Membrane potential

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Differences in concentration of ions on opposite sides of a cellular membrane produce a voltage difference called the membrane potential. The largest contributions usually come from sodium ( a!" and chloride (#l$" ions %hich have high concentrations in the e&tracellular region, and potassium ('!" ions, %hich along %ith large protein anions have high concentrations in the intracellular region. #alcium ions, %hich sometimes play an important role, are not sho%n. Membrane potential (or transmembrane potential" is the difference in voltage (or electrical potential difference" bet%een the interior and e&terior of a cell (Vinterior ( Vexterior". )ll animal cells are surrounded by a plasma membrane composed of a lipid bilayer %ith a variety of molecular structures embedded in it. The membrane potential arises from the interaction of ion channels and ion pumps embedded in the membrane, %hich maintain different ion concentrations on the intracellular and e&tracellular sides of the membrane. The membrane potential has t%o basic functions. First, it allo%s a cell to function as a battery, providing po%er to operate a variety of *molecular devices* embedded in the membrane. +econd, in electrically e&citable cells such as neurons, it is used for transmitting signals bet%een different parts of a cell. ,pening or closing of ion channels at one point in the membrane produces a local change in the membrane potential, %hich causes electric current to flo% rapidly to other points in the membrane. -n non.e&citable cells, and in e&citable cells in their baseline states, the membrane potential is held at a relatively stable value, called the resting potential. For neurons, typical values of the resting potential range from $/0 to $10 millivolts2 that is, the interior of a cell has a negative baseline voltage of a bit less than one tenth of a volt. ,pening and closing of ion channels can induce a departure from the resting potential, called a

depolari3ation if the interior voltage rises (say from $/0 m4 to $56 m4", or a hyperpolari3ation if the interior voltage becomes more negative (changing from $/0 m4 to $10 m4, for e&ample". -n e&citable cells, a sufficiently large depolari3ation can evoke a short.lasting all.or.nothing event called an action potential, in %hich the membrane potential very rapidly undergoes a large change, often briefly reversing its sign. )ction potentials are generated by special types of voltage.dependent ion channels. -n neurons, the factors that influence the membrane potential are diverse. They include numerous types of ion channels, some that are chemically gated and some that are voltage.gated. 7ecause voltage.dependent ion channels are controlled by the membrane potential, %hile the membrane potential itself is partly controlled by these same ion channels, feedback loops arise %hich allo% for comple& temporal dynamics, including oscillations and regenerative events such as action potentials.

Contents
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: ;hysical basis o :.: 4oltage o :.< +alts and ions in an a=ueous medium o :.> ;lasma membrane o :.? Facilitated diffusion and transport o :.6 -on pumps o :.5 -on channels :.5.: @eakage channels :.5.< @igand.gated channels :.5.> 4oltage.dependent channels o :./ Aeversal potential o :.1 B=uivalent circuit < Aesting potential > Craded potentials ? )ll other values of membrane potential 6 Bffects and implications 5 +ee also / otes 1 Aeferences D Further reading :0 B&ternal links

[edit] Physical basis

The membrane potential in a cell derives ultimately from t%o factors: electrical force and diffusion. Blectrical force arises from the mutual attraction bet%een particles %ith opposite electrical charges (positive and negative" and the mutual repulsion bet%een

particles %ith the same type of charge (both positive or both negative". Diffusion arises from the statistical tendency of particles to redistribute from regions %here they are highly concentrated to regions %here the concentration is lo%.

