles and glands to do its bidding. In turn, the motor division
is also divided into two parts. There is the somatic nervous system,
which sends signals to the skeletal muscles we consciously control,
and the autonomic nervous system, which sends signals to smooth
muscle and cardiac muscle that operates without conscious control.
And finally, the autonomic nervous system is also split into two parts.
Those are the sympathetic division and parasympathetic division,
which operate in conjunction and typically have opposing functions.
There is so much to discuss at each tier of structure that we
have just mentioned, so let’s go through these tiers one at a time,
starting with the central nervous system.
The Central Nervous System: The Brain and Spinal Cord
Professor Dave here, let’s look at some brains. We just learned
about nervous tissue, and the structure of a neuron, as well as the
divisions of the nervous system. The two main divisions are the
central nervous system and the peripheral nervous system, so let’s go
over the first of these in more detail now.
As we said, the central nervous system consists of the brain and
spinal cord. The human brain is the single most complex object in the
known universe. With a dizzying number of neuronal connections,
and the mechanism by which it produces consciousness not yet
well-understood, we will have to be satisfied with a mere
introduction to this organ. Rest assured, the brain and cognition will
be discussed in far greater detail in the upcoming biopsychology
course, but for now, we will just cover the basics.
The best way to approach learning the structure of the brain is
to examine early brain development. Within an embryo, the brain and
spinal cord begin as a single neural tube. The anterior or rostral end
begins to expand and constrictions soon demarcate the three primary
brain vesicles.
These are the prosencephalon, or forebrain, the mesencephalon,
or midbrain, and rhombencephalon, or hindbrain. The posterior or
caudal end of the neural tube will eventually become the spinal cord,
which we will discuss later. From the primary brain vesicles, the
secondary brain vesicles eventually develop. The forebrain becomes
the telencephalon, or endbrain, and the diencephalon, or interbrain.
The midbrain stays as it is, and the hindbrain becomes the
metencephalon, or afterbrain, and myelencephalon, or spinal brain.
These will then all develop further to become the regions of
the adult brain. The telencephalon sprouts two lateral regions called
cerebral hemispheres, which together form the cerebrum, and the
midbrain and hindbrain segments collectively become the brain stem.
Confined to the volume of the skull, the fast-growing brain
produces many folds, a process called gyrification, in order to best
occupy the available space. This eventually results in the
representation of the brain we are all familiar with, which we
typically divide into four main regions. Those are the cerebral
hemispheres, diencephalon, brain stem, and cerebellum. There are
also hollow cavities called ventricles, which are filled with
cerebrospinal fluid and lined with glial cells called ependymal cells.
Many of these have cilia to help circulate the fluid. The other types
of neuroglia we will find in the central nervous system include
astrocytes, with lots of branches to perform a variety of regulatory
functions, microglial cells, which monitor neuron health, and
oligodendrocytes, which form myelin sheaths.
Getting back to brain structure, the majority of the mass of the
brain sits in the cerebral hemispheres. The spongy appearance is
produced by ridges called gyri that are separated by grooves called
sulci, while deeper grooves are called fissures, like the longitudinal
fissure that separates the hemispheres
Each hemisphere is divided into five lobes, those being frontal,
parietal, temporal, occipital, and insula, the first four of which are
named after the cranial bones that are adjacent to them. We can also
describe each hemisphere as exhibiting three regions. There is a
cerebral cortex made of gray matter, consisting mainly of nerve cell
bodies and nonmyelinated fibers, an internal region of white matter,
which is a dense collection of myelinated fibers, and basal nuclei, or
regions of gray matter within the white matter.
As it’s the most complex, let’s focus on the cerebral cortex.
This is the most recently evolved section of the animal brain, and as
such it is where the conscious mind is found. It is made of gray
matter comprised of six layers of interneurons, as well as glia and
blood vessels, and there are specific regions in the cortex called
domains, which are responsible for particular motor and sensory
functions.
In other words, certain parts of the cortex are in charge of
certain aspects of bodily domains — motor and sensory functions
function. We call these motor areas, sensory areas, and association
areas. The highest mental functions, however, like memory and
language, are spread around much of the cortex, and overlap
numerous domains. In addition, each hemisphere is responsible for
the sensory and motor functions of the opposite side of the body, so
the left side of the brain controls the right side of the body, and vice
versa.
There are other aspects of the brain that are lateralized,
meaning focused more on one some functions are lateralized side of
the cortex than the other, although that whole “left-brain/right-brain”
personality type is a complete myth. Going back to the domains we
mentioned, let’s discuss the motor areas first.
First is the primary cortex. This region controls voluntary
motion, and each part of the body is relegated to a particular part of
the primary cortex. The most delicate voluntary motion occurs in the
face, tongue, and hands, so a disproportionate amount of this cortex
is devoted to those areas. The motor homunculus is an image that
depicts the human body with all of its body parts of a size that is
proportional to the quantity of neurons that control them, which is
why some features seem dramatically oversized.
