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Muscle Spindles - Motor and Sensory

So what are muscle spindles? Are they muscles that let you move, or are they
sensory organs like Merkle's discs and Pacinian Corpuscles? And for that
matter, what the heck is a spindle?
Let's get the last question out of the way first. What's a spindle? Back long
ago, before even I was a boy, if you wanted some wool yarn you didn't have
to drive all the way to the mall to get it. There were no malls. Really.
Instead, you simply sheared a sheep and took it's wool to a spinning wheel.
The spinning wheel would spin the wool into a long thread that was collected
on a spindle.
And since the neuroanatomists of old saw spindles all the time, when they
saw a spindle shaped fiber in a muscle that looked different from the rest,
they called it a muscle spindle!
Now for most of us these days, the closest we ever got to seeing a spinning
wheel was in Snow White. So this analogy isn't too helpful. So what does a
spinning wheel spindle look like? Well, it's a cylinder that's tapered at the
ends. In other words, it looks like a ... muscle spindle.
So much for analogies.

Now that that's settled, what about the other issue. Earlier this quarter, I said
that muscle spindles were somatosensory receptors, well then they must be
sensory receptors. Right?
But why call them muscles? Muscles aren't sensory organs, right? Correct,
muscle aren't. But muscle spindles are a mixture of tissues, both sensory
neurons and muscle. They are sensory receptors that can contract and
Muscle spindles are made of thin cells called muscle fibers, just like the rest
of the muscle that make up most of the muscle (called extrinsic muscle
fibers). Like all muscle fibers, there are areas in the muscle spindle fibers that
are striated. They have Z lines in this striated regions, there's actin and

myosin there, and whey calcium flows... contraction happens

(You remember Z lines, right?).
Still, there is an important difference between the muscle
spindle fibers and extrinsic muscle fibers. Only the ends of the
muscle spindle fibers can contract. The central 4/5s or so
cannot contract at all. This is the sensory area of the fiber
which, like a Merkle's disc is connected to a primary afferent
Actually there are two types of muscle spindle fibers, not just
one, the nuclear bag fibers and the nuclear chain fibers.
The difference in the anatomy of the fibers is pretty subtle and
based on the location of the nuclei. Oh, just in case you were
wondering, these nuclear fibers are not radioactive.
Muscle fibers, unlike neurons, have more than one nuclei. They
are multinucleated. In this diagram, the nuclei are the little
round balls inside the soma of the muscle fibers. On the left is a
chain fiber. In this fiber, the nuclei spread out along the fiber
like a chain. A nuclear chain!
For the nuclear bag fiber, the nuclei hang out in the center of the fiber, which
bulges out into a bag. A nuclear bag!

Notice that at the end of the fibers are some red stripes. This represents the
striated, contractile part of these fibers. Since such a small part of the fiber
can contract, your never going to be able to lift a Buick with your muscle
spindles. Or for that matter, you aren't going to even lift you arm with them.
As muscles, they are wimps.
So what good are they?

The long non-contractile region is connected to primary afferent fibers. You

know, the ones with cells bodies in the Dorsal Root Ganglia. The ones that
terminate in ... humm. I think I feel a question coming on.
Where would the primary afferent fiber that is attached to a muscle spindle
located in the gastronemius muscle terminate?
The Dorsal Horn
Nucleus Gracilis

Nucleus Gracilis - One of Many Targets

Very good. You remembered that muscle spindles are one of the sensory
receptors that encode proprioception. And their primary afferent fibers form
part of the DC/ML that runs without crossing or synapsing all the way to the
You did remember this. Right?
Anyway, the primary afferent fibers from muscle spindles go not to just one
place, but several. These include:
1. Nucleus gracilis or cuneatis
2. Lamina 7 at the base of the dorsal horn (e.g. Clark's) and nucleus in the medulla we will
soon meet called the external cuneate nucleus. These nuclei send proprioceptive
information to the cerebellum. The unconscious proprioception pathway.
3. Lamina nine of the spinal cord. That's where the alpha motor neurons live.
So these primary afferent fibers are pretty busy fibers. And as you might
imagine there are more than on type of fiber. Actually two types --The Ia and
II fibers. These fibers begin at specialized connections to the nuclear chain
and bag fibers that are called the annulospiral endings and flower spray
As you might imagine, these "endings" get their name from there
appearance. Well, at least how they appeared to some ancient
neuroanatomists who may have spent a little too long at the microscope. I
say this because they called these fiber endings. These aren't nerve endings.
They are nerve beginnings. Sensory nerve receptors.