[edit] Voltage

Blectric field (arro%s" and contours of constant voltage created by a pair of oppositely. charged obEects. The electric field is at right angles to the voltage contours, and the field is strongest %here the spacing bet%een contours is the smallest. 4oltage, %hich is synonymous %ith electrical potential, is the ability to drive an electric current. -f a voltage source such as a battery is placed in an electrical circuit, the higher the voltage of the source, the greater the amount of current that it %ill drive. -n a functioning circuit, each point can be assigned a voltage levelFthe voltage difference bet%een any t%o points determines the amount of current that %ould flo% through a %ire hooked directly from one point to the other. -n practical electronics, the voltage difference bet%een t%o points can be measured by connecting them to the t%o leads of a volt meter (voltmeter". The functional significance of voltage lies only in voltage differencesFthe absolute value of voltage has no significance. ) volt meter can measure the voltage difference bet%een t%o locations in a circuit, but there is no instrument that can measure the voltage at a single point: the concept has no meaning. -t is conventional in electronics to assign a voltage of 3ero to some arbitrarily chosen element of the circuit, and then assign voltages for other elements on the basis of the measured or calculated voltage differences, but there is no significance in %hich element is chosen as the 3ero pointFthe function of a circuit depends only on the differences, not on voltages per se.

The same principle applies to voltage in cell biology. -n electrically active tissue, the voltage difference bet%een any t%o points can be measured by inserting an electrode at each point and connecting both electrodes to the leads of a volt meter. There is no %ay, ho%ever, to measure the voltage of a single point. Thus, a statement that the voltage difference across the membrane of a cell is 50 millivolts can be verified by placing electrodes inside and outside the cellFbut %hether the e&terior is assigned a voltage of 50 m4 and the interior 0 m4, or the e&terior is assigned a voltage of 0 m4 and the interior $50 m4, has no significance2 only the difference bet%een the t%o matters, not the absolute number assigned to either. -n mathematical terms, the definition of voltage begins %ith the concept of an electric field E, a vector field assigning a magnitude and direction to each point in space. -n many situations, the electric field is a conservative field, %hich means that it can be e&pressed as the gradient of a scalar function V, that is, E G $V. This scalar field V is referred to as the voltage distribution. ote that the definition allo%s for an arbitrary constant of integrationFthis is %hy absolute values of voltage are not meaningful. -n general electric fields can only be treated as conservative if magnetic fields do not significantly influence them, but this condition usually applies %ell to biological tissue. 7ecause the electric field is the gradient of the voltage distribution, rapid changes in voltage %ithin a small region imply a strong electric field2 conversely, if the voltage remains appro&imately the same over a large region, the electric fields in that region must be %eak. ) strong electric field, e=uivalent to a strong voltage gradient, implies that a strong force is e&erted on any charged particles that lie %ithin the region.

[edit] Salts and ions in an aqueous medium

The fluid both inside and outside of animal cells (intracellular and e&tracellular" contains a high concentration of dissolved salts. When salts dissolve in %ater, they break apart into ionsFfor e&ample sodium chloride ( a#l" breaks up almost entirely into positively charged sodium ions ( a!" and negatively charged chloride (#l$" ions. +mall ions such as sodium ( a!", potassium ('!", calcium (#a!!", and chloride (#l$" are present in high concentrations, and are capable of diffusing freely from place to place, unless some type of barrier impedes them.

[edit] Plasma membrane

The cell membrane, also called the plasma membrane or plasmalemma, is a semipermeable lipid bilayer common to all living cells. -t contains a variety of biological molecules, primarily proteins and lipids, %hich are involved in a vast array of cellular processes. Bvery animal cell is enclosed in a plasma membrane, %hich has the structure of a lipid bilayer %ith many types of large molecules embedded in it. 7ecause it is made of lipid molecules, the plasma membrane intrinsically has a high electrical resistivity, in other %ords a lo% intrinsic permeability to ions. Ho%ever, 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 %hich they can move. -n electrical terminology, the plasma membrane functions as a combined resistor and capacitor. Aesistance arises from the fact that the membrane impedes the movement of charges across it. #apacitance 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 to%ard the other side. The capacitance of the membrane is relatively unaffected by the molecules that are embedded in it, so it has a more or less invariant value estimated at about < IFJcm< (the total capacitance of a patch of membrane is proportional to its area". The conductance of a pure lipid bilayer is so lo%, on the other hand, that in biological situations it is al%ays dominated by the conductance of alternative path%ays provided by embedded molecules. Thus the capacitance of the membrane is more or less fi&ed, but the resistance is highly variable. The thickness of a plasma membrane is estimated to be about /.1 nanometers. 7ecause the membrane is so thin, it does not take a very large transmembrane voltage to create a strong electric field %ithin it. Typical membrane potentials in animal cells are on the order of :00 millivolts (that is, one tenth of a volt", but calculations sho% that this generates an electric field close to the ma&imum that the membrane can sustainFit has been calculated that a voltage difference much larger than <00 millivolts could cause dielectric breakdo%n, that is, arcing across the membrane.