Then there is the premotor cortex. This helps plan movements, and
sequences them into complex tasks, like playing a musical
instrument.
Next is Broca’s area. This controls muscles involved in speech
production, among other things. And then there is the frontal eye
field, which controls voluntary eye movement.
Moving on, let’s list the sensory areas. The primary
somatosensory cortex receives information from receptors in the skin
and other areas. This information goes to the somatosensory
association cortex where it is integrated to produce a rational
understanding of an object that is being perceived. The primary
visual cortex and visual association area receive and integrate visual
information, the primary auditory cortex and auditory association
area do the same for auditory information, the olfactory cortex
processes odors, the gustatory cortex perceives taste, the visceral
sensory area produces conscious perception of visceral sensations in
the stomach and other organs, while the vestibular cortex allows for
our perception of balance or equilibrium. There are also multimodal
association areas that send and receive information to and from
multiple areas.
These are the anterior and posterior association areas, and the
limbic association area. Moving on from the cerebral hemispheres,
we get to the diencephalon, which sits at the very center of the brain.
This consists of the thalamus, hypothalamus, and epithalamus. The
thalamus receives and directs all of the information headed to the
cerebral cortex. This means it is intimately involved with learning
and memory, among other things. The hypothalamus sits
immediately below the thalamus. This controls the autonomic
nervous system, regulates body temperature, hunger and thirst, sleep
cycles, physical response to emotions, and the endocrine system. It
also houses the pituitary gland. Lastly the epithalamus houses the
pineal gland, and helps regulate sleep. After the diencephalon we get
to the brain stem. This consists of the midbrain, pons, and medulla
oblongata, the last of which blends into the spinal cord.
Finally, we get to the cerebellum. This region, which consists
of two hemispheres, regulates muscle contraction to generate smooth,
coordinated motion. In addition, we should be aware of the structures
that protect the brain.
Meninges are connective tissue membranes that sit between the
brain and the skull. On top is dura mater, consisting of a periosteal
layer and a meningeal layer. Immediately below is arachnoid mater,
filled with blood vessels. And lastly is pia mater, made of more
delicate connective tissue. That wraps up a basic survey of the brain.
We will go into more detail at another time, for now let’s finish
off the central nervous system with a quick look at the spinal cord.
We learned about the vertebral column when we looked at the
skeletal system, and right in the middle of the column sits the spinal
cord, spanning from the base of the skull to just past the ribs. Other
than the vertebral column, the spinal cord is protected by
cerebrospinal fluid and the same meninges that we saw for the brain.
Thirty one pairs of spinal nerves attach to the cord, and we can
get a better look at the cord by examining cross sections. The gray
matter towards the center takes on a butterfly shape, made of
multipolar neurons. From these dorsal horns and ventral horns,
neurons connect with skeletal muscles and other structures around
the body, and these stem from four zones. Somatic sensory, visceral
sensory, visceral motor, and somatic motor.
Surrounding the gray matter is white matter, made of nerve
fibers that allow for communication between the cord and the brain.
These can be ascending, descending, or transverse, depending on
their direction of travel.
These participate in an incredible number of pathways that we
will investigate in more detail later.For now, let’s continue through a
survey of the branches of the nervous system.
The Peripheral Nervous System: Nerves and Sensory Organs
Professor Dave again, let’s discuss the peripheral nervous
system. The nervous system is divided into two main divisions, the
central nervous system, which consists of the brain and spinal cord,
as we just described in the previous clip, and the peripheral nervous
system.
This is the rest of it, the part that receives information from the
world outside our bodies, the information that the brain needs to be
able to tell the body what to do, so the peripheral nervous system is
the interface between a human being and its surroundings. It consists
of tiny white nerves that thread through every single part of the body,
and this includes all neural structures apart from the brain and spinal
cord.
Let’s learn a bit more about how this system is organized. The
peripheral nervous system is divided into a sensory, or afferent
division, and a motor, or efferent division. The first of these is what
gives us the ability to sense and perceive our surroundings, and the
second is what gives us the ability to physically respond to them.
Let’s talk about the sensory system first, since that’s the entry
point for any stimulus. A major component of this is called the
somatosensory system, which is comprised of all the receptors along
the exterior of your body. If something touches you, on your leg, or
arm, or the back of your head, you have this system to thank for
being able to feel it. Some processing occurs even before the signal
reaches the brain, and information about the stimulus is encoded in
the resulting nerve impulses.
These impulses are integrated at the circuit level so as to send
them to the correct area of the cerebral cortex, so that the stimulus
can be perceived properly. It is here where the information is
interpreted, its magnitude is estimated, and its quality is assessed.
Sometimes the perception of a stimulus is pain, and although
unpleasant, this is a valuable biological response that causes us to
protect ourselves from something that is actively damaging us, like a
flame, or a sharp object, by moving away from it.
There are a wide variety of sensory receptors that have the
ability to respond to stimuli. We can categorize these by the type of
stimulus they detect, their location on the body, and their structure.
