This will result in a greater accumulation of sodium ions than chloride ions in solution B and a lesser number of sodium ions than chloride ions in solution A. This means that there is a net positive charge in solution B from the higher concentration of positively charged sodium ions than negatively charged chloride ions. Likewise, there is a net negative charge in solution A from the greater concentration of negative chloride ions than positive sodium ions.
Since opposite charges attract and like charges repel, the ions are now also influenced by electrical fields as well as forces of diffusion. Therefore, positive sodium ions will be less likely to travel to the now-more-positive B solution and remain in the now-more-negative A solution. The point at which the forces of the electric fields completely counteract the force due to diffusion is called the equilibrium potential.
At this point, the net flow of the specific ion in this case sodium is zero. Every animal cell is enclosed in a plasma membrane , which has the structure of a lipid bilayer with many types of large molecules embedded in it. Because it is made of lipid molecules, the plasma membrane intrinsically has a high electrical resistivity, in other words a low intrinsic permeability to ions. However, some of the molecules embedded in the membrane are capable either of actively transporting ions from one side of the membrane to the other or of providing channels through which they can move.
In electrical terminology, the plasma membrane functions as a combined resistor and capacitor. Resistance arises from the fact that the membrane impedes the movement of charges across it. Capacitance arises from the fact that the lipid bilayer is so thin that an accumulation of charged particles on one side gives rise to an electrical force that pulls oppositely charged particles toward the other side.
The conductance of a pure lipid bilayer is so low, on the other hand, that in biological situations it is always dominated by the conductance of alternative pathways provided by embedded molecules. Thus, the capacitance of the membrane is more or less fixed, but the resistance is highly variable.
The thickness of a plasma membrane is estimated to be about nanometers.
Because the membrane is so thin, it does not take a very large transmembrane voltage to create a strong electric field within it. Typical membrane potentials in animal cells are on the order of millivolts that is, one tenth of a volt , but calculations show that this generates an electric field close to the maximum that the membrane can sustain—it has been calculated that a voltage difference much larger than millivolts could cause dielectric breakdown , that is, arcing across the membrane.
The resistance of a pure lipid bilayer to the passage of ions across it is very high, but structures embedded in the membrane can greatly enhance ion movement, either actively or passively , via mechanisms called facilitated transport and facilitated diffusion. The two types of structure that play the largest roles are ion channels and ion pumps , both usually formed from assemblages of protein molecules.
Ion channels provide passageways through which ions can move. In most cases, an ion channel is permeable only to specific types of ions for example, sodium and potassium but not chloride or calcium , and sometimes the permeability varies depending on the direction of ion movement. Ion pumps, also known as ion transporters or carrier proteins, actively transport specific types of ions from one side of the membrane to the other, sometimes using energy derived from metabolic processes to do so.
Ion pumps are integral membrane proteins that carry out active transport , i. The ion pump most relevant to the action potential is the sodium—potassium pump , which transports three sodium ions out of the cell and two potassium ions in. If the numbers of each type of ion were equal, the sodium—potassium pump would be electrically neutral, but, because of the three-for-two exchange, it gives a net movement of one positive charge from intracellular to extracellular for each cycle, thereby contributing to a positive voltage difference. The pump has three effects: 1 it makes the sodium concentration high in the extracellular space and low in the intracellular space; 2 it makes the potassium concentration high in the intracellular space and low in the extracellular space; 3 it gives the intracellular space a negative voltage with respect to the extracellular space.
The sodium-potassium pump is relatively slow in operation. If a cell were initialized with equal concentrations of sodium and potassium everywhere, it would take hours for the pump to establish equilibrium. The pump operates constantly, but becomes progressively less efficient as the concentrations of sodium and potassium available for pumping are reduced. Ion pumps influence the action potential only by establishing the relative ratio of intracellular and extracellular ion concentrations.
The action potential involves mainly the opening and closing of ion channels not ion pumps. If the ion pumps are turned off by removing their energy source, or by adding an inhibitor such as ouabain , the axon can still fire hundreds of thousands of action potentials before their amplitudes begin to decay significantly. Another functionally important ion pump is the sodium-calcium exchanger. Because the net flow of charge is inward, this pump runs "downhill", in effect, and therefore does not require any energy source except the membrane voltage. Its most important effect is to pump calcium outward—it also allows an inward flow of sodium, thereby counteracting the sodium-potassium pump, but, because overall sodium and potassium concentrations are much higher than calcium concentrations, this effect is relatively unimportant.
The net result of the sodium-calcium exchanger is that in the resting state, intracellular calcium concentrations become very low. Ion channels are integral membrane proteins with a pore through which ions can travel between extracellular space and cell interior. Most channels are specific selective for one ion; for example, most potassium channels are characterized by selectivity ratio for potassium over sodium, though potassium and sodium ions have the same charge and differ only slightly in their radius.
The channel pore is typically so small that ions must pass through it in single-file order. When a channel is open, ions permeate through the channel pore down the transmembrane concentration gradient for that particular ion. Rate of ionic flow through the channel, i. A channel may have several different states corresponding to different conformations of the protein , but each such state is either open or closed. In general, closed states correspond either to a contraction of the pore—making it impassable to the ion—or to a separate part of the protein, stoppering the pore.
For example, the voltage-dependent sodium channel undergoes inactivation , in which a portion of the protein swings into the pore, sealing it. Ion channels can be classified by how they respond to their environment. Ligand-gated channels form another important class; these ion channels open and close in response to the binding of a ligand molecule , such as a neurotransmitter. Other ion channels open and close with mechanical forces.