[edit] Facilitated diffusion and transport

Facilitated diffusion in cell membranes, sho%ing ion channels and carrier proteins 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 t%o types of structure that play the largest roles are ion channels and ion pumps, both usually formed from assemblages of protein molecules. -on channels provide passage%ays through %hich ions can move. -n most cases an ion channel is only permeable to specific types of ions (for e&ample sodium and potassium but not chloride or calcium", and sometimes the permeability varies depending on the direction of ion movement. -on pumps, also kno%n 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.

[edit] Ion pumps

The sodium.potassium pump uses energy derived from )T; to e&change sodium for potassium ions across the membrane. ) maEor contribution to establishing the membrane potential is made by the sodium. potassium e&change pump. This is a comple& of proteins embedded in the membrane that derives energy from )T; in order to transport sodium and potassium ions across the membrane. ,n each cycle, the pump e&changes three a! ions from the intracellular space for t%o '! ions from the e&tracellular space. -f the numbers of each type of ion %ere e=ual, the pump %ould be electrically neutral, but because of the three.for.t%o e&change, it gives a net movement of one positive charge from intracellular to e&tracellular for each cycle, thereby contributing to a positive voltage difference. The pump has three effects: (:" it makes the sodium concentration high in the e&tracellular space and lo% in the intracellular space2 (<" it makes the potassium concentration high in

the intracellular space and lo% in the e&tracellular space2 (>" it gives the e&tracellular space a positive voltage %ith respect to the intracellular space. The sodium.potassium e&change pump is relatively slo% in operation. -f a cell %ere initiali3ed %ith e=ual concentrations of sodium and potassium every%here, it %ould take hours for the pump to establish e=uilibrium. The pump operates constantly, but becomes progressively less efficient as the concentrations of sodium and potassium available for pumping are reduced. )nother functionally important ion pump is the sodium.calcium e&changer. This pump operates in a conceptually similar %ay to the sodium.potassium pump, e&cept that in each cycle it e&changes three a! from the e&tracellular space for one #a!! from the intracellular space. 7ecause the net flo% of charge is in%ard, this pump runs *do%nhill*, effectively, and therefore does not re=uire any energy source e&cept the membrane voltage. -ts most important effect is to pump calcium out%ardFit also allo%s an in%ard flo% 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 e&changer is that in the resting state, intracellular calcium concentrations become very lo%.

[edit] Ion channels

)s e&plained above, a pure lipid bilayer has a very lo% permeability to ions of any type. Ho%ever, animal cell membranes contain a very diverse set of ion channels, %hich are protein structures embedded in the membrane that allo% passage of specific types of ions under specific conditions. These can be divided into three types: leakage channels, ligand.gated channels, and voltage.dependent channels. This categori3ation is not e&haustiveFit leaves out sensory receptors, many of %hich depend on ion channels that are activated by physical stimuli such as light, temperature, or stretching. [edit] ea!age channels @eakage channels are the simplest type, 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. -t should be noted that 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 %ords, they are rectifiers"2 second, some of them are capable of being shut off by chemical ligands even though they do not re=uire ligands in order to operate. [edit] igand"gated channels