Going by the first method, we can identify mechanoreceptors,
that respond to touch or pressure, thermoreceptors, that respond to
temperature change, photoreceptors, that respond to light, and
chemoreceptors, that respond to specific chemicals. There are also
nociceptors, which respond to stimuli that can damage the body, and
in turn stimulate other receptors of the varieties we mentioned to
elicit a response.
If going by location, exteroceptors respond to stimuli outside of
the body, interoceptors respond to stimuli inside the body, and
proprioceptors respond to stimuli inside the body as well, although
specifically in skeletal muscles and related connective tissue. Lastly,
if considering structure, there are many, including nonencapsulated
or encapsulated nerve endings, lamellar and bulbous corpuscles,
muscle spindles, tendon organs, and joint kinesthetic receptors.
These are each best suited for monitoring their own particular type of bones of the middle ear. This area is known as the tympanic cavity,
stimulus. which is spanned by three tiny bones called auditory ossicles. These
Of course, most of our sensory perception comes from the are the malleus, the incus, and the stapes. This leads to the internal
senses, each of which requires a complex organ to receive and ear, also known as the labyrinth, due to its complex structure. It is
transmit data. These include the eyes for vision, a nose for smelling, divided into the bony labyrinth and the membranous labyrinth. The
a tongue for tasting, and ears for hearing. Beyond this there is the main cavity in the bony labyrinth is the vestibule, which connects
sense of touch that is not as localized, and can be divided into semicircular ducts within semicircular canals. Also extending from
different modalities associated with pain, temperature, vibration, and the vestibule is the cochlea, a snail-like spiral chamber that coils
other phenomena. But looking at the other senses and their respective around the modiolus. The cavity of the cochlea is divided into three
organs, each of these structures could be discussed for several hours, chambers, or scalae. These are the scala vestibuli, scala media, and
so we will offer just a brief description here, and expand upon them scala tympani. And running through the center is the cochlear duct,
later. which houses the spiral organ, which contains cochlear hair cells, the
First, the eyes. These come with accessory structures, those receptors for hearing. The ear also contains the vestibular apparatus,
being the eyebrows, which prevent sweat from trickling into the eyes, which helps us to maintain equilibrium, or balance.
and eyelids, which cover and protect the eyes. They are activated That covers the sensory organs, so before we move forward,
reflexively to blink every few seconds, which spreads secretions let’s just mention a few things about nerves. A nerve is a bundle of
across the eyes to keep them moist. Eyelashes project from the axons enclosed by connective tissue. These are the highways that
eyelids, which are full of nerve endings, and if anything touches allow for the transfer of information around the body. Each axon
them, reflex blinking will occur. Lining the eyelids and folding back within a nerve is surrounded by some loose connective tissue called
across part of the eyeball is a mucous membrane called the an endoneurium that also encloses the Schwann cells. Another layer
conjunctiva. There is also a lacrimal apparatus, comprised of a gland of connective tissue called the perineurium binds groups of fibers
that produces a secretion that we call tears, and ducts that drain into together to form fascicles, and a tough fibrous sheath called the
the nasal cavity. Tears are meant to clean the eye and destroy epineurium encloses all the fascicles to form the nerve. Apart from
pathogens that enter the area. neurons, there are also blood vessels and lymphatic vessels in the
Six muscles control the motion of the eye, while the eyeball nerve. Most nerves are mixed nerves that contain both sensory and
itself is a hollow sphere filled with a clear fluid called the vitreous motor neurons, and thus transmit impulses both to and from the
humor. The eye has an outer fibrous layer, made of a sclera and a central nervous system. But there are also sensory nerves, that
cornea. Next is a vascular layer comprised of the choroid, ciliary transmit towards the central nervous system, and motor nerves, that
body, and iris. The iris is the colored part of the eye, with its central transmit away from the central nervous system.
pupil, which is where light enters. The inner portion of the eye is the There are twelve pairs of cranial nerves that are associated with
retina, and this is where the photoreceptors sit, which convert the brain, most of which are associated with the brain stem. These
photons into information for the brain, thus allowing us to see our run between the brain and the sensory organs, and several other
surroundings. The retina has an outer pigmented layer and an inner places. Then there are thirty one pairs of spinal nerves that go from
neural layer. The photoreceptors are of two types, rods and cones. the spinal cord to most of the areas of the body, forming complicated
The sensitive rods are for dim light and peripheral vision, while the networks called nerve plexuses, with each plexus innervating a
cones are for bright light and colors. The lens is the biconvex, particular region of the body. We will get more specific with these
transparent structure that can focus light onto the retina, and it is plexuses and nerves a bit later, as well as all of the reflexes they
comprised of the lens epithelium and lens fibers. The actual produce, when we take a look at kinesiology, which is the study of
mechanisms of refraction and phototransduction are very complex body movement. For now let’s move forward and wrap up our survey
and will not be discussed at this time, so let’s move on to the nose. of the nervous system.
