In this course, students learn to recognize and to apply the basic concepts that govern integrated body function (as an intact organism) in the body's nine organ systems.
Action Potentials
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Из курса от партнера Duke University
Introductory Human Physiology
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The Nervous System
We hope you are enjoying the course! Last week's lectures can be challenging because we introduce many concepts that may be new to you. This module will allow you to apply some of the concepts that you learned last week and provide you with more concrete examples. In this module we will begin our tour of the various organ systems with the nervous system.
We start by considering the function of the individual cells (neurons) and then how they interact as an integrative system. The nervous system provides rapid communication throughout the body coordinating the actions of trillions of cells. It responds to internal changes to the body as well as to changes in our external environment. This is a busy week. The things to do this week are to watch the 5 videos, to answer the in-video questions, to read the notes, and to complete the Nervous System problem set. We suggest that you read the notes, watch the videos, and answer the in-video questions before you start on the problem sets. The problem sets require you to apply your knowledge from the lectures so it is best to be fairly familiar with the material before tackling them. The problem sets are not graded, and there is no due-date for them.
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Jennifer CarbreyAssistant Research Professor
Department of Cell Biology
Emma JakoiAssociate Research Professor
Department of Cell Biology
Welcome back.
We're going to be talking this session about action potentials.
So these are going to be the action potentials are going to be
another major way that neurons are going to communicate.
And we'll talk more about specifically, they have a role in communication but
we're going to start talking about first the ion channels that are important and
that are necessary to have in action potential.
So an action potential is going to be a very specific kind
of change in membrane potential.
And the reason why action potentials happen instead of a graded potential is
because of the types of channels that are present where we have action potentials.
So instead of being channels like the ligand gated ion channels we talked about
with graded potentials.
For action potentials, you must have voltage-gated channels.
And so, we're going to talk about the voltage-gated sodium and
potassium channels that are going to be so common in forming an action potential.
So, the fact that their voltage gated means that
a depolarization of the membrane.
The membrane becoming more positive, is what gates them,
which what causes them to open, and
that's what's shown in this fist spot where they start off closed.
And then we have depolarization, and that causes them to open.
However, keep in mind the sodium channel is always fast to make
these changes to its structure, and the potassium channels slow.
So the sodium is going to be fast and the potassium are going to be slow.
And so, the sodium is going to open first and
the sodium channels are going to allow sodium to rush into the cells, so
that's what happening here in this next spot.
Whereas, the potassium channels they've still been stimulated by
that depolarization.
However, they just haven't opened yet.
After a little bit of time, then what happens is
the potassium channels have finally opened in response to the stimulus.
But in that same amount of time, you see this little structure
has now closed or filled in the opening of the sodium channel.
And the reason why it has done that is because now it has moved to being in
an inactivated state.
Which means that it not only is no longer moving sodium ions in and
out of the cell but it also can't be opened.
That's why it's called inactivated, it's different from being closed.
So at this third time point,
then we only have potassium moving out of the cell.
As potassium's moving out of the cell, then that means that the membrane
potential is starting to head towards the equilibrium potential for potassium.
And it's that repolarization of the cell that then causes the closing
of both the potassium channel and the closing of the sodium channel.
Now, we know it was in the inactivated state and
it wasn't moving sodium ions then.
But now since it's in the closed state,
then it can at least be opened with another change in the voltage.
So this properties of these channels are what form and
give the properties of the action potential.
And this property of the sodium channel being inactivated until the membrane
potential comes back down to rest.
Is also a very important aspect of the action potential and
we'll talk about why it's important.
And it's responsible for
the property that we call the refractory period of the action potential
when there's a period while those sodium channels are inactivated.
That no matter what you do to stimulate the membrane, they're not going to open,
and you're not going to have another action potential and
we'll see why that's important soon.
So, let's put together these properties of these channels and
what that's going to mean for the membrane potential when an action potential occurs.
So, in our state 1 here, we are at rest and
then, you can see the next step is that we have some stimuli.
And these stimuli are very often going to be graded potentials.
So that say that on our membrane we're releasing a little bit of neurotransmitter
that's binding to some sodium channels, causing them to open.
Which then sodium is going to rush in because of its chemical and
electrical gradient that's going to cause a little bit of increase in
the memory potential in that spot.
And this diagram showing two sub-threshold stimuli where they don't go high enough,
they don't depolarize the membrane enough, to open these voltage-gated channels.