@igand.gated calcium channel in closed and open states @igand.gated ion channels are channels %hose permeability is greatly increased %hen some type of chemical ligand binds to the protein structure. )nimal cells contain hundreds, if not thousands, of types of these. ) large subset function as neurotransmitter receptorsFthey occur at postsynaptic sites, and the chemical ligand that gates them is released by the presynaptic a&on terminal. ,ne e&ample of this type is the )K;) receptor, a receptor for the neurotransmitter glutamate that %hen activated allo%s passage of sodium and potassium ions. )nother e&ample is the C)7)) receptor, a receptor for the neurotransmitter C)7) that %hen activated allo%s passage of chloride ions. eurotransmitter receptors are activated by ligands that appear in the e&tracellular area, but there are other types of ligand.gated channels that are controlled by interactions on the intracellular side. [edit] Voltage"dependent channels 4oltage.gated ion channels, also kno%n as voltage dependent, are channels %hose permeability is influenced by the membrane potential. They form another very large group, %ith each member having a particular ion selectivity and a particular voltage dependence. Kany are also time.dependentFin other %ords, they do not respond immediately to a voltage change, but only after a delay. ,ne of the most important members of this group is a type of voltage.gated sodium channel that underlies action potentialsFthese are sometimes called Hodgkin-Huxley sodium channels because they %ere initially characteri3ed by )lan @loyd Hodgkin and )ndre% Hu&ley in their obel ;ri3e.%inning studies of the physiology of the action potential. The channel is closed at the resting voltage level, but opens abruptly %hen the voltage e&ceeds a certain threshold, allo%ing a large influ& of sodium ions that produces a very rapid change in the membrane potential. Aecovery from an action potential is partly dependent on a type of voltage.gated potassium channel %hich is closed at the resting voltage level but opens as a conse=uence of the large voltage change produced during the action potential. +ome voltage.dependent ion channels are also at the same time ligand.gated. ,ne of the best kno%n of these is the KD) receptor, a type of calcium channel that is gated by the

neurotransmitter glutamate but also re=uires the membrane potential to be elevated substantially above baseline in order to open.

[edit] #e$ersal potential

The reversal potential (or equilibrium potential" of an ion is the value of transmembrane voltage at %hich diffusive and electrical forces counterbalance, so that there is no net ion flo% across the membrane. This means that the transmembrane voltage e&actly opposes the force of diffusion of the ion, such that the net current of the ion across the membrane is 3ero and unchanging. The reversal potential is important because it gives the voltage that acts on channels permeable to that ionFin other %ords, it gives the voltage that the ion concentration gradient generates %hen it acts as a battery. The e=uilibrium potential of a particular ion is usually designated by the notation Eion.The e=uilibrium potential for any ion can be calculated using the ernst e=uation.8:9 For e&ample, reversal potential for potassium ions %ill be as follo%s:

%here

Ee=,'! is the e=uilibrium potential for potassium, measured in volts R is the universal gas constant, e=ual to 1.>:? EoulesL'(:Lmol(: T is the absolute temperature, measured in kelvins (G ' G degrees #elsius ! </>.:6" z is the number of elementary charges of the ion in =uestion involved in the reaction is the Faraday constant, e=ual to D5,?16 coulombsLmol(: or JL4(:Lmol(: 8'!9o is the e&tracellular concentration of potassium, measured in molLm(> or mmolLl(: 8'!9i is the intracellular concentration of potassium

Bven if t%o different ions have the same charge (i.e. '! and a!", they can still have very different e=uilibrium potentials, provided their outside andJor inside concentrations differ. Take, for e&ample, the e=uilibrium potentials of potassium and sodium in neurons. The potassium e=uilibrium potential E' is .1? m4 %ith 6 mK potassium outside and :?0 mK inside. The sodium e=uilibrium potential, on the other hand, E a is appro&imately !?0 m4 %ith appro&imately :< mK sodium inside and :?0 mK outside.8note :9