Here are our muscle fibers again, all wrapped up by the

annulospiral "endings". The green fibers are the beginning of
type Ia fibers that will travel all the way back to the central
nervous system.
The purple fiber looks like a flower spray doesn't it? These
"endings" form the beginning of the type II fibers which also
travel far and wide in the CNS. Although most of the type II
fibers originate in the chain fibers, some do originate in the bag
fibers too.
The cell bodies of these fibers are located in the Dorsal Root
Ganglia. Such a long way to go for such small fibers. But then
that's what neurons are all about. Sending information over
long distances.
So, what about the gamma efferent fibers? Where do they fit in?
They Don't Fit In
At the Ends

Catching a Gamma with Your Tail

The contractile region at the end is the target for the axons of the gamma
motor neurons. These neurons are located far away in the ventral horn. In
good old lamina 9 along with the alpha motor neurons.
Notice the red fiber. To any color blind souls, it's the one in the middle. This
fiber really is the ending of an axon. It's the end of the axon of a gamma
motor neuron. It's synapsing on the contractile portion of the muscle fibers
and when it releases it's neurotransmitter, the muscle spindles will contract.
Just like the extrinsic muscle fibers. But don't expect to see any movement
from this. Well, at least not at first.
So, that's the anatomy of the muscle spindles. Your textbook has a nice
chapter on this anatomy, so if you are confused, feel free to read.
In summary, the muscle spindle is a muscle fiber, but it is also a sensory
receptor. It is innervated by motor fibers, the gamma motor neuron axons
and contracts when these fibers are active, but can't produce much force. It

is also innervated by sensory fibers, like all good sensory

receptors. These are the type Ia and type II fibers.

The Monosynaptic Reflex - Not a Deep Tendon

Why do health care professionals whack us with hammers? Whack, whack,
whack! Well, we will get to the motive soon, but now we need to review the
anatomy of the monosynaptic reflex, often somewhat unfortunately
referred to by clinicians as the deep tendon reflex.
Key features are:

The Muscle Spindle

The Ia afferent fiber (I'm ignoring the II fibers, sorry)
The Alpha Motor Neuron
The Extrinsic Muscle Fibers

Oops. Did I forget to mention the gamma motor neurons.

How about a question.
Striking the Patella Tendon with a hammer results in activation of:
The Golgi Tendon Organ
The Muscle Spindle

The Monosynaptic Reflex - Sensory in Motor Out

Very good. These Ia fibers go to many places, but right now we want to focus
on the termination in the spinal cord, ventral horn, lamina 9. You can't get too
much more specific than that.

The Ia fibers make a direct synaptic contact with the alpha motor neurons.
This is a very powerful contact that has a powerful excitatory effect on these
ventral horn cells. This is the sensory part of the reflex, and all reflexes have
a sensory part. No sensory input, no reflex. But to complete the reflex, there
is a missing axon. I wonder where?

Ah ha! The missing fiber. The motor fiber. Reflexes consist of two parts a
sensory and a motor part, and now you see the motor.
The axon of the alpha neuron (I won't insult you intelligence by mentioning
that this is the blue fiber) travels back to the muscle and synapses not on the
muscle spindle, but on the other muscle fibers in the muscle. The ones that
connect to the tendons outside of the muscle -- the extrinsic muscle fibers
(muscle spindles are sometimes called intrinsic muscle fibers). In fact
most alpha neuron axons terminate on many extrinsic muscle fibers and can

cause quite a lot of contraction of these fibers. So movement occurs. The

reflex is complete. Sensory in motor out.
A few other points and we are finished with the anatomy and basic
physiology of the MSR. First, you probably noticed that the motor fiber
returned to the same muscle from which the Ia fiber emerged. This is called
the homonymous muscle, where homonymous, in Latin, means.