And so they just die out as grated potentials do and then nothing happens.
But if we get a stimulus that’s great enough,
that gets the membrane to threshold,
that is when the sodium and potassium channels will open.
And that's when we'll get to stage 2, which we’ll remember we said both channels
are going to be activated by this reaching threshold.
But that sodium channels are going to open really quickly and so,
that means in stage 2, we have only sodium channels open.
So sodium's going to rush in, that's going to make the membrane
more positive or the membrane potential more positive,
which then is going to open more sodium channels.
And so we're going to have a positive feedback cycle going on where we bring
in more and more sodium and open up more and more More sodium channels.
Then the membrane potential is going to head towards the equilibrium potential for
sodium because we have only sodium channels open.
So that's what's happening in stage 2,
we're getting a huge depolarization and overshoot.
In stage 3 now, we've had a little bit of time pass and so
we've got 2 things happening.
One is now there's enough time that the potassium channels have opened and there's
been enough time that now the sodium channels are in the inactivated state.
Which, remember, means they're no longer moving sodium and
they can't be opened right now.
So, if we've got only potassium channels open, then membrane
potential is going to head towards the equilibrium potential for potassium.
Those potassiums are going to rush out of the cell based on both the chemical and
electrical gradient at this point.
And so, we're going to have a big repolarization of the membrane in stage 3.
In stage 4, now we've come back down to rest and
so, now we're starting that reset of the channels.
Where now the sodium channel is going to be able to close,
which is what's shown here.
So, it could, in theory, be opened at this point,
the potassium channels are going to be slow to close.
And so, they're still going to stay open and so, that's why we're
going to go below And get to the equilibrium potential for potassium.
But then in stage five,
finally the potassium channels have caught up and now they're closed.
And so then,
we're going to get back to rest through the actions of the sodium-potassium ATPs
which is going to get our gradients back in order where they should be.
Keep in mind, with all of these big changes in the voltage of the membrane,
we're moving very few ions and so our chemical gradients,
our concentration gradients are not changing appreciably.
Even though we're having these huge swings in the membrane potential.
So based on this, you can see how this is different from a graded potential.
If you don't reach threshold, nothing happens, but as soon as you reach
threshold, you're going to have the same response no matter what.
You're going to have an action potential, so it's an all or nothing phenomenon.
And we'll talk many times about this property and
what that means for what the nervous system can accomplish.
And we'll also talk about how this is not going to decay over a distance
and that's what shown here.
So, remember we've talked a little about or we're going to be
talking about the fact that these voltage gated channels that
mean that an action potential can occur are going to be in an axon.
So here's a neuron with an axon and we're saying that right here,
we're having an action potential and that it's going to travel down the axon.
And so, at this point where this action potentials occurring,
sodium is rushing in and it's going to then
depolarize the membrane in the neighboring area.
Such that it's going to get that part of the axon above threshold and
so it will have an action potential.
So we had an action potential right here and
then it stimulated this area right here to have an action potential.
And so that's what's happening right here, which is then has stimulated this
area to have an action potential, so we've got sodium rushing in right here.
Now, you can see when we're having action potential right here,
we have sodium rushing in and it's diffusing in all directions.
However, this area right next door is still having its action potential and
so, since it's still highly depolarized.
Then those sodium channels are still in the inactivated state and so
we can't have an action potential right here where we're still having one.
We can only have one downstream in this direction and so in that way,
once we get started here we can only move the action potential down the axon.
Now, if we started in the middle that would be a different issue but
since we always start at the initial segment for the most part.
Then we're going to have a unidirectional propagation of the action potential and
it's because of that refractory period of the sodium channels when
they're inactivated.
So, this is where we're going to talk about the different parts of the neuron
and what channels are expressed where.
So, really to make things simple, you can think about the fact that the dendrites
and the cell bodies are going to tend to have ligand gated ion channels.
Which means these are going to the parts of the neuron
that are going to be responsive to neurotransmitter.
And so, they're going to be the parts that are going to
have a synapse from another neuron and they're going to release neurotransmitter.
And the dendrites in the cell body are going to have the ion
channel that respond to those neurotransmitters and
that means that the dendrites and cell body will have graded potentials.
The axon is going to be different, it's going to be where
you have the concentration of the voltage gated channels and so
that means that's where you're going to have the action potential.