[edit] Equi$alent circuit

B=uivalent circuit for a patch of membrane, consisting of a fi&ed capacitance in parallel %ith four path%ays each containing a battery in series %ith a variable conductance Blectrophysiologists model the effects of ionic concentration differences, ion channels, and membrane capacitance in terms of an e=uivalent circuit, %hich is intended to represent the electrical properties of a small patch of membrane. The e=uivalent circuit consists of a capacitor in parallel %ith four path%ays each consisting of a battery in series %ith a variable conductance. The capacitance is determined by the properties of the lipid bilayer, and is taken to be fi&ed. Bach of the four parallel path%ays comes from one of the principal ions, sodium, potassium, chloride, and calcium. The voltage of each ionic path%ay is determined by the concentrations of the ion on each side of the membrane2 see the Aeversal potential section above. The conductance of each ionic path%ay 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.dependent channels.

Aeduced circuit obtained by combining the ion.specific path%ays using the Coldman e=uation For fi&ed ion concentrations and fi&ed values of ion channel conductance, the e=uivalent circuit can be further reduced, using the Coldman e=uation as described belo%, to a circuit containing a capacitance in parallel %ith a battery and conductance. Blectrically this is a type of A# circuit (resistance.capacitance circuit", and its electrical properties are very simple. +tarting from any initial state, the current flo%ing across either the conductance or capacitance decays %ith an e&ponential time course, %ith a time constant of M G A#, %here # is the capacitance of the membrane patch, and A G :Jgnet is the net resistance. For realistic situations the time constant usually lies in the :F:00 millisecond range. -n most cases changes in the conductance of ion channels occur on a faster time scale, so an A# circuit is not a good appro&imation2 ho%ever the differential e=uation used to model a membrane patch is commonly a modified version of the A# circuit e=uation.

[edit] #esting potential

When the membrane potential of a cell can go for a long period of time %ithout changing significantly, it is referred to as a resting potential or resting voltage. This term is used for the membrane potential of non.e&citable cells, but also for the membrane potential of e&citable cells in the absence of e&citation. -n e&citable cells, the other possible states are graded membrane potentials (of variable amplitude", and action potentials, %hich are large, all.or.nothing rises in membrane potential that usually follo% a fi&ed time course. B&citable cells include neurons, muscle cells, and some secretory cells in glands. Bven in other types of cells, though, the membrane voltage can undergo changes in response to environmental or intracellular stimuli. For e&ample, depolari3ation of the plasma membrane appears to be an important step in programmed cell death.8<9 The interactions that generate the resting potential are modeled by the Coldman e=uation. 8>9 This is similar in form to the ernst e=uation sho%n above, in that it is based on the charges of the ions in =uestion, as %ell as the difference bet%een their inside and outside concentrations. Ho%ever, it also takes into consideration the relative permeability of the plasma membrane to each ion in =uestion.

The three ions that appear in this e=uation are potassium ('!", sodium ( a!", and chloride (#l(". #alcium is omitted, but can be added to deal %ith situations in %hich it plays a significant role.8?9 7eing an anion, the chloride terms are treated differently than the cation terms2 the intracellular concentration is in the numerator, and the e&tracellular concentration in the denominator, %hich is reversed from the cation terms. !i stands for the relative permeability of the ion type i. The Coldman formula essentially e&presses the membrane potential as a %eighted average of the reversal potentials for the individual ion types, %eighted by permeability. -n most animal cells, the permeability to potassium is much higher in the resting state than the permeability to sodium. #onse=uently, the resting potential is usually close to the potassium reversal potential.869859 The permeability to chloride can be high enough to be significant, but unlike the other ions, chloride is not actively pumped, and therefore e=uilibrates at a reversal potential very close to the resting potential determined by the other ions. 4alues of resting membrane potential in the most animal cells usually vary bet%een the potassium reversal potential (usually around .10 m4" and around .?0 m4. The resting potential in e&citable cells (capable of producing action potentials" is usually near .50 m4Fmore depolari3ed voltages %ould lead to spontaneous generation of action potentials. -mmature or undifferentiated cells sho% highly variable values of resting voltage, usually significantly more positive than in differentiated cells.8/9 -n such cells, the

