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Neurophysiology – lecture 7

January 27, 2011

1 Review:
1. The concentration of cations in the cytoplasm = the concentration of anions in the cytoplasm. This is
a state of Electroneutrality.
2. The simplified model of a cell has K+ and organic anions (A− ) in the cytoplasm and Na+ and Cl− in
the extracellular fluid.
3. In solutions ions are good conductors of electricity.
4. We showed why cells can be represented in Electrical Models as a wire where the cytoplasm is, a wire
where the extracellular fluid is and an array of resistors connecting those 2 wires where the lipid bilayer
membrane is. This model can be simplified further by replacing the array of parallel resistors with one
equivalent resistor, Req , where
n
X
−1
Req = r1−1 + . . . + rn−1 = ri−1 (1)
i=1

5. This latter equation depends on Kirchhoffs Current Law which states that the sum of current going to
a junction = the sum of the currents coming out of the junction.
6. If a constant current is passed into a cell, I, produces a change in transmembrane voltage equal to
∆Vm then Rin = ∆VI m where Rin is called the input resistance.
7. Larger cells have more surface area and therefore more current pathways out of or into the cell and
hence have lower resistance.
8. But we expect a given area of membrane to have a similar resistance to other areas of the cell membrane
no matter where on the cell it came from or even which cell it came from.
To see if this is true we define Rm = the resistance of a unit area of membrane = Rin × cell surface
area,A. Rm is quantified in Ω · cm2
9. Rm for cells is generally between 1-10 KΩ · cm2 .
10. This value for Rm is much larger than it would be if the cell membrane were composed of pure protein
or carbohydrate, but is much lower than it would be if the membrane were composed of pure lipid.

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2 But why is the Rm of the membrane less than that of pure
lipid?
1. The patch clamp studies indicate that membranes contain ion channels. Anatomical studies indicate
that there are 100 to 10000 ion channels
mm2 of neuronal membrane.
2. Since a small number of these channels are open at any given time and these channels open from one
side of the lipid membrane to the other, these ion channels should be considered as resistors of lower
value than the resistors representing the lipids and in parallel with the resistors representing the lipids
spanning the membrane.
3. So for a membrane:
n m
1 XX 1 1
= + (2)
Rin r
i=1 j=1 Li
rchj

where rch is the resistance of a channel and rL represents the resistance of a small area of lipid
membrane.
4. To simplify such an equation and evaluate what it states, we define a new parameter: Conductance
symbolized by G or g = R1 or 1r and is quantified in S, Siemens.
n X
X m
5. The Gin of a cell membrane: Gin = gLi + gchj
i=1 j=1

6. Since gch  gL , Gin = n · gL + m · gch > (n + m) · gL


So the relatively high Gin of cell membranes is due to the relatively high g of the ion channels in the
membrane.

3 Needed modifications of the present electrical model of cell


membranes
1. As described above, our present electrical model of cell membranes is of an array of parallel resistors
spanning between the intracellular solution and the extracellular solution.
2. This model can be reduced to 1 equivalent resistor. If there were only 1 resistor between the inside and
the outside of the cell, then when current is passed across this resistor the voltage difference across it
should “jump” instantly to a value determined by Ohms Law.
3. However, when this experiment is done in the laboratory records like those in Figure 6A are produced.
In these records the transmembrane voltage takes some time to reach the value predicted by Ohms
Law.
4. So we need to change our Electrical Model to make it correspond to what is observed in the laboratory.
5. I bluntly stated and hope to demonstrate in next Tuesdays lecture that if we add a “capacitor” to the
Electrical Model, the Electrical Model will also change voltage gradually when a rectangular current
is run through it.