The Same Thing

Homonymous is a word that will keep popping up I'm afraid. Clinicians love
Latin. I don't know why. In any case it's a great word. Impressive at parties. I
can almost hear you at the next party saying, "What! Pin the tail on the
donkey again! Why do we always do the homonymous thing at every party?!"
A homonymous muscle is necessarily a synergistic muscle, that is, a muscle
that has the same action. While the Ia fibers do indeed synapse on alpha
motor neurons of homonymous synergists, they also synapse on the alpha
motor neurons of heteronymous synergists. That is, muscles that are
different, but have the same action. So keep in mind that this simple picture
of a monosynaptic reflex is just that, simplistic. The Ia fibers innervate much
more than just one alpha motor neuron.

Nevertheless, the Ia fiber doesn't innervate all alpha motor neurons. It

doesn't, for instance, synapse on the alpha motor neurons that project to
antagonist muscles. Multiple alpha motor neurons are activated, but they all
have one common purpose. To resist the stretch.
So, we have MSRs. Often called DTRs by rather confused practitioners. When
you stretch a muscle, the muscle contracts. Strongly!
Well, that's nice, but why in the world do we have such a resistive reflex? Or
in other words, did the monosynaptic reflex evolve to provide health
care workers with a handy way to test for lower motor neuron

A Synergist is not always the Same

It means having the same action, but not literally the same.

The Monosynaptic Reflex Servomotor Control

Here we see the components of the MSR, the muscle spindle, the Ia afferent
fiber, the alpha motor neuron and it's axon and the extrinsic motor fibers all
happily in homeostasis, holding up a 10 pound weight acting across a joint.
Since there is a force stretching the extrinsic muscle fibers and the muscle
spindle is attached to these fibers, the muscle spindle is stretched. And the Ia
fibers are activated. The Ia fibers synapse on the alpha motor neurons, so
these cells are activated and produce action potentials that race down their
axons and cause the extrinsic muscle fibers to contract. Just enough to resist
the 10 pound force.

And this keeps the joint from moving.

Gamma Activation- A Little Pop to Get You Going

So what do Gamma motor neurons do, and what do they have to do with the
MSR? If I said that the gamma motor neurons, which we will now simply call
gammas, adjust the contractile state of the muscle spindle to extend its
dynamic range to the full extent of the range of motion of the extrinsic
muscle, would you have any idea what I mean?
If so, you can probably skip the rest of this lecture. In fact you may want to
consider leaving Physical Therapy altogether for a promising career in
physiology ... or in government service.

Otherwise, let's put the gammas in the picture.

Here's our gamma motor neuron, green for gamma. It's smaller than the
alpha motor neuron which is drawn in blue because... I don't know. Notice
that the Ia fiber terminates on the alphas not the gammas. Notice that the
alpha's axon terminates on the extrinsic muscle while the gamma terminates
on the contractile part of the muscle spindle. What would happen if the
gamma motor neuron fired, and the alpha motor neuron didn't?
The Muscle Spindle would Decrease in Length
The Muscle Spindle would Stay the same Length

Isometric Contraction
The muscle spindle stays the same length since the extrinsic muscle fibers
don't contract; an isometric contraction. But something has to change

since force has increased, and what increases is the tension in the muscle
spindle. The central sensory region gets stretched.

Much to the displeasure of the now rather tense nuclear bag and chain fibers
in this unhappy spindle. And we know what an increase in tension in the
sensory part of the muscle spindle does to the activity of the Ia fibers. Right?
The activation of the gamma motor neuron converts the muscle spindle from
a laid back sensory receptor to a lean, mean sensing machine. Now, any
change in the length of the external muscle will change the tension in the
muscle spindle and alter the activity of the Ia afferent fiber. Or in other
"the gamma motor neurons adjust the contractile state of the muscle spindle
to extend its dynamic range to the full extent of the range of motion of the
extrinsic muscle".

Gamma Activation- Tensing the Spindle

Okay, here's that sentence again.
The gamma motor neurons, which we will now simply call gammas,
adjust the contractile state of the muscle spindle to extend its
dynamic range to the full extent of the range of motion of the
extrinsic muscle.
Let's see if we can turn this into something a bit more understandable.
First, it's important that the muscle spindle be able to detect a change in
length of the extrinsic muscle, the big muscle, whether it is extended or
flexed. That's their job as sensory receptors after all. When muscles are
extended, the muscle spindles are also extended, and have an easy time
detecting small changes in muscle length, and therefore limb movement or
lack thereof.