And in specifically right here in the initial site
where that axon is meeting the cell body is where you're going to have a super
high concentration of this voltage gated channels.
So that if whatever's happening with the dendrites or
the cell body with the forming graded potentials is
great enough to get this little spot above threshold.
Now remember, they're going to decay over time and space and
be proportional to the stimulus but
if they're great enough, then we will cause an action potential.
So in this way, you can think of the cell body and
the dendrites as kind of the place that's summed,
all those signals are summed through graded potentials.
And that if the stimulus is high enough, then the initial
segment will take that information and decide whether or
not to have an action potential and we'll talk about this much more right now.
So this is the idea of having multiple synapses so
multiple inputs through greater potentials.
And imagining that we're measuring the membrane potential
at the axon initial segment.
So that we know if a membrane potential goes above fresh hold at the axon
initial segment.
We're going to send an axon that tend to down the axon.
So here we have synapse 1,
where when that neuron releases
a little bit of neurotransmitter.
It causes a little bit of depolarization of the membrane that
reaches the initial acts that acts on initial segment,
so let's say it causes some sodium to enter the neuron.
Synapse 2, when it fires, it, in this case, causes an even larger stimulus,
so maybe it's closer to the axon initial segment.
So that there's more of those positive charges that come in when those sodium
channels open, reaches the Initial segment to make a larger depolarization there.
Then we have synapse 3,
which is actually causing a hyper-polarization of the membrane,
so perhaps it's causing ion channels to open that.
Let chloride into the cell or let potassium out of the cell,
so that we get a hyperpolarization when synapse 3 fires.
Since these graded potentials are additive,
then they are additive when you look at the axon initial segment.
So if we have firing of one and two at the same time, we get a larger effect and we
get closer to threshold than we did when either one of them fired by themselves.
And we can also have and additive affect with a positive or an excitatory and
an inhibitory synapse, so that they kind of cancel each other out.
And then, remember we can also have repeated stimuli, so
we might not just have a greater stimulus.
But very often the nervous system is going to signal by having
repeated actions potential, so repeated stimuli.
And when that happens, then they can be additive and then,
in that way, get to threshold so that we have an action potential.
So this is what we're talking about when we're talking about the axon
initial segment integrating information.
It's allowing these gray potentials to be additive, and
if they're strong enough, to cause an action potential.
The last thing we're going to talk about with action potentials
is the role of myelination.
So we talked about how and myelinate portions of the axons and
then leave a space, which we call the node of Ranvier, which is
a bare spot of the membrane that isn't electrically insulated by the myelin.
And has a large concentration of these voltage gated channels.
And so what that means is if we have
an action potential, for instance, let's say it's happening at this node,
the sodium is going to rush in, which positive charges are going to rush in.
Because of that myelination and not allowing leak
of the sodium across the membrane and really insulating it.
Then that means that the effect of those positive charges of the sodium,
instead of just being felt here, are felt all the way down here in the next node.
They're also felt down here in the previous node but
we're saying that's the spot that just had an action potential and
so here the sodium channels are inactivated.
And so, it doesn't matter that the depolarization is felt at the node.
They just fired but it does matter that it's felt at this downstream node.
And so, in this way the fact that the action potential can jump from node to
node, because the effects of the action potential are fell further away
because of the insulation that allows the transmission of the action potential or
the conduction of the action potential to happen much more quickly.
So that instead of having to have an action potential here and
then an action potential here and then an action potential here,
it just has one at the node and then at the next node.
So it's very rapid.
So there are two main ways that an axon
can have characteristics that allow for
very fast transmission, which is important for certain modalities such as touch.
So certain sensory axons are going to be myelinated,
some will be unmyelinated if it's not as important for fast transmission.
The other factor is the diameter of the axon.
So it's just like a wire where if you have a thicker wire,
a wire with a larger diameter,
then current will travel down it more quickly because there's less resistance.
Because there's more paths to choose because it has a larger diameter.
And so, an axon behaves just in the same way.
A larger diameter axon has lower resistance and so
axon potentials will travel faster down a large axon that's myelinated.
So to wrap this up, we've talked about an action
potential which is going to be where you're going to have sodium,
come in to the cell which is going to cause a large deep polarization.
And then, potassium is going to leave the cell and so we're going
to have a large repolarization and the fact that that's going to be all or
none, and that were going to have our refractory period,
which means that the cell needs to come back to rest in that
location before another axon potential can fire.
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