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Membrane potential

Membrane potential (also transmembrane potential or membrane voltage) is the difference in electric potential between the interior and the exterior of a biological cell. That is, there is a difference in the energy required for electric charges to move from the internal to exterior cellular environments and vice versa, as long as there is no acquisition of kinetic energy or the production of radiation. The concentration gradients of the charges directly determine this energy requirement. For the exterior of the cell, typical values of membrane potential, normally given in units of milli volts and denoted as mV, range from –80 mV to –40 mV.

All animal cells are surrounded by a membrane composed of a lipid bilayer with proteins embedded in it. The membrane serves as both an insulator and a diffusion barrier to the movement of ions. Transmembrane proteins, also known as ion transporter or ion pump proteins, actively push ions across the membrane and establish concentration gradients across the membrane, and ion channels allow ions to move across the membrane down those concentration gradients. Ion pumps and ion channels are electrically equivalent to a set of batteries and resistors inserted in the membrane, and therefore create a voltage between the two sides of the membrane.


Almost all plasma membranes have an electrical potential across them, with the inside usually negative with respect to the outside.[2] The membrane potential has two basic functions. First, it allows a cell to function as a battery, providing power to operate a variety of "molecular devices" embedded in the membrane.[3] Second, in electrically excitable cells such as neurons and muscle cells, it is used for transmitting signals between different parts of a cell. Signals are generated by opening or closing of ion channels at one point in the membrane, producing a local change in the membrane potential. This change in the electric field can be quickly sensed by either adjacent or more distant ion channels in the membrane. Those ion channels can then open or close as a result of the potential change, reproducing the signal.


In non-excitable cells, and in excitable cells in their baseline states, the membrane potential is held at a relatively stable value, called the resting potential. For neurons, resting potential is defined as ranging from –80 to –70 millivolts; that is, the interior of a cell has a negative baseline voltage of a bit less than one-tenth of a volt. The opening and closing of ion channels can induce a departure from the resting potential. This is called a depolarization if the interior voltage becomes less negative (say from –70 mV to –60 mV), or a hyperpolarization if the interior voltage becomes more negative (say from –70 mV to –80 mV). In excitable cells, a sufficiently large depolarization can evoke an action potential, in which the membrane potential changes rapidly and significantly for a short time (on the order of 1 to 100 milliseconds), often reversing its polarity. Action potentials are generated by the activation of certain voltage-gated ion channels.


In neurons, the factors that influence the membrane potential are diverse. They include numerous types of ion channels, some of which are chemically gated and some of which are voltage-gated. Because voltage-gated ion channels are controlled by the membrane potential, while the membrane potential itself is influenced by these same ion channels, feedback loops that allow for complex temporal dynamics arise, including oscillations and regenerative events such as action potentials.

Eeq,K+= equilibrium potential for potassium, measured in

volts

R = universal , equal to 8.314 joules·K−1·mol−1

gas constant

T = , measured in kelvins (= K = degrees Celsius + 273.15)

absolute temperature

z = number of of the ion in question involved in the reaction

elementary charges

F = , equal to 96,485 coulombs·mol−1 or J·V−1·mol−1

Faraday constant

[K+]o= extracellular concentration of potassium, measured in ·m−3 or mmol·l−1

mol

[K+]i= intracellular concentration of potassium

Driving force is the net electrical force available to move that ion across the membrane. It is calculated as the difference between the voltage that the ion "wants" to be at (its equilibrium potential) and the actual membrane potential (Em). So, in formal terms, the driving force for an ion = Em - Eion

For example, at our earlier calculated resting potential of −73 mV, the driving force on potassium is 7 mV : (−73 mV) − (−80 mV) = 7 mV. The driving force on sodium would be (−73 mV) − (60 mV) = −133 mV.