resting potential value correlates %ith the degree of differentiation: undifferentiated cells in some cases may not sho% any transmembrane voltage difference at all. Kaintenance of the resting potential can be metabolically costly for a cell because of its re=uirement for active pumping of ions to counteract losses due to leakage channels. The cost is highest %hen the cell function re=uires an especially depolari3ed value of membrane voltage. For e&ample, the resting potential in daylight.adapted blo%fly ("alliphora vicina" photoreceptors can be as high as .>0 m4.819 This elevated membrane potential allo%s the cells to respond very rapidly to visual inputs2 the cost is that maintenance of the resting potential may consume more than <0N of overall cellular )T;.8D9 ,n the other hand, the high resting potential in undifferentiated cells can be a metabolic advantage. This apparent parado& is resolved by e&amination of the origin of that resting potential. @ittle.differentiated cells are characteri3ed by e&tremely high input resistance8/9 %hich implies that fe% leakage channels are present at this stage of cell life. )s an apparent result, potassium permeability becomes similar to that for sodium ions, %hich places resting potential in.bet%een the reversal potentials for sodium and potassium as discussed above. The reduced leakage currents also mean there is little need for active pumping in order to compensate, therefore lo% metabolic cost.

[edit] %raded potentials

)s e&plained above, the potential at any point in a cellOs membrane is determined by the ion concentration differences bet%een the intracellular and e&tracellular areas, and by the permeability of the membrane to each type of ion. The ion concentrations do not normally change very =uickly (%ith the e&ception of calcium, %here the baseline intracellular concentration is so lo% that even a small inflo% may increase it by orders of magnitude", but the permeabilities can change in a fraction of a millisecond, as a result of activation of ligand.gated or voltage.gated ion channels. The change in membrane potential can be large or small, depending on ho% many ion channels are activated and %hat type they are. #hanges of this type are referred to as graded potentials, in contrast to action potentials, %hich have a fi&ed amplitude and time course. )s can be derived from the Coldman e=uation sho%n above, the effect of increasing the permeability for a particular type of ion is to shift the membrane potential to%ard the reversal potential for that ion. Thus, opening sodium channels pulls the membrane potential to%ard the sodium reversal potential, usually around !:00 m4. ,pening potassium channels pulls the membrane potential to%ard about .D0 m42 opening chloride channels pulls it to%ard about ./0 m4. 7ecause .D0 to !:00 m4 is the full operating range of membrane potential, the effect is that sodium channels al%ays pull the membrane potential up, potassium channels pull it do%n, and chloride channels pull it to%ard the resting potential.

Craph displaying an B;+;, an -;+;, and the summation of an B;+; and an -;+;. Craded membrane potentials are particularly important in neurons, %here they are produced by synapsesFa temporary rise or fall in membrane potential produced by activation of a synapse is called a postsynaptic potential. eurotransmitters that act to open sodium channels cause the membrane potential to rise, %hile neurotransmitters that act on potassium channels cause it to fall. 7ecause the membrane potential in a neuron must rise past the threshold value to produce an action potential, a rise in membrane potential is e&citatory, %hile a fall is inhibitory. Thus neurotransmitters that act to open sodium channels produce a so.called e&citatory postsynaptic potential, or B;+;, %hereas neurotransmitters that act to open potassium channels produce an inhibitory postsynaptic potential, or -;+;. When multiple types of channels are open %ithin the same time period, their postsynaptic potentials summate.

[edit] &ll other $alues of membrane potential

From the vie%point of biophysics, the resting membrane potential is merely the membrane potential that results from the membrane permeabilities that predominate %hen the cell is resting. The above e=uation of %eighted averages al%ays applies, but the follo%ing approach may be more easily visuali3ed. )t any given moment, there are t%o factors for an ion that determine ho% much influence that ion %ill have over the membrane potential of a cell. :. That ionOs driving force and, <. That ionOs permeability -ntuitively, this is easy to understand. -f the driving force is high, then the ion is being *pushed* across the membrane hard (more correctly stated: it is diffusing in one direction faster than the other". -f the permeability is high, it %ill be easier for the ion to diffuse across the membrane. 7ut %hat are Odriving forceO and OpermeabilityOP

Driving force: the driving force is the net electrical force available to move that ion across the membrane. -t is calculated as the difference bet%een the voltage that the ion *%ants* to be at (its e=uilibrium potential" and the actual membrane potential (Em". +o formally, the driving force for an ion G Em . Eion

For e&ample, at our earlier calculated resting potential of (/> m4, the driving force on potassium is / m4 : ((/> m4" ( ((10 m4" G / m4. The driving force on sodium %ould be ((/> m4" ( (50 m4" G (:>> m4. ;ermeability: is simply a measure of ho% easily an ion can cross the membrane. -t is normally measured as the (electrical" conductance and the unit, siemens, corresponds to : #Ls(:L4(:, that is one charge per second per volt of potential.

+o in a resting membrane, %hile the driving force for potassium is lo%, its permeability is very high. +odium has a huge driving force, but almost no resting permeability. -n this case, potassium carries about <0 times more current than sodium, and thus has <0 times more influence over Em than does sodium. Ho%ever, consider another caseFthe peak of the action potential. Here permeability to a is high and ' permeability is relatively lo%. Thus the membrane moves to near E a and far from E'. The more ions are permeant, the more complicated it becomes to predict the membrane potential. Ho%ever, this can be done using the Coldman.Hodgkin.'at3 e=uation or the %eighted means e=uation. 7y simply 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 CH' e=uations says, basically, is that at any time, the value of the membrane potential %ill be a %eighted average of the e=uilibrium potentials of all permeant ions. The *%eighting* is the ions relative permeability across the membrane.

[edit] Effects and implications

While cells e&pend 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 )T;, %hich is the common currency of biological energy. #ells may dra% on the energy they store in the resting potential to drive action potentials or other forms of e&citation. These changes in the membrane potential enable communication %ith other cells (as %ith action potentials" or initiate changes inside the cell, %hich happens in an egg %hen it is fertili3ed by a sperm. -n neuronal cells, an action potential begins %ith a rush of sodium ions into the cell through sodium channels, resulting in depolari3ation, %hile recovery involves an out%ard rush of potassium through potassium channels. 7oth these flu&es occur by passive diffusion.

[edit] See also

)ction potential Blectrochemical potential Coldman B=uation +altatory conduction Kembrane biophysics +ignal (biology"

[edit] 'otes
:. ( ote that the sign of E a and E' are opposite. This is because the concentration gradient for potassium is directed out of the cell, %hile the concentration gradient for sodium is directed into the cell. Kembrane potentials are defined relative to the e&terior of the cell2 thus, a potential of (/0 m4 implies that the interior of the cell is negative relative to the e&terior.

[edit] #eferences
:. <. >. ?. 6. ( ;urves et al#, pp. <1$><2 7ullock, ,rkand, and Crinnell, pp. :>>$:>?2 +chmidt. ielsen, pp. ?/1$?10, 6D5$6D/2 Junge, pp. >>$>6 ( Franco A, 7ortner #D, #idlo%ski J) (January <005". *;otential roles of electrogenic ion transport and plasma membrane depolari3ation in apoptosis*. $# %embr# &iol# )*+ (:": ?>$61. doi::0.:00/Js00<><.006.01>/.6. ;K-D :5516500. ( ;urves et al#, pp. ><$>>2 7ullock, ,rkand, and Crinnell, pp. :>1$:?02 +chmidt. ielsen, pp. ?102 Junge, pp. >6$>/ ( +pangler +C (:D/<". *B&pansion of the constant field e=uation to include both divalent and monovalent ions*. 'la $ %ed (ci + (<": <:1$<>. ;K-D 60?60?:. ( ;urves