The nose is full of chemoreceptors. If a substance is volatile,
meaning easily evaporated at room temperature, some of it can
interact with these receptors, allowing us to decide whether that
substance is to be avoided. The structure responsible for this is called
the olfactory epithelium, up in the roof of the nasal cavity. This
contains millions of olfactory sensory neurons, which are receptors
capable of distinguishing thousands of odors. If an odorant binds to a
receptor in the olfactory cilium membrane, an impulse is generated
that runs up to an olfactory bulb and then down an olfactory tract all
the way to the olfactory cortex in the brain, where a number of
responses may be triggered depending on the nature of the odor.
Moving on to the tongue, this is the main structure that allows
for the perception of taste, as the majority of the ten thousand or so
taste buds we possess are found on the papillae of the tongue. A taste
bud is made of fifty to a hundred cells, which are a combination of
gustatory epithelial and basal epithelial cells. The former of these are
the ones that act as chemoreceptors, while the latter act as stem cells,
replacing gustatory epithelial cells that get damaged or burnt. The
five taste sensations that can be produced by these receptors are
sweet, sour, salty, bitter, and umami, the last of which is not familiar
to everyone, but it means savory, like a steak. Most foods produce
some combination of these sensations to generate a unique
perception.
Lastly, let’s look at the ear. This is divided into an external ear,
a middle ear, and internal ear. The external ear contains the familiar
features, an aurical, helix, and lobule or earlobe. The auditory canal
also begins here, which leads up to the tympanic membrane, or
eardrum. Acoustic energy in the form of sound waves will reach the
eardrum and cause it to vibrate, and this energy is transferred to the
 MUSCLE TISSUE
Muscle Histology Explained for Beginners | Corporis
In this video, I’ll teach you the differences between smooth,
cardiac, and skeletal muscle so hopefully you can see a slide under a
microscope and know exactly which kind of muscle you’re looking
at. If you’re new to the channel, welcome, my name is Patrick and
this channel is all about anatomy and how we learn about it. As
always, I have the accompanying notes for this video linked in the
description if you want to check those out. Otherwise, let’s get
started.
Muscle is one of the four types of tissues in the human body,
and it’s the simplest one to start learning. The others, nervous,
epithelial, and connective tissue all have a ton of different
subcategorizations, but there are only three types of muscle tissue
and they all look and behave differently. But to make sense of the
microscopic anatomy, or what we deal with in the world of histology,
we need to take a step back and look at big picture anatomy where
things are more familiar.
Like you already know that the whole purpose of muscle is to
produce force. When muscle fibers contract, no matter which type of
muscle fiber it is, it will produce force. But all three types work in
slightly different ways. Skeletal muscle contracts forcefully along a
set path, so it has big bundles of parallel fibers. Meanwhile smooth
muscle just needs to squeeze, so it’s organized into broad sheets. So
as we go over these three muscle types, remember that form
influences function. Once we know that we’re looking at some kind
of muscle, it’s our job to tell what kind of muscle it is, and there are
only three types. You have cardiac muscle only in your heart
hopefully. There’s skeletal muscle in all of the muscles you’re used
to seeing, biceps, quads, etc. Finally you have Smooth Muscle which
surrounds different organs that need to constrict and expand like your
blood vessels, sphincters, or uterus.
We’ll start with skeletal muscle. This stuff is optimized for
quick and strong contractions, it’s gotta be able to move your
skeleton against gravity and heavy weights. When you actually see it
on a slide, you’ll typically see one of two perspectives. The first is
the classic transverse cross section — it’s as if you took a chicken
breast, cut it in half and looked at all the muscle fibers bundled
together, which usually looks like a bunch of circles with white
connective tissue wrapping them together. The other perspective is
from the side, or a longitudinal section. Here, you’ll see all the
parallel fibers, and if you zoom in way further, you can see the
functional unit of the muscle called the sarcomere.
Most of the time when professors present the cross section
bundle view, you’ll just need to identify the muscle fiber and the
connective tissue around it. But because a muscle is effectively
bundles of bundles of muscle cells, your task is to figure out which
layer of connective tissue and organization you’re looking at. The
whole muscle is wrapped in dense connective tissue called the
epimysium — epi- for upon, -mysium for muscle. But within the big
bulky muscle, we see clearly segregated clusters of muscle tissue
called fascicles which literally translates to bundles. And any time
you bundle something, you need something to hold it together. Each
fascicle is held together by perimysium, peri- for around.
Within each of these fascicles are individual muscle fibers,
coated with a type of loose connective tissue called the endomysium,
literally meaning within the muscle. Each muscle fiber has a cell
membrane called the sarcolemma. And this is a big stumbling block,
but sarcolemma and endomysium are not the same thing. The
sarcolemma is the cell’s membrane, it‘s part of the muscle fiber itself
while the endomysium is connective tissue, it’s a different type of
cell entirely. At this point, each of these strands is a living muscle
cell with nuclei and mitochondria and other organelles. And each
fiber has a bunch of tiny myofibrils, which are proteins, not living
things. If you have a cross sectional view of skeletal muscle, one
thing I find helpful is to take a step back and think about what you’re
looking at. Is it visible with the naked eye? That’s a muscle. Is it
magnified at a mid level and you can see individual nuclei? That’s
probably a bunch of muscle fibers surrounded by epimysium.