Permeability is a measure of how easily an ion can cross the membrane. It is normally measured as the (electrical) conductance and the unit, , corresponds to 1 C·s−1·V−1, that is one coulomb per second per volt of potential.

siemens

From the viewpoint of biophysics, the resting membrane potential is merely the membrane potential that results from the membrane permeabilities that predominate when the cell is resting. The above equation of weighted averages always applies, but the following approach may be more easily visualized. At any given moment, there are two factors for an ion that determine how much influence that ion will have over the membrane potential of a cell:


If the driving force is high, then the ion is being "pushed" across the membrane. If the permeability is high, it will be easier for the ion to diffuse across the membrane.


So, in a resting membrane, while the driving force for potassium is low, its permeability is very high. Sodium has a huge driving force but almost no resting permeability. In this case, potassium carries about 20 times more current than sodium, and thus has 20 times more influence over Em than does sodium.


However, consider another case—the peak of the action potential. Here, permeability to Na is high and K permeability is relatively low. Thus, the membrane moves to near ENa and far from EK.


The more ions are permeant the more complicated it becomes to predict the membrane potential. However, this can be done using the Goldman-Hodgkin-Katz equation or the weighted means equation. By plugging in the concentration gradients and the permeabilities of the ions at any instant in time, one can determine the membrane potential at that moment. What the GHK equations means is that, at any time, the value of the membrane potential will be a weighted average of the equilibrium potentials of all permeant ions. The "weighting" is the ions relative permeability across the membrane.

Effects and implications[edit]

While cells expend energy to transport ions and establish a transmembrane potential, they use this potential in turn to transport other ions and metabolites such as sugar. The transmembrane potential of the mitochondria drives the production of ATP, which is the common currency of biological energy.


Cells may draw on the energy they store in the resting potential to drive action potentials or other forms of excitation. These changes in the membrane potential enable communication with other cells (as with action potentials) or initiate changes inside the cell, which happens in an egg when it is fertilized by a sperm.


Changes in the dielectric properties of plasma membrane may act as hallmark of underlying conditions such as diabetes and dyslipidemia.[40]


In neuronal cells, an action potential begins with a rush of sodium ions into the cell through sodium channels, resulting in depolarization, while recovery involves an outward rush of potassium through potassium channels. Both of these fluxes occur by passive diffusion.


A dose of salt may trigger the still-working neurons of a fresh cut of meat into firing, causing muscle spasms.[41][42][43][44][45]

Bioelectrochemistry

Chemiosmotic potential

Electrochemical potential

Goldman equation

Membrane biophysics

Microelectrode array

Saltatory conduction

Surface potential

Gibbs–Donnan effect

Synaptic potential

Alberts et al. Molecular Biology of the Cell. Garland Publishing; 4th Bk&Cdr edition (March, 2002).  0-8153-3218-1. Undergraduate level.

ISBN

Guyton, Arthur C., John E. Hall. Textbook of medical physiology. W.B. Saunders Company; 10th edition (August 15, 2000).  0-7216-8677-X. Undergraduate level.

ISBN

Hille, B. Ionic Channel of Excitable Membranes Sinauer Associates, Sunderland, MA, USA; 1st Edition, 1984.  0-87893-322-0

ISBN

Nicholls, J.G., Martin, A.R. and Wallace, B.G. From Neuron to Brain Sinauer Associates, Inc. Sunderland, MA, USA 3rd Edition, 1992.  0-87893-580-0

ISBN

Ove-Sten Knudsen. Biological Membranes: Theory of Transport, Potentials and Electric Impulses. Cambridge University Press (September 26, 2002).  0-521-81018-3. Graduate level.

ISBN

National Medical Series for Independent Study. Physiology. Lippincott Williams & Wilkins. Philadelphia, PA, USA 4th Edition, 2001.  0-683-30603-0

ISBN

Functions of the Cell Membrane

Nernst/Goldman Equation Simulator

Nernst Equation Calculator

Goldman-Hodgkin-Katz Equation Calculator

Electrochemical Driving Force Calculator

- Online interactive tutorial (Flash)

The Origin of the Resting Membrane Potential