6. Membrane potential and the passive electrical properties of the neuron Flashcards
What are the two types of ion channels?
resting and gated
What are the roles of resting and gated ion channels?
Resting channels are primarily important in maintaining the resting membrane potential, the electrical potential across the membrane in the absence of signalling. Some types of resting channels are constitutively open and are not gated by changes in membrane voltage; other types are gated by voltage but can open at the negative resting potential of neurons. Most voltage-gated channels, in contrast, are closed when the membrane is at rest and require membrane depolarisation to open.
What causes the membrane potential (at an atomic level)
The neuron’s cell membrane has thin clouds of positive and negative ions spread over its inner (-) and outer (+) surfaces. At rest the extracellular surface of the membrane has an excess of positive charge and the cytoplasmic surface an excess of negative charge
How is the membrane potential (Vm) defined as?
Vm = Vin - Vout
where Vin is the potential on the inside of the cell and Vout the potential on the outside.
What is the membrane potential at rest called? What previously mentioned variable is it equal to?
The membrane potential of a cell at rest is called the resting membrane potential (Vr). Since by convention the potential outside the cell is defined as zero, the resting potential is equal to Vin.
What is the typical range of the membrane potential?
−60 mV to −70 mV
How is the direction of current defined?
The direction of current is conventionally defined as the direction of net movement of positive charge.
What does this (definition of direction of current) mean in terms of the cations and anions in the cell?
Thus, in an ionic solution cations move in the direction of the electric current and anions move in the opposite direction.
Describe the net charge movement of the cell at rest
In the nerve cell at rest there is no net charge movement across the membrane. When there is a net flow of cations or anions into or out of the cell, the charge separation across the resting membrane is disturbed, altering the electrical potential of the membrane.
What is meant by depolarisation and hyperpolarisation?
A reduction or reversal of charge separation, leading to a less negative membrane potential, is called depolarisation. An increase in charge separation, leading to a more negative membrane potential, is called hyperpolarisation.
What are meant by electrotonic potentials?
Changes in membrane potential that do not lead to the opening of gated ion channels are passive responses of the membrane and are called electrotonic potentials.
What kind of responses are typically electrotonic potentials?
Hyperpolarising responses are almost always passive, as are small depolarisations.
When is an action potential triggered?
When depolarisation approaches a critical level, or threshold, the cell responds actively with the opening of voltage-gated ion channels, which produces an all-or-none action potential.
Describe how the membrane potential can be recorded from a cell
Glass micropipettes filled with a concentrated salt solution serve as electrodes and are placed on either side of the cell membrane. Wires inserted into the back ends of the pipettes are connected via an amplifier to an oscilloscope, which displays the amplitude of the membrane potential in volts. Because the diameter of a microelectrode tip is small (< 1 μm), it can be inserted into a cell with relatively little damage to the cell membrane.
How can you know you’re recording from inside the cell?
When both electrodes are outside the cell, no electrical potential difference is recorded. But as soon as one microelectrode is inserted into the cell, the oscilloscope shows a steady voltage, the resting membrane potential. In most nerve cells at rest the membrane potential is approximately −65 mV.
How could causal claims be made about changes to the membrane potential?
The membrane potential can be experimentally changed using a current generator connected to a second pair of electrodes—one intracellular and one extracellular. When the intracellular electrode is made positive with respect to the extracellular one, a pulse of positive current from the current generator causes positive charge to flow into the neuron from the intracellular electrode. This current returns to the extracellular electrode by flowing outward across the membrane.
How are differences seen between small depolarising currents and larger currents being injected into the cell?
Small depolarising current pulses evoke purely
electrotonic (passive) potentials in the cell—the size of the change in potential is proportional to the size of the current pulses. However, a sufficiently large depolarising current triggers the opening of voltage-gated ion channels. The opening of these channels leads to the action potential, which differs from electrotonic potentials in the way in which it is generated as well as in magnitude and duration.
How are differences seen between small hyperpolarising currents and larger currents being injected into the cell?
The responses of the cell to hyperpolarisation are usually purely electrotonic—as the size of the current pulse increases, the hyperpolarisation increases proportionately. Hyperpolarisation does not trigger an active response in the cell.
Comment on the concentrations of different ion species inside and outside the cell
Of the four most abundant ions found on either side of the cell membrane, Na+ and Cl− are concentrated outside the cell and K+ and organic anions (A -, primarily amino acids and proteins) inside.
What is meant by the equilibrium potential?
The membrane potential at which there is no net flux of the ion species across the cell membrane
Roughly describe the distribution of these ions inside and outside of one particularly well-studied nerve cell process, the giant axon of the squid, whose extracellular fluid has a salt concentration similar to that of seawater. Give the exact equilibrium potential for the following:
K+
Na+
Cl-
A-
Species; Conc Cyto (Mm); Conc Extr (Mm); Eq pot
K+; 400; 20; -75
Na+; 50; 440; +55
Cl-; 52; 560; -60
A-; 385; none ; none
What is the permeability of a cell membrane to a particular ion species determined by?