Every now and then, you’ll use this view to tell the difference
between Type 1 muscle, or slow twitch muscle, and Type 2 Muscle,
fast twitch. The slow twitch fibers rely on aerobic metabolism, so
they’ve got a bunch of mitochondria and myoglobin. This gives it a
distinct darker red color compared to the pale white meat, which has
less mitochondria and myoglobin. Important caveat, but you
wouldn’t be able to tell the difference between the human skeletal
muscle types with the naked eye. In reality, each of your skeletal
muscles has a mix of muscle types and we can only see the
difference with specific dyes under magnification.
Shifting perspectives, if we want to look at muscle from the
longitudinal section, we’ll see all these stripes, or striations, as well
as all these dots which are the multiple nuclei of each cell. Pro tip,
striations refer to the stripes along the muscle cell itself. When I was
a beginner, I thought the stripes being referred to were the long
parallel muscle fibers, but that’s not the case. Those striations are the
visible sarcomeres, the functional unit of muscle, we have to zoom in
a bit more. Each of these things is really tiny, only about 2 to 3
thousandths of a millimeter long, and photos of them are usually
super blurry, so your professor will usually use this diagram instead.
For this video, I’ll overlay the diagram on the microscopy so
you can see what we’re working with. From this view, we can see the
Z line, the thick boundary of each sarcomere that anchors the thin
filament, the actin. You can see the I band, which is this section in
the middle where there’s only actin, and the H zone where there’s
only myosin, the thick filament. Then there’s the A band where you
see them overlap. So for skeletal muscle, if you remember to keep in
mind which view your slide is taking, and how magnified you are,
you’ve already done most of the work.
Next up is cardiac muscle, the muscle of the heart. And we’re
gonna do the same thing here — big picture first, then we’ll see how
that plays out microscopically. The heart is just hanging out in
between the lungs, pumping blood, and its connective tissue serves to
separate the inside from the outside and keep it anchored in place. If
you were to dissect a cadaver, the first layer you’d see is the fibrous
pericardium, a layer of dense connective tissue around the whole
package. Cut that away and you’ll find a layer of serous fluid then
the epicardium. Sometimes the outer layer is referred to as the
parietal pericardium and this is the visceral pericardium, but it’s just
a different name for the epicardium. The myocardium is the muscle
tissue that we usually see histological slides of.
Finally, the innermost layer of tissue is endocardium, a
combination of epithelial cells and connective tissue that lines the
chambers of the heart. Usually that double layer pericardium mode is
analogous to putting your fist into a balloon filled with liquid. Your
hand is the heart muscle while the rubber balloon is the parietal
pericardium on the outside and the visceral pericardium, or
epicardium on the inside — the layer touching your hand heart. It’s
one continuous sac of connective tissue, but they’re clearly separated.
And of these structures, we usually focus on the myocardium
because that’s the tissue that contracts — it’s muscle. Quick
terminology note. Remember the old biological levels of
organization?
Organ systems are made of organs which are made of tissues
which are made of cells? In the case of the heart organ, when we say
myocardium we’re talking about the tissue level, but when we talk
about cardiomyocytes, we’re talking about cells. Literally heart
muscle cells. And those cells have some unique characteristics
compared to skeletal muscles. To me, they look like rectangles
instead of cylinders. And each of them usually has a single nucleus
per cell and a boatload of mitochondria. Of course, the heart isn’t just
muscle, so you’ll see some capillaries scattered around. Since each
heart chamber’s muscle cells need to contract in unison, the heart
needs to make sure there is like, zero lag in electrical impulses from
cell to cell. So these junctions, what are called intercalated discs,
allow cardiac muscle to send transmission signals super quickly
between cells.
Our final type of muscle cell is smooth muscle. They’re called
smooth because they don’t have the striations of cardiac and skeletal
muscle. Each smooth muscle cell is spindle shaped, with narrow tips
and wider cell body, and a single nucleus in each cell. So instead of
neatly organized muscle cells, you get a broad mosaic of cells
organized into sheets. Just like with cardiac muscle, smooth muscle
uses gap junctions to contract all the cells within a muscle at the
same time. That lets a smooth muscle contract at the same time, and
like cardiac muscle, smooth muscle is controlled involuntarily by the
autonomic nervous system. You’ll find smooth muscle around things
that expand and contract.
Anatomy like your blood vessels which dilate and constrict, or
the uterus, bladder, ans gastrointestinal tract, all of which have to
contract and expand at times. All in all, when you’re looking at
muscle histology, my number one tip is to take a step back and
consider how the form of each muscle cell influences function.
If you’re studying this for class and want a little extra help, I’ve
got links for my notes and example problems linked in the
description. 40% You can also check out this playlist here for more
histology help or this playlist for more musculoskeletal videos. Have
fun, be good, thanks for watching.