Still other ion channels—such as those of sensory neurons —open and close in response to other stimuli, such as light, temperature or pressure. Leakage channels are the simplest type of ion channel, in that their permeability is more or less constant. The types of leakage channels that have the greatest significance in neurons are potassium and chloride channels. Even these are not perfectly constant in their properties: First, most of them are voltage-dependent in the sense that they conduct better in one direction than the other in other words, they are rectifiers ; second, some of them are capable of being shut off by chemical ligands even though they do not require ligands in order to operate.
Ligand-gated ion channels are channels whose permeability is greatly increased when some type of chemical ligand binds to the protein structure. Animal cells contain hundreds, if not thousands, of types of these. A large subset function as neurotransmitter receptors —they occur at postsynaptic sites, and the chemical ligand that gates them is released by the presynaptic axon terminal.
One example of this type is the AMPA receptor , a receptor for the neurotransmitter glutamate that when activated allows passage of sodium and potassium ions.
Neurotransmitter receptors are activated by ligands that appear in the extracellular area, but there are other types of ligand-gated channels that are controlled by interactions on the intracellular side. Voltage-gated ion channels , also known as voltage dependent ion channels , are channels whose permeability is influenced by the membrane potential. They form another very large group, with each member having a particular ion selectivity and a particular voltage dependence.
Many are also time-dependent—in other words, they do not respond immediately to a voltage change but only after a delay. One of the most important members of this group is a type of voltage-gated sodium channel that underlies action potentials—these are sometimes called Hodgkin-Huxley sodium channels because they were initially characterized by Alan Lloyd Hodgkin and Andrew Huxley in their Nobel Prize-winning studies of the physiology of the action potential. The channel is closed at the resting voltage level, but opens abruptly when the voltage exceeds a certain threshold, allowing a large influx of sodium ions that produces a very rapid change in the membrane potential.
Recovery from an action potential is partly dependent on a type of voltage-gated potassium channel that is closed at the resting voltage level but opens as a consequence of the large voltage change produced during the action potential. The reversal potential or equilibrium potential of an ion is the value of transmembrane voltage at which diffusive and electrical forces counterbalance, so that there is no net ion flow across the membrane.
This means that the transmembrane voltage exactly opposes the force of diffusion of the ion, such that the net current of the ion across the membrane is zero and unchanging. The reversal potential is important because it gives the voltage that acts on channels permeable to that ion—in other words, it gives the voltage that the ion concentration gradient generates when it acts as a battery.
The equilibrium potential of a particular ion is usually designated by the notation E ion. The equilibrium potential for any ion can be calculated using the Nernst equation. Even if two different ions have the same charge i.
Take, for example, the equilibrium potentials of potassium and sodium in neurons. A neuron 's resting membrane potential actually changes during the development of an organism. In order for a neuron to eventually adopt its full adult function, its potential must be tightly regulated during development. As an organism progresses through development the resting membrane potential becomes more negative.
The drop in extracellular potassium can lead to a decrease in membrane potential of 35 mV. Electrophysiologists model the effects of ionic concentration differences, ion channels, and membrane capacitance in terms of an equivalent circuit , which is intended to represent the electrical properties of a small patch of membrane. The equivalent circuit consists of a capacitor in parallel with four pathways each consisting of a battery in series with a variable conductance.
The capacitance is determined by the properties of the lipid bilayer, and is taken to be fixed. Each of the four parallel pathways comes from one of the principal ions, sodium, potassium, chloride, and calcium. The voltage of each ionic pathway is determined by the concentrations of the ion on each side of the membrane; see the Reversal potential section above.
The conductance of each ionic pathway at any point in time is determined by the states of all the ion channels that are potentially permeable to that ion, including leakage channels, ligand-gated channels, and voltage-gated ion channels. For fixed ion concentrations and fixed values of ion channel conductance, the equivalent circuit can be further reduced, using the Goldman equation as described below, to a circuit containing a capacitance in parallel with a battery and conductance.
In electrical terms, this is a type of RC circuit resistance-capacitance circuit , and its electrical properties are very simple. For realistic situations, the time constant usually lies in the 1— millisecond range. In most cases, changes in the conductance of ion channels occur on a faster time scale, so an RC circuit is not a good approximation; however, the differential equation used to model a membrane patch is commonly a modified version of the RC circuit equation.
When the membrane potential of a cell goes for a long period of time without changing significantly, it is referred to as a resting potential or resting voltage.
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This term is used for the membrane potential of non-excitable cells, but also for the membrane potential of excitable cells in the absence of excitation. In excitable cells, the other possible states are graded membrane potentials of variable amplitude , and action potentials, which are large, all-or-nothing rises in membrane potential that usually follow a fixed time course. Excitable cells include neurons , muscle cells, and some secretory cells in glands.
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Even in other types of cells, however, the membrane voltage can undergo changes in response to environmental or intracellular stimuli. For example, depolarization of the plasma membrane appears to be an important step in programmed cell death. The interactions that generate the resting potential are modeled by the Goldman equation.
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However, it also takes into consideration the relative permeability of the plasma membrane to each ion in question. Calcium is omitted, but can be added to deal with situations in which it plays a significant role. P i stands for the relative permeability of the ion type i. In essence, the Goldman formula expresses the membrane potential as a weighted average of the reversal potentials for the individual ion types, weighted by permeability.
Although the membrane potential changes about mV during an action potential, the concentrations of ions inside and outside the cell do not change significantly. They remain close to their respective concentrations when then membrane is at resting potential. In most animal cells, the permeability to potassium is much higher in the resting state than the permeability to sodium. As a consequence, the resting potential is usually close to the potassium reversal potential. Values of resting membrane potential in most animal cells usually vary between the potassium reversal potential usually around mV and around mV.
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