The relative proportions of the various types of ion channels that are open.
The simplest case is that of the glial cell to demonstrate this permeability. Why is that?
The simplest case is that of the glial cell, which has a resting potential of approximately −75 mV. Like most cells, a glial cell has high concentrations of K+ and negatively charged organic anions on the inside and high concentrations of Na+ and Cl− on the outside. However, most resting channels in the membrane are permeable only to K +.
Why is the membrane potential of glial cells negative then?
Because K+ ions are present at a high concentration inside the cell, they tend to diffuse across the mem- brane from the inside to the outside of the cell down their chemical concentration gradient. As a result, the outside of the membrane accumulates a net positive charge (caused by the slight excess of K+) and the inside a net negative charge (because of the deficit of K+ and the resulting slight excess of anions).
Why do these potassium ions line the membrane of the extracellular fluid?
Because opposite charges attract each other, the excess positive charges on the outside and the excess negative charges on the inside collect locally on either surface of the
membrane
What does it mean to say that the flux of K+ out of the cell is self-limiting?
The efflux of K+ gives rise to an electrical potential difference; positive outside, negative inside. The greater the flow of K+, the more charge is separated and the greater is the potential difference. Because K+ is positive, the negative potential inside the cell tends to oppose the further efflux of K+.
Thus, how can ions be driven across a membrane?
Thus ions are subject to two forces driving them across the membrane: (1) a chemical driving force, a function of the concentration gradient across the membrane, and (2) an electrical driving force, a function of the electrical potential difference across the membrane.
How do these two forces relate to the equilibrium potetial?
Once K+ diffusion has proceeded to a certain point, the electrical driving force on K+ exactly balances the chemical driving force. That is, the outward movement of K+ (driven by its concentration gradient) is equal to the inward movement of K+ (driven by the electrical potential difference across the membrane). This potential is called the K+ equilibrium potential, Ek
How is the equilibrium potential related to the resting membrane potential?
In a cell permeable only to K+ ions, Ek determines the resting membrane potential, which in most glial cells is approximately −75 mV.
How can the equilibrium potential for any ion X be calculated?
The equilibrium potential for any ion X can be calculated from an equation derived in 1888 from basic thermodynamic principles by the German physical chemist Walter Nernst:
Ex = (RT/ zF) (ln [X]o / [X]i)
where R is the gas constant, T the temperature (in degrees Kelvin), z the valence of the ion, F the Faraday constant, and [X]o and [X]i the concentrations of the ion outside and inside the cell. (To be precise, chemical activities rather than concentrations should be used.)
What is the value of the first part of the equation at 25C? (RT/F)
RT/F is 25 mV at 25°C (room temperature)
Describe how you would get the value of -75mV for K+ in a glial cell given that Ko = 20 and Ki = 400
Since RT/F is 25 mV at 25°C (room temperature), and the constant for converting from natural logarithms to base 10 logarithms is 2.3, the Nernst equation can also be written as follows:
Ex = (58mV / z) log(Xo/Xi)
Thus, for K+, since z = +1 and given the concentrations inside and outside the squid axon:
Ex = (58mV / 1) log(20/400) = -75mV
How are nerve cells different to glial cells in their permeability?
Unlike glial cells, nerve cells at rest are permeable to Na+ and Cl− ions in addition to K+ ions. Of the abundant ion species in nerve cells, only the large organic anions (A−) are unable to permeate the cell membrane.
Describe what would happen with the aforementioned K+ permeable cell with concentration gradients for K+, Na+, Cl-, and A- if a few resting Na+ channels are added to the membrane, making it slightly permeable to Na+.
Two forces drive Na into the cell: Na+ tends to flow into the cell down its chemical concentration gradient, and it is driven into the cell by the negative electrical potential difference across the membrane. The influx of Na+ depolarises the cell, but only slightly from the K+ equilibrium potential (−75 mV). The new membrane potential does not come close to the Na+ equilibrium potential of +55 mV because there are many more resting K+ channels than Na+ channels in the membrane.
As soon as the membrane potential begins to
depolarise from the value of the K+ equilibrium potential, K+ flux is no longer in equilibrium across the membrane. The reduction in the negative electrical force driving K+ into the cell means that there is now a net flow of K+ out of the cell, tending to counteract the Na+ influx. The more the membrane potential is depolarised and driven away from the K equilibrium potential, the greater is the net electrochemical force driving K+ out of the cell and consequently the greater the net K+ efflux. Eventually, the membrane potential reaches a new resting level at which the increased outward movement of K+ just balances the inward movement of Na+. This balance point (usually approximately −65 mV) is far from the Na+ equilibrium potential (+55 mV) and is only slightly more positive than the K+ equilibrium potential (−75 mV).
What is the magnitude of the flux of an ion the product of?
The magnitude of the flux of an ion across a cell membrane is the product of its electrochemical driving force (the sum of the electrical and chemical driving forces) and the conductance of the membrane to the ion
ion flux = (electrical driving force + chemical driving force) × membrane conductance.