Types of Tissue Part 3: Muscle Tissue
Hey it’s Professor Dave, let’s talk about muscles. When you
hear the word muscle, what comes to mind? You might think about
the muscles in your legs that help you to run fast, or maybe
body-builders working on their biceps in the gym. These muscles
belong to a particular type of muscle called skeletal muscle, but we
actually have three different types of muscles in our bodies, and the
other two are not as obvious. In addition to skeletal muscle, humans
also have cardiac muscle, and smooth muscle, so let’s discuss all
three of these types right now.
First, what these different types of muscle have in common is
that they are all vital muscles are vital for movement. Muscle tissue
is predominately comprised of muscle cells, and these are highly
specialized for a process called contraction, which is what allows for
mobility. The three types of muscles are quite different in their
functions, but there are some similarities in their structure, so let’s go
through them one at a time.
Let’s start with skeletal muscle, as it’s the most abundant.
These are the muscles that cover our skeleton and give our body its
shape. The exact number of skeletal muscles in the body is not
precisely known, but it’s thought to be over six hundred and fifty.
These attach to bones either directly or via connective tissue called
tendons, and they are responsible for a variety of functions. Skeletal
muscles produce voluntary movements, which are the ones that are
under our control. This includes simple movements like extending
limbs, or more coordinated movements like running or swimming,
and they achieve this by pulling on the bones of the skeleton. Our
skeletal muscles also maintain our posture. When you’re sitting down,
it might feel as though you’re totally relaxed, but in actuality, many
of these muscles are constantly working by making small
adjustments to maintain your precise body position. Skeletal muscle
even forms sphincters in the digestive and urinary tracts to allow for
the control of actions such as swallowing and urination. Beyond
these functions, they help us maintain optimal body temperature by
releasing heat as a byproduct of muscle contraction.
So what makes up skeletal muscle? We will see that skeletal
muscle is different from the cardiac and smooth muscles that will be
examined later, in that skeletal muscle cells are multinucleated. This
means they have multiple nuclei. Cells called myoblasts, which have
one nucleus each, fuse together to form a long cylindrical
multinucleated cell called a muscle fiber, with its nuclei located at
the periphery, and this is surrounded by a sheath of connective tissue
called the endomysium.
Inside each muscle fiber are many myofibrils, and these contain
myofilaments, which are themselves organized into units called
sarcomeres. These are the contractile machinery which actively
shorten, and are therefore responsible for skeletal muscle contraction.
It is also these sarcomeres that give skeletal muscle a striated or
striped appearance, and they are the smallest functional unit in the
muscle. The myofilaments within are of two types, thinner actin
filaments and thicker myosin filaments, and the sliding of these
filaments past one another is what produces contraction, which we
will discuss in more detail later. So that’s what a muscle fiber is
made up of.
Now zooming out from there, each muscle fiber is arranged
into a bundle of muscle cells tiers of structure within skeletal muscle
called a fascicle, which is surrounded by a layer of fibrous
connective tissue called the perimysium, and multiple muscle
fascicles will in turn group together to form an even larger structure,
which is surrounded by a layer of dense irregular connective tissue
called the epimysium. This group of fascicles, along with blood
vessels, nerves, and all the tissue, comprise an entire skeletal muscle,
and these will span joints and attach to bones in at least two places.
Attachments can be direct, where the epimyseum of the muscle
is fused to the periosteum of a bone or perichondrium of a cartilage,
or it can be indirect, where the muscle’s connective tissue extends
beyond the muscle as a ropelike tendon, or a sheetlike aponeurosis,
which anchors to a skeletal element, or other muscles. For a skeletal
muscle to be able to contract, it needs to be activated or innervated
by neurons in the central nervous system, and in particular the
somatic nervous system, which essentially means the voluntary
nervous system. We will learn about the nervous system a little bit
later, for now, we can just understand that skeletal muscle is the only
muscle type under voluntary control, which means these are the
muscles that move when we decide to move them. Now let’s move
onto cardiac muscle.
Cardiac means “relating to the heart”, so as one might guess,
cardiac muscle is only found in the heart, and cardiac muscle cells
are called cardiomyocytes. There is just one role for cardiac muscle,
and that is to contract in order to push blood out of the heart to the
rest of the body, which will supply tissues around the body with vital
oxygen and nutrients. Cardiac muscle is similar to skeletal muscle in
the sense that they also have a striated appearance, which is due to
being comprised of myofibrils and sarcomeres for contractility.
However, cardiac muscle is different from skeletal muscle in other
ways.