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4 So what is a capacitor and how does it work?
1. A capacitor is composed simply of 2 surfaces composed of conducting material separated by a layer of
material which has a high resistance. This separating material is also called a dielectric.
2. To consider how a capacitor operates we will start with the analog model, the Hydraulic Model (Figure
6B).
3. In A. Initially, we assume there is no force applied to the pump handle, hence no fluid movement.
The 2 springs in the capacitor chamber are under low and equal tension and hence the piston in the
chamber does not move.
4. In B. A short time later, we assume that force has now been applied to the pump handle and the pump
piston has move ∆s distance. This movement displaced water from the pump and pushed fluid into
the capacitor chamber. While moving fluid into the capacitor chamber, the piston in the capacitor
has been moved upward slightly compressing the upper spring and extending the lower spring. These
springs will then exert some force on the piston to restore it to the central position in the capacitor
chamber.
5. In C. After a long time, the pump has pushed considerable fluid into the capacitor chamber, displacing
the piston upward farther and compressing the upper spring more and extending the lower spring
farther. A some juncture there will be enough downward force exerted by the compressed upper spring
and downward force exerted by the stretched lower spring to together sum to produce a downward
force equal to force being applied to the pump handle. Since these 2 forces are equal and opposite, no
more pumping will occur. The fluid that has been pumped into the capacitor chamber will stay there
as long as the same force is applied to the pump handle.
6. You were then challenged to go through the same series of steps using the Electrical Model.
7. In A. Initially, we assume the plates in the capacitor each contain an equal number of positive and
negative charges and therefore have no net charge. While the battery or current source is poised to
place charge onto the capacitor plates it has not yet done so simply because this is 10-100 seconds after
the circuit has been completed.
8. In B. A short time later, the battery or current source has driven a small amount of positive charge
onto the lower plate and removed an equal amount of positive charge from the upper plate. Thus the
lower plate has a slight amount of excess positive charge and the upper plate has a slight amount of
excess negative charge. Thus, a small electrical field is generated between the two plates which serves
to generate an attractive force between the positive charges on the lower plate and the negative charges
on the upper plate. That is to say a small voltage difference is generated between the plates. (In other
words, work could be done if the positive charge on the lower plate were allowed to move to the upper
plate, but this does not happen because there is no conducting pathway between the two plates.)
9. In C. After a long time, the battery or current source has added sufficient positive charge to the lower
plate and removed an equal amount of positive charge from the upper plate that results in the voltage
difference between the plates having grown to be equal to the voltage difference supplied by the battery
or current source. But the voltage difference supplied by the battery or current source is oppositely
directed from the voltage difference generated across the capacitor plates. Hence the sum of these 2
voltage differences around the circuit will be 0. At this juncture there will be no more charge movement
(current). At this juncture the capacitor would be said to be “charged”.
10. Note:
(a) No charges actually moved from one plate to another, so the current that did occur is unlike
resistive or ionic current in which there is actual movement of charge.

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(b) The effect on the charges located on opposite plates of the capacitor was produced by electric
fields.
(c) The current was large at first and then decayed to 0 as the capacitor got more charges placed on
it.
(d) This current is called capacitative current, Ic .

5 RC circuits as seen with biological membranes


1. While membranes have capacitance they also have resistance as we have previously seen. So we must
combine these 2 electrical components into our Electrical Model. This is done by wiring the membrane
equivalent resistor in parallel with a capacitor with both spanning the membrane.
2. If we now pass a rectangular current pulse through this Electrical Model of a membrane we expect to
see the following sequence of transmembrane voltage changes:
At the start of the current pulse there has been no charge previously placed on the membrane capacitor
so little work is need to place an initial amount of charge on it, whereas it always takes work to pass
charge through a resistor. For this reason at the very beginning of the current pulse essentially all of
the current will pass onto the capacitor plates as Ic .
As + charge is added to the lower capacitor plate and removed from the upper, it becomes harder
(requires more work) to add additional + charges to the lower plate and remove them from the upper
plate, because of the repulsive force produced by the charges that were previously placed on the lower
plate and the attractive force produced by the negative charges left on the upper plate. These forces
will slow the addition of new charges to the capacitor plates. But the current source is still passing
the same amount of current so some of this current is passed through the resistor rather than the onto
the capacitor.
After current has been passed through this circuit for a while the + charges on the lower plate of
the capacitor and the - charges on the upper plate of the capacitor will have increased to the point
that it takes as much work to place addition charge onto the capacitor plates as the current source
is capable of producing. At this final stage all of the current supplied by the current source will be
passing through the resistor and this resistive current will be maintained as long as current is being
passed from the current source.
3. To summarize, given a circuit with a resistor and a capacitor in parallel the current from a current
source while initially pass onto the capacitor plates and not through the resistor, but as the charge on
the capacitor plates increases more and more current will pass through the resistor until the capacitor
becomes “completely charged” and all of the current passes through the resistor.
4. Note that Ohms Law then tells you that the ∆V produced by the passage of current over time will
have the same form as the form of the resistive current has; i.e., initially there is no change in ∆V in
response to the current pulse at its onset but then the ∆V rises slowly and exponentially to a maximum
or asymptote.
This is the pattern of ∆Vm that is seen when a rectangular current pulse is passed through a biological
membrane (see Figure 6A).

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