Neurophysiology – lecture 20

March 29, 2011

1 Review:
1. The soma of neurons is large relative to axons and is filled with a relatively good conducting medium
(cytoplasm). Hence there are many current conducting pathways between various areas in the soma
and so there is only a small resistance between one site in the soma and another. As a result everywhere
inside a soma is at approximately the same potential – the soma is said to be “isopotential.”
2. Axons on the other hand are long and narrow which means there is substantial resistance between one
site along an axon and a somewhat distant site along the same axon.
3. Therefore in the electrical model of an axon handed out together with Figure 13 the axon is subdivided
into segments and there is a small resistance, labeled ri , placed between the segments and inside the
axon. It is also to be noted that each segment has Na+ , K+ , L and capacitive channels, thus each
segment is capable of generating an action potential.
4. Neuroscientists have learned about cable properties from the analysis of transatlantic cable properties.
The properties of transatlantic cable are dominated by 2 parameters: ri – the longitudinal resistance
of 1 cm of axon – and rm – the membrane resistance of 1 cm of axon.
5. ri has Ω/cm as its units while rm has Ωcm as its units.
6. If a change in transmembrane potential, V0 , occurs at one locus along an axon and is maintained for a
−t

rthen the transmembrane potential, Vm , at any locus along the axon equals Vrest + V0 e λ , where
while,
rm
λ= .
ri
7. The data describing voltage differences along an axon as collected by Hodgkin in 1937 are well fit by
this exponential decay equation.
8. Along axons λ = less than 1 cm. Since any depolarization along an axon is not noticeable 5 or more
λ away from the site of the depolarization, one can see that the spread of voltage along the length of
an axon does not provide a good means for communication along axons.

2 What then is a good system for communication along axons? A

Qualitative Model.
1. Assume an action potential is elicited at point X along an axon →
On either side of this action potential there will be a more or less exponentially decaying spread of
voltage into neighboring areas of the axon →

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 Those neighboring areas of the axon nearest to the site generating the action potential will be depo-
larized above “threshold” →
Therefore in the millisecond or so it takes to generate an action potential these neighboring areas of
axon membrane will have generated an action potential →
This new action potential will depolarize the areas of axon membrane neighboring on it →
This sequence: A.P. generation at an axonal site → spread of depolarization → triggering an A.P. in
neighboring areas of membrane → a new A.P. developing in that neighboring area – forms a reoccurring
cycle of events until the end of the axon is reached and there is no more “neighboring” axon membrane.
2. Note that “axon potential propagation” is really a sequence of generating new action potentials.
Each of these New action potentials is generated by an axon of uniform diameter, with the same
concentrations of ions inside and outside the axon and very similar densities of Na+ and K+ channels
in the axon membrane.
Thus each newly triggered action potential looks like the action potential that triggered it.
However, this new action potential is produced by a different area of axon membrane containing a
different set of Na+ and K+ channels and bathed inside and out by a different collection of Na+ and
K+ ions, so it’s an entirely different action potential.
3. I used the analogy that action potential “propagation” is like knocking over a series of upright dominoes.
Each falling domino is a separate entity whose fall is merely triggered by the preceding fall of another
domino. The end result is a propagated falling though the individual dominoes are all different.
4. The source of the energy needed to trigger a fall in the next domino in the chain is in the potential
energy present in the upright dominoes. This energy I had to place in them by setting them upright.
5. In similar manner the energy needed to depolarize neighboring areas of axonal membrane is present in
the ionic concentration gradients present between the inside and outside of the axon. These gradients
were established by the metabolism of ATP by the Na+ - K+ exchange pump.

3 The importance of refractory periods

1. Since each segment of axonal membrane contains all the ion channels and ion concentrations required
to generate an action potential and each segment is separated from adjacent segments by a relatively
small resistance, an action potential in segment A can trigger and action potential in segment B and
an action potential in segment B can trigger an action potential in segment A.
2. Action potentials being propagated along an axon away from the neuronal soma are called ortho-
dromic; whereas, action potentials propagated along an axon toward the neuronal soma are called
antidromic.
3. If I apply an electrical stimulus to the center of a previously quiescent axon and elicit an action
potential, there will be an action potential generated that travels in both directions away from the site
of stimulation.
4. Once a sequentially generated action potential begins to move in a particular direction along an axon
it cannot turn around and move in the opposite direction.
This is because the axonal membrane which has just supported an action potential is refractory for 10
– 20 ms after having just produced an action potential. Thus any area of axonal membrane which has
just produced an action potential cannot do so again for a sufficiently long time after the peak of the
action potential that the transmembrane potential is back to near resting potential level.

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 5. Behind every action potential propagated along an axon there is an area of refractory membrane that
cannot support a new action potential.
6. These areas of refractory membrane insure that action potential propagation is unidirectional once the
action potential is initiated.

4 Can we describe these propagated A.P. in a more quantitative

way?
1. During an action potential rm is changing so the simple model of exponential decay along the axon
does not really apply.
2. What then is the distribution of transmembrane potential along an axon when an action potential is
being produced in one area along the axon?
3. As an action potential is propagated past a site in which a recording microelectrode is placed, we are
able to record the changes in transmembrane potential over time which we call an action potential.
dVm
Call these changes .
dt
dVm dVm dx dVm dx dx
4. Then = = . is a speed, in this case we can think of it as the conduction
dt dt dx dx dt dt
dVm dVm
velocity of the action potential. We designate conduction velocity with a θ. So = . That is
dt dx
the distribution of transmembrane potential in time at a point along an axon is equal to a constant
times the distribution of transmembrane potential along an axon at a specific time.
5. Since we can easily evaluate we know the distribution of transmembrane potential along the axon at
any given time.
6. It is discovered that at the leading edge of a propagated action potential the transmembrane potential
rises exponentially for 10 to 15 mV just as the cable equation predicts. Sorthe cable equation model
rm
fits this small portion of the action potential voltage distribution and λ = is applicable here.
ri
7. But then the leading edge of a propagated action potential is the area where neighboring areas of
axonal membrane are being depolarized and the effectiveness of this depolarization is a major part of
what controls the speed with which action potentials are propagated.

5 Increasing θ by making ri small

1. The longitudinal resistance along the axon depends on the size of the axon and the conductivity or
resistivity of the axoplasm.
2. The specific cytoplasmic resistivity, Ri , is defined as the resistance between 2 opposing faces of a cube
1cm on a side filled with cytoplasm. Actual measurements are made on volumes of cytoplasm much
smaller than that.
30 Ω cm ≤ Ri ≤ 100 Ω cm for mammalian tissues

3. But the cross sectional area of an axon is much smaller than 1 cm2 , so the ri of 1 cm of axon will be
Ri
much larger than Ri (remember resistors in parallel add by the inverse law). So, ri = where a is
πa2
the radius of the axon.

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4. But as the axon radius changes the surface area of 1 cm of axon also changes and so rm changes.
Rm Rm
rm = =
S.A. 2π a

5. Therefore, s r
Rm πa2
r
rm Rm a
λ= = =
ri 2π a Ri 2 Ri

6. Thus, as the radius increases λ increases.

7. As λ increases, θ increases. The squid got it right!
8. A slide was shown to indicate that this is true not only for unmyelinated axons but also for myelinated
axons.