Cardiac muscle cells are short, branched cells, which is a
structure that allows them to communicate with other cells, and they
typically have one or maybe two centrally located nuclei. In between
each cardiomyocyte there is an intercalated disc, which contains gap
junctions. These gap junctions allow the electrical stimulus that is
required for contraction to rapidly spread across the cardiac tissue, by
allowing ions to move from one cell to the next. This means that the
cardiomyocytes can contract in a coordinated fashion, allowing the
heart to work as an efficient pump. Another difference from skeletal
muscle is that we do not voluntarily contract the muscles in our heart,
meaning that we don’t consciously think about our heartbeat, it just
happens automatically. This means that our cardiac muscle is
innervated by the autonomic nervous system, which regulates the
speed at which electrical stimuli are generated by specialized cardiac
muscle cells called pacemaker cells. More on the autonomic nervous
system later. Lastly, let’s take a look at the third type of muscle
tissue, known as smooth muscle.
Smooth muscle is found in most of our organs, and it contracts
to regulate many of our bodily functions as we go about our daily
business. Smooth muscle in blood vessel walls will contract or relax
to regulate blood pressure and the distribution of blood to our
different organs. It is present in our airways to allow more or less air
to pass through. It contracts in the walls of our digestive tract to push
material along. The smooth muscle in our bladder contracts to expel
urine out of the body, and it has numerous functions in the
reproductive system, including contraction of the walls of the uterus
to push a baby out of its mother during childbirth.
Smooth muscle is typically organized into two layers. First, a
longitudinal layer, where muscle fibers run parallel to the long axis
of the organ, such that contraction will dilate and shorten the organ.
Then there is a circular layer, where muscle fibers run around the
circumference of the organ, such that contraction will constrict and
lengthen the organ. When these layers alternate contraction, it can
push material through the organ, a process called peristalsis.
Similarly to cardiac muscle, smooth muscle is innervated by the
autonomic nervous system, so it contracts and relaxes automatically.
Smooth muscle cells also have a single centrally located nucleus, but
they have a different shape than cardiomyocytes. The cells are short
but spindle-shaped, which is referred to as fusiform.
Interestingly, smooth muscle differs from skeletal muscle and
cardiac muscle because it is not striated, due to the lack of myofibrils
and sarcomeres. Their myofilaments are scattered throughout the
cytoplasm of the muscle cells instead.
So with that, we now know a bit about the three different types
of muscle tissue, those the three types of muscle tissue being skeletal
muscle, cardiac muscle, and smooth muscle. We can clearly see how
these three types are quite different in terms of functionality, but
remarkably, they are all extremely similar in structure. Now that we
have a decent grasp on the structure of a muscle, let’s talk about
muscle contraction on the molecular level, so that we can better
understand this incredible process.
The Mechanism of Muscle Contraction: Sarcomeres,
Action Potential, and the Neuromuscular Junction
Professor Dave here, let’s talk about action potential. We just
learned about the structure of muscle, like skeletal muscle, which is
what allows you to move your body around. But how exactly does
this work? If you decide to lift your arms up in the air, what is
happening on the molecular level that produces this motion? It’s not
magic, it’s just a staggering amount of chemistry, so let’s dive back
down into those muscle fibers and get a closer look.
As we recall, any skeletal muscle is made of fascicles, and each
fascicle is made of muscle fibers, which are the individual
multinucleated muscle cells. From there, we can zoom in farther on
one of these myofibrils, which are in turn comprised of myofilaments
arranged into sarcomeres, and this is the contractile unit, the one that
lets muscles do what they do. Now let’s get a closer look at these
structures, because this is where all the action happens.
Looking at a sarcomere, we can label a few regions. We can
see these darker A bands and the lighter I bands, which are aligned
and responsible for the striated appearance of skeletal muscle. Within
each A band we can find a lighter region called the H zone, and this
H zone is split down the middle by something called the M line,
made of a protein called myomesin. The I bands are also split down
the middle, by a region called the Z disc. We can identify an
individual sarcomere as being a section from one Z disc to the next,
and this is the functional unit of skeletal muscle.
If we zoom in even more, we can see the myofilaments that
make up these regions. As we discussed previously, the thick
filaments contain myosin, and these extend across the A band,
connected at the M line. The thin filaments contain actin, extending
across the I band and into the A band. There are also elastic filaments
made of titin, spanning from the Z disc to the thick filament and then
continuing on, acting as the core of the thick filament.
Now let’s zoom in even further on these two main types of
filaments. Myosin is a protein with two globular heads pointing
outwards, and a long tail which contributes to the structure of the
filament. The heads are the site of all the activity, since there are
ATP binding sites, as well as actin binding sites, meaning this is
where the thick and thin filaments will interact, by making cross
bridges. It’s important to note the lack of myosin heads in the center
of the sarcomere, just as there also are no thin filaments.
Speaking of those, as we said, they are made predominately of
actin, and two actin filaments will twist together to form the
backbone of the thin filament. Each actin subunit has an active site
where myosin can bind, and when a muscle fiber is relaxed, these are
blocked by spiraling strands of tropomyosin. There is also troponin, a
globular complex of three polypeptides, one of which binds to actin,
one of which binds to tropomyosin, and one of which binds calcium.
Beyond this, we must be aware of the sarcoplasmic reticulum, a
series of tubules that surround each myofibril. These regulate levels
of calcium, which is needed for muscle contraction, through storage
and release. This structure includes the T tubules, which sit at each A
band – I band junction, encircling each sarcomere and helping
signals reach every region of the muscle cell. Now that we’ve
examined each structural component, let’s describe how everything
works, using the sliding filament model of contraction.
This says that when the nervous system stimulates muscle
fibers, the myosin heads on the thick filaments will interact with the
binding sites on the actin subunits. These attachments will form and
break several times as the thick filaments pull the thin filaments in
towards the center of the sarcomere, thus pulling the Z discs towards
the M line. The I bands shorten, and the H zone disappears. The
overall effect is that the A bands from adjacent sarcomeres get closer
together, so the entire muscle cell will shorten. This is the
mechanism by which muscles contract.
Now the question is, how is all of this activity initiated? As we
said, this begins with a signal from the nervous system. We will
examine that system in detail a bit later, but for now we can just
examine the interface between the nervous system and a skeletal
muscle. This is called the neuromuscular junction.
Each muscle fiber has one, and these sections, called axon
terminals, are nearly touching the muscle fiber, separated only by a
thin space called the synaptic cleft, where the muscle produces
junctional folds within the postsynaptic membrane. The axon
terminal has lots of synaptic vesicles, which are like little bubbles,
containing acetylcholine, which is a neurotransmitter. When a nerve
impulse reaches the end of an axon, by a mechanism we will discuss
later, the axon terminal will release acetylcholine into the synaptic
cleft. The junctional folds contain acetylcholine receptors, so these
will bind the approaching acetylcholine, which causes a
conformational change. This protein will then act as an ion channel,
which allows for sodium ions to enter, and potassium ions to leave,
although not in equal number. Sodium will cross the membrane in
greater quantity, and this affects the membrane potential, or the
potential difference across the membrane.
For more information on electric potential, check out my
physics tutorial on this subject, otherwise, we just need to understand
that the way that positive and negative charges are distributed within
a structure can be a driving force for a particular process. Prior to the
opening of the ion channels, the inside of the cell was more
negatively charged than the outside, but as sodium ions enter,
depolarization will occur. This causes other nearby sodium channels
to open, and more sodium ions will enter, following the
electrochemical gradient. Once a particular threshold voltage is
reached, this will generate an action potential. Sodium can now
diffuse into the cell anywhere along the membrane. While this is
happening, an enzyme in the synaptic cleft called
acetylcholinesterase will break down acetylcholine, and the ion
channel closes, preventing further muscle contraction until another
nerve impulse arrives. But the action potential will continue to
propagate along the sarcolemma, and down the T tubules, which
opens up calcium ion release channels, which we will discuss in a
moment.
Finally, after the depolarization wave has completely
propagated, repolarization will begin, due to the changes in charge
density. Potassium channels will now open, and potassium ions that
are more highly concentrated within the cell will diffuse out of the
muscle fiber, which restores negative charge inside, and the
sarcolemma goes back to normal. At this point, the muscle fiber can
be stimulated again if another impulse arrives.
So we can see that charge distribution and electric potential are
key concepts here, but how does that relate to muscle contraction?
Well the action potential generated will initiate excitation-contraction
coupling. As we said, the propagation of the action potential causes a
rise in levels of calcium in the cytosol, and this is what causes the
filaments to slide.
Let’s recall that when a muscle cell is relaxed, tropomyosin
blocks the myosin binding sites on the actin subunits. But as more
and more calcium ions become present, these will bind to troponin,
and once two ions bind, it will change shape, which will push
tropomyosin off of the myosin binding sites, making them available
for cross bridge cycling, the process we mentioned before. In this
process, the myosin heads will pivot and bend, pulling the actin
filament along and using ATP in the process. The two positions
possible for the heads allow it to pull, detach, change position, bind,
pull, detach, and so forth, many times, until contraction is complete.
As calcium levels deplete, troponin returns to its original shape,
tropomyosin blocks actin’s binding sites once more, and the muscle
fiber relaxes.
There is a lot more to discuss regarding muscle contraction. We
could talk about the time frames associated with each step in muscle
contraction. We could talk about graded muscle responses, or the
differences in contraction for smooth muscle versus skeletal. But we
will have to leave that for another day, since it’s best to really
understand the basics first. Before moving forward, let’s quickly
review what we discussed.
Muscle contraction in a skeletal muscle begins when a signal
arrives at the neuromuscular junction. Acetylcholine is released,
which binds to receptors on the sarcolemma. Sodium and potassium
ions move through the ion channels, which results in a local change
in membrane voltage, also called depolarization. This initiates an
action potential, which travels across the sarcolemma in all directions,
eventually along T tubules. This is where calcium ions are released,
which interact with the myofilaments such that myosin and actin are
able to bind, and contraction begins.
Now that we understand the structure of a muscle, as well as
the mechanism by which muscles contract, let’s move forward and
look at the entire muscular system as a whole, so that we can get the
big picture regarding how we move around.