Lecture Slides Week 2 Slides

Bioelectricity week 2
Bioelectricity week 2
Bioelectricity week 2: A) Membrane Patch, the idea B) Get energy in the form of transmembrane voltage Vm Train system: Get money by selling tickets
Excitable membrane is not just some dead plastic shrink wrap!
Bioelectricity Week2 Lectures

Get energy in the form of transmembrane voltage Vm Membrane Patch, the idea
1. Introduction to week 2 2. A Membrane Patch, the idea of it. 3. Energy as Trans-membrane Voltage Vm 4. Sodium-Potassium Pumps 5. Ionic equilibrium 6. Battery Lifetime 7. Problem session 1 8. Membrane Resistance Rm 9. Membrane Capacitance Cm 10. Why is Cm so big? 11. Problem session, R and C 12. Week 2 summary
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Membrane Patch, the idea of it

What is a patch?
Answer: A segment of membrane large enough to have all the pumps and channels present In their true proportions.
How big is a patch?

Answer: Maybe a rectangle 10 to 100 micrometers on a side, more or less.
This week discussion of a patch leads in 2 directions First, a discussion of voltage across the patch (fascinating) Second, a discussion of patch resistance and capacitance (boring but necessary)
Membrane Patch in a Cell Sketch

A Shocking Answer to the question: Where is the electrical action that drives the cells electrical system?
Inside?
In teeny-tiny patches of the membrane?
Outside?
Duke University, Durham, NC, USA
Bioelectricity week 2.3

Bioelectricity: Get energy in the form of transmembrane voltage Vm
1. Introduction to week 2 2. A Membrane Patch 3. Energy as Trans-membrane Voltage Vm 4. Sodium-Potassium Pumps 5. Ionic equilibrium 6. Battery Lifetime 7. Problem session 8. Membrane Resistance Rm 9. Membrane Capacitance Cm 10. Why is Cm so big? 11. Problem session, R and C 12. Week 2 summary
Vm Across the Membrane Patch

What is the problem? Answer: How to get energy in the form of a trans-membrane voltage Vm.
How does the membrane charge itself up?
1 2
First the membrane pumps Potassium and Sodium Ions, both at the same time.
Then the membrane leaks the ions back out, selectively.

Sodium-Potassium Pump
K+ Membrane Outside Inside K+ K+ Na+
Na+ Na+

1. Introduction to week 2 2. A Membrane Patch 3. Energy as Trans-membrane Voltage Vm 4. Sodium-Potassium Pumps
5. Ionic equilibrium
6. Battery Lifetime 7. Problem session 8. Membrane Resistance Rm 9. Membrane Capacitance Cm 10. Why is Cm so big? 11. Problem session, R and C 12. Week 2 summary
Nernst for Potassium, K+ permeable
1) Membrane permeable to K+ only
Not an easy trick to do
Nernst for Potassium, Diffusion 1) Membrane permeable to K+ only 2) Diffusion pushes K+ out
Diffuses because of the random motion at the membrane temperature.
Nernst for Potassium, E field

1) Membrane permeable to K+ only 2) Diffusion pushes K out+
The positive charge created by K+ moving out creates the electric field pointing back in.
3) Electric Field pushes K + back in
Nernst for Potassium, Equilibrium

1) Membrane permeable to K+ only 2) Diffusion pushes K+ out 3) Electric Field pushes K+ back in
Equilibrium when the flow due to diffusion equals that from the electric field. It is a dynamic equilibrium, not static equilibrium It occurs close to the Nernst Vm for K+, but not quite at that Vm, because the pearmeability to K alone is not quite perfect. ln means the natural log, sometimes writen just log, or sometimes loge. RT/F was discussed earlier.
4) Equilibrium occurs near the

Nernst Vm for K+
Vm = (RT/F) ln(Ke/Ki) Vm = 26*ln(20/397) squid
Nernst for Potassium, Vm at equilibrium

1) Membrane permeable to K+ only 2) Diffusion pushes K+ out 3) Electric Field pushes K+ in 4) Dynamic equilibrium near the Nernst Vm
Notably negative, even compared to standard battery EK is a constant, for fixed concentrations and temperature, but Vm moves around from time to time. When Vm = EK there is Nernst equilibrium for K+ EK often is written with K subscript.
Vm at K+ equilibrium about -77mV (squid) Given the special symbol EK.
Nernst for Sodium Ions Na+