But when the limbs are flexed, the muscle spindles shorten. This makes it
more difficult for the spindles to detect small changes in the length of the
muscle. Nevertheless, you still need to be able to regulate limb postition
when the limb is flexed, so something must be done to tighten up the
Physiologists long ago showed the excellent sensitivity of muscle spindles
long ago by taking out a muscle and hooking it up to a device that allowed
them to stretch the muscle over and over while measuring the activity in the
Ia fibers. While I thought about requiring you all to do this experiment
yourselves, thanks to the miracle of the web, we can simply view the
experiment which I conducted on one of the muscles of a
former neuroanatomy student, Raymond E. Kaygel.
Raymond didn't quite have a passing average last winter, so
he asked if there was anything he could do to pass the
course. Anything. Perhaps a special project.
I was happy to oblige.
Here you can see one of Ramon's muscles. I think it was a
hamstring muscle but I forget. The muscle is stretched by the
stimulating machine nearly to its maximum length, which
would normally happen if the leg was fully extended. For hour
after hour, the stimulating machine stretched the muscle by a
small amount. The Ia fiber detected the change in length
faithfully, firing the most when the muscle was most
stretched. Unfortunately, physiologists don't deal with faith
very well. Instead, they use graphs. Here's the graph of
Raymond's Extended Muscle Experiment.

As the machine's arm went up and down, the extrinsic muscle's tension was
decreased and increased, respectively. The blue wavy line shows the tension
of the muscle, but strangely up means less tension. I did this so that it would
match the animation, and to confuse any physiologists who might be
As the blue line drops, the muscle gets longer, and the tension increases. And
look what else happens.
The line at the bottom of the graph, which looks to normal people like, well,
not much, looks to the physiologist like the activity in the Ia fiber. Each
vertical line is one action potential. And the number of lines, the frequency of
the action potentials, increases when the muscle is stretched. The activity in
the Ia fiber faithfully represents the tension in the muscle, and thus the

length of the muscle, and thus the position of the machine's

arm. There's that faith stuff again. Propriofaith.
So when the limb is extended, the muscle spindle accurately
determines limb position. Now what happens if the limb was
The Muscle Spindle Will Still Be Faithful
The Muscle Spindle Will Be A Jerk

Gamma Activation - Keeping the

Spindle At Home

When the machine's arm is flexed, the extrinsic muscle is much shorter and
compressed. And so is the muscle spindle inside since it's fiber run in parallel
to the extrinsic fibers and are attached to them.
Up and down up and down, goes the machine. The arm is flexing and
extending by the same amount as it was in the previous experiment. So what
does the activity in the Ia fibers do?

No, it's not a misprint. The Ia fibers are silent. Even though the muscle
spindle is being stretched over and over, now it's fat and happy. Floppy,
flaccid, lax, limp, detumescent. (Okay, not exactly detumescent, but I put it in
for emphasis)

So, what's wrong with the little guy? Why won't he fire off his Ias? Up and
down, up and down for hours, and ... nothing!

Well, in case you haven't guessed, the muscle spindle can't reach ...
threshold. (Why, what were you thinking?) The tension on the sensory region
of the muscle spindle is increased when the extrinsic muscle is stretched, but
not to the point of ... emission ... of an action potential. So, currently, the
muscle spindle cannot provide faithful information about the position of the
limb as it moves up and down in this flexed position. And that's no good.
Fortunately for the nervous system, the spinal cord has a way to adjust the
sensitivity of the muscle spindle. A way to make it sensitive to limb
movement whether the limb is flexed, extended or whatever. The nervous
system can adjust the muscle spindle so that it is sensitive throughout the
entire range of motion of the limb. And we call that:

Gamma Activation - Increasing the Dynamic

The following is a transcript from the Society for Neurophysiologists
conference on the Monosynaptic Reflex, held in 1950. The speaker is the
eminent and very aged Dr. Charles Tonofsherry.
Dr. Tonofsherry concludes ...
In my final experiment, I will demonstrate the physiological effects of gamma
stimulation on the dynamic range of the muscle spindles and their
dependencies, the Ia afferent fibers. In the previous experiment, we saw that
the muscle spindles were only, fai..., I mean, reliable, when the limb was in a
extended position. In the flexed position, the Ia fibers were silent, reflecting
the flaccid state of the muscle spindle fibers.
Now, if my assistant will activate the electrical stimulator that
is attached to the gamma motor neuron axons, we will see if
this changes the output of this system. You will note that the
mechanical stimulation parameters are identical to the
previous experiment. The limb is in exactly the same flexed
position, the extrinsic muscles are compressed, and prior to
activating the electrical stimulator, the muscle spindle is limp
as angel hair spaghetti after a hour's boil.
With stimulation of the gamma motor neuron axons, however,
the tension in the sensory portion of the spindle is increased
secondary to the isometric nature of the contraction. This

increase in tension in the sensory region has profound effects on the activity
of the Ia afferent fibers as the muscle is repeatedly stretched.

As you can see in the oscillogram, the activation of the gamma motor
neurons (bottom trace) restores the sensitivity to the muscle spindle. The
muscle spindle now provides a completely, ummm, accurate representation
of the limb position in this flexed state. Thus the dynamic range of the
muscle spindle is extended to the full range of motion of the limb by gamma
Thank you. I will take questions now.

Dr. Granite asks: "So one might say that the gamma motor neuron evolved to
compensate for the demands placed on the MSR as the limb mobility
Dr. Tonofsherry's response: "An interesting question. Yes, I suppose that is
possible.As the range of motion increased and the precision of movement
became more exacting, the gamma motor neurons may have developed to
allow the MSR to continue to function."
Dr. Poorluck asks: "So even though the gamma motor neurons aren't actually
part of the monosynaptic reflex, without them, the reflex would only work
when the limbs were extended."
Dr. Tonofsherry's response: "Yes, that would be the case."
Dr. Liceberg ask: "Yeah, but isn't this supposed to be a conference on the
Monosynaptic Reflex? Why are we talking Gammas here?"

Dr. Tonofsherry's response: "Well, ah, I thought you would find my results
interesting in light of the relation between the dynamic range of the spindle
and the MSR."
Dr. Icicles asks: "Gee, Chuck. Wasn't it pretty obvious that gamma stimulation
would increase the dynamic range of the spindles? I mean, like, duh!"
Dr. Tonofsherry's response: " Well John, in the word's of my ancestors, "Yo
lion. Eat this!"
Following this comment a brouhaha ensued which continued for some time
until a young physiologist named, Dr. Mirthton, suddenly shouted,"Hey, guys
stop fighting! Let's be happy. I just thought of a way the gamma motor
neuron might initiate movement via the MSR!"
He continued. "Although activation of the gamma motor neuron and the
concomitant contraction of the spindle produces insufficient force to move
the limb, the gamma motor neuron might provide an easily damped
servocontrol mechanism with intrinsic load compensation through which
gamma activation leads to a cascade of events encompassed within the
structural components of the MSR! In essence, a loop. The Gamma Loop!"
The room became quiet. Stunned by the eloquence of his words. A mummer
was heard throughout the room which grew quickly to become a unified
expression of the sentiment felt through this august assembly. The murmur
was of one word. And that word was....

The Gamma Loop- Round and Round We Go

Isn't the Web great?! While sitting in one place you can surf to conferences
about almost anything but current and past. Even a conference on the
monosynaptic reflex! I know the conference that I linked you all to was pretty
old, but hey, everything we do in this course is pretty old knowledge.
Okay, back to your present professor. What is the gamma loop? And how
does this help answer the discussion question we are working on, that is, why
do we have a MSR? An also we finally get to show how the MSR is to involved
in spasticity. And that is certainly important to clinicians.

The Gamma Loop idea is pretty simple. If the gamma

motor neuron is activated, then this would increase the
tension in the muscle spindle and, therefore, increase
the activity in the Ia afferent fibers. The increase in Ia
activity increases the activity of the alpha motor
neurons, and this then causes the extrinsic muscle fibers
to contract.
So, movement of a limb could be produced by
contraction of a muscle spindle. Just not directly. The gamma motor neurons
and muscle spindles produce movement by activating the MSR.