Same but different than K+
Membrane permeable to Na+ Diffusion pushes Na+ In Electric Field pushes Na+ Out Equilibrium near Nernst Vm for Na+ Vm about +57 mV (note positive, not negative)
Because
The Na+ is mostly outside
Nernst for Chlorine Ions ClSame but different than K+

Membrane permeable to Cl Equation for equilibrium is slightly different Vm = (RT/F) ln (Cl_i/Cl_e) Vm about -68 mV Vm at equilibrium is called ECl
Because
Different because Cl- is a negatively charged ion
Multi-ion expression (Goldmans equation)
With Im = 0 Then Vm is
The Ps are permeability coefficients
Duke University Engineering, Durham, NC

Energy: Battery Lifetime? What is the problem? Batteries we make discharge Die immediately in salt water Die in a few days in continuous use Die in a month to a year with infrequent use How to get 100 years?
Energy: Battery Lifetime? Charges before birth

Keeps on going, no matter how old
Imagine billions of spinning Na-K pumps in each of these devices
Duke University Divinity School

Problem session questions

Using the ionic concentrations for frog muscle, what is the Nernst potential (a) for K+ (b) for Na+ (c) the change in Vm from K+ to Na+ equilibrium?
Problem session K+
Using the ionic concentrations for frog muscle, what is the Nernst potential (a) for K+
Problem session Na+

Using the ionic concentrations for frog muscle, what is the Nernst potential (b) for Na+
Problem session, Vm shift

Using the ionic concentrations for frog muscle, what is the Nernst potential (c) the change in Vm from K+ to Na+ equilibrium? Vm shift = ENa EK ENa = EK = Vm shift =
Duke University Library

Membrane Resistance
Where does the membrane resistance come from?
Membrane Resistance
What is membrane resistance as compared to resistivity?
Membrane Resistance
Why use resistivity anyway?
Membrane Resistance
How does one get resistance R from resistivity Rm?
Membrane Resistance
Is membrane resistance a constant? Fairly constant when the tissue is passive. When the tissue is active, resistance changes around all over the place by huge factors.
Membrane Resistance
Why do we care about membrane resistance, if It changes around in active tissue? That is, it seems to be useful only for passive tissue, so do we care?
Duke University Engineering

Membrane Capacitance
Why does membrane have capacitance? Answer: because 2 conductors are separated by an insulating region
For membrane, where are the conductors? Answer: They are the conducting solutions inside and outside the membrane.
Cm = Q/Vm Q = Cm V dQ/dt = Cm dmV / dt Im = Cm dVm / dt
Duke University Chapel, Durham, NC, USA

Because membrane is so thin.
Why is Cm so big?
In biological membrane, Cm is about 1 microfarad per cm squared. That is a huge amount. That means that membrane has 10,000 times the capacitance of two flat conductors separated by a piece of paper, for the same surface area. The difference comes from the thickness of membrane as compared to the thickness of paper.
Membranes thickness versus papers
For flat plates, capacitance is C = eps A / d, where A is the area, and d is the separation between the plates For membranes, d is the membrane thickness, only 40 angstroms. In comparison, a stack of 1000 sheets of office paper is about 4cm high.
Membranes thickness versus papers
That means that the thickness of 1 sheet of office paper can be found as follows: Thickness 1 sheet paper = 4cm for 1000 sheets / 100 So thickness 1 sheet paper = 4e8 Angstroms / 1000 = 4e5 Angstroms So thickness 1 sheet paper = 400,000 Angstroms, approximately
So what is the capacitance of conductors separated by paper as compared to separated by membrane?

Thickness paper / Thickness membrane = 400,000/40 = 10,000 That is, paper is 10,000 times the thickness of membrane. That means that two flat conductors separated by membrane Would have 10,000 times the capacitance of two flat conductors separated by a piece of paper, for the same surface area. Conclusion: membranes have a lot of capacitance, about 1 micro-Farad per cm-squared.
Membrane capacitance, conclusion

Conclusion: Membranes have a lot of capacitance, about 1 micro-Farad per cm-squared. All electrically active membranes, from whatever organ or system, have about this much capacitance.

1. Introduction to week 2 2. A Membrane Patch 3. Energy as Trans-membrane Voltage Vm 4. Sodium-Potassium Pumps 5. Ionic equilibrium 6. Battery Lifetime 7. Problem session 8. Membrane Resistance Rm 9. Membrane Capacitance Cm 10. Why is Cm so big? 11. Problem Session, R and C 12. Week 2 summary
Problem Session, R and C, a)

Question: A cylindrical fiber 10um in diameter has segments that are 100um long. Cm is 1uF/cm2, and Rm is 1500 Ohms-cm2. For one segment, what is R and C?
Solution: a) Find the surface area As of one segment
Problem Session, R and C, b) and c)

Solution: a) As is b) R = Rm / As = c) C = Cm * As =
Problem Session, R and C, notes

a) Notation that does not use super and subscripts, such as cm2, nonetheless means the same thing. It is a form brought on by computer programming. b) Note that the units of Rm, membrane resistivity, are Ohm-cm2 and not the same as those for the bulk resistivity, Ohm-cm as used in week 1. c) Note that one divides Rm by As to get R, while one multiplies Cm by As to get C.
Duke University Medical Center

Week 2 Summary-1
A membrane patch is small but large enough to include pumps and channels in proportion to their presence in the whole membrane If circular it might be 25-50 microns in radius.
Week 2 Summary-2
A membrane patch is small but large enough to include pumps and channels in proportion to their presence in the whole membrane The Na-K pump moves Na+ ions outward across a patch membrane, and K+ ions inward, so that large concentration differences are established.
Week 2 Summary-3
A membrane patch is small but large enough to include pumps and channels in proportion to their presence in the whole membrane The Na-K pump moves Na+ ions outward across a patch membrane, and K+ ions inward, so that large concentration differences are established. Consequently, equilibrium voltages for K+ and for Na+ ions are well below and above 0 volts.
Week 2 Summary-4
A membrane patch is small but large enough to include pumps and channels in proportion to their presence in the whole membrane The Na-K pump moves Na+ ions outward across a patch membrane, and K + ions inward, so that large concentration differences are established. Consequently, equilibrium voltages for K+ and for Na+ ions are well below and above 0 volts.
There is a trans-membrane voltage when the membrane is selectively permeable to one or the other of these ions.
Week 2 Summary-5
These steps, taken as a whole, are a mechanism for using metabolic energy to run the membrane pumps. The concentration differences that result are used to create transmembrane voltages by selectively opening channels to either K+ or Na+ The trans-membrane voltages created in this way then drive currents across the membrane, as well as through and around the tissue structure.
Week 2 Summary-6
Voltages created in this way mean that the membrane can charge itself up The mechanism is distributed and reliable, and lasts a lifetime
Duke University Library

Bioelectricity, end of week 2

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Bioelectricity week 2

Excitable membrane is not just some dead plastic shrink wrap!

Bioelectricity Week2 Lectures

Train system: Get money by selling tickets

Membrane Patch, the idea of it

How big is a patch?

Membrane Patch in a Cell Sketch

In teeny-tiny patches of the membrane?

Duke University, Durham, NC, USA

Bioelectricity week 2.3

Train system: Get money by selling tickets

Vm Across the Membrane Patch

How does the membrane charge itself up?

Then the membrane leaks the ions back out, selectively.

Duke University, Durham, NC, USA

Bioelectricity week 2.4

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K+ Membrane Outside Inside K+ K+ Na+

Duke University, Durham, NC, USA

Bioelectricity week 2.5

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Nernst for Potassium, K+ permeable

1) Membrane permeable to K+ only

Not an easy trick to do

Nernst for Potassium, E field

3) Electric Field pushes K + back in

Nernst for Potassium, Equilibrium

4) Equilibrium occurs near the

Vm = (RT/F) ln(Ke/Ki) Vm = 26*ln(20/397) squid

Nernst for Potassium, Vm at equilibrium

Vm at K+ equilibrium about -77mV (squid) Given the special symbol EK.

Nernst for Sodium Ions Na+

Nernst for Chlorine Ions ClSame but different than K+

Multi-ion expression (Goldmans equation)

The Ps are permeability coefficients

Duke University Engineering, Durham, NC

Bioelectricity week 2.6

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Energy: Battery Lifetime? Charges before birth

Imagine billions of spinning Na-K pumps in each of these devices

Duke University Divinity School

Bioelectricity week 2.7

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Problem session questions

Problem session Na+

Problem session, Vm shift

Duke University Library

Bioelectricity week 2.8

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Duke University Engineering

Bioelectricity week 2.9

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Duke University Chapel, Durham, NC, USA

Bioelectricity week 2.10

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Because membrane is so thin.

Membranes thickness versus papers

Membranes thickness versus papers

So what is the capacitance of conductors separated by paper as compared to separated by membrane?

Membrane capacitance, conclusion

Bioelectricity week 2.11

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Problem Session, R and C, a)

Solution: a) Find the surface area As of one segment