Perhaps you have noticed that I have studiously avoided the question of what
turns on gamma motor neurons? What synapses on the gamma motor
neuron? Well, now it's time for you to find out why I've avoided this important
question. Basically, I don't know.
Gamma motor neurons may be spontaneously active. They may just fire, fire,
fire when left on their own. Local spinal interneurons or descending
projections from the brainstem and cortex may activate them. I just don't
know. What I do know is that gamma motor neurons are busy little cells, and
a very important function of descending connections form the cortex and
brainstem is to slow them down. To inhibit the Gamma Loop.
Inhibiting the Gamma Loop
Muscle normally has tone, a state of tonic contraction. The gamma loop is
very important in the production of this tone. If the gamma motor neurons
are active, and they are active little buggers, they tense the spindles, fire the
Ias and keep the alpha motor neurons busy. But if the gamma motor neurons
became overactive, the gamma loop would go wild. The muscle spindles
would contract and contract and the Ia fibers would continue to fire driving
the alpha motor neurons into a frenzy of activity resulting in extreme
contraction of the extrinsic mucle fibers. If left alone, the Gamma Loop would
go wild, driving tone through the roof.
Thus, something has to shut them down. And descending projections do just
that, they inhibit the gamma loop.
The Gamma Loop Goes Wild
Damage to the descending pathways produces spasticity -- a chronic increase
in tone in the physiological extensor muscles. Although the exact mechanism
of spasticity is not known, here's a plausible sequence of events that might
just do the trick.

1. Damage to descending spinal projections results in a loss of inhibition of the gamma

motor neurons.
2. The gamma motor neurons increase firing to maximum levels.
3. This increase in gamma activity puts the squeeze on the muscle spindles.
4. Therefore, the Ia fibers go nuts.
5. Thus, driving up the activity of the alpha motor neurons.
6. And causing a massive increase in tone.
While this certainly isn't the only sequence of events involved in the
production of spasticity, the "Gamma Loop Gone Mad" hypothesis does help
to explain what happens with a fairly common surgical treatment for
A surgeon can reduce spasticity by severing:
The Dorsal Roots
The Ventral Roots

The Dorsal Rhizotomy

Severing the ventral roots would work of course, but with a greater functional
lose than the spasticity produces. A flaccid muscle cannot help with
movement. Severing the dorsal roots, however, can reduce the spasticity
dramatically and allow the patient to regain use of the muscle. With proper
physical therapy of course.
Cutting the dorsal root opens the gamma loop, the gamma motor neurons
are still hyperactive after the loss of descending inhibition, and the muscle
spindles are contracted as tight as a tin drum, but the Ia fibers are severed,
so the excess activity in the gamma motor neurons is not relayed to the
alpha motor neurons. Tone is reduced.
But not without a cost.
The total dorsal rhizotomy has a side effect of:

GTOs Do What?

There are several ideas on what Golgi Tendon Organs do since they have
nothing at all to do with the monosynaptic reflex. We know that they can't be
part of this reflex because:
1. They don't synapse on alpha motor neurons.
2. When stimulated they reduce the activity of alpha motor neurons through inhibitory
But what do they do?
The Clasp Knife Reflex
One popular idea is that the Golgi Tendon Organ has a protective function.
When weight lifters attempt to lift so much weight that they risk damaging
their muscle, they often report that the force they are producing suddenly
drops, as does the weight, hopefully not on their foot.
Something caused their alpha motor neurons to shut down when the tension
across the muscle exceeded a high threshold. Since the Golgi Tendon Organ
senses tension, it has long been though to be responsible for this protective
release of tension.
Unfortunately, when the actual threshold of the GTO was measured, it was
found to be very low. The Ib fibers that carry the GTO response would reach
maximum firing rates long before extreme tensions were reached. Thus the
GTO couldn't produce this effect.
In some neurological patients, an abnormal reflex called the clasp knife reflex
occurs at much lower levels of muscle tension. The patient resists the
extension of a limb by the clinician. The patient's muscle tension builds but
then suddenly releases as the limb is stretched out. The feels to some
clinicians like the resistance of a clasp-type knife as you open it.
An overly active GTO might explain this abnormal reflex, but their seems to
be little clear data on this.
So, what do GTOs do? I don't know. But I do know that they don't produce the
Deep Tendon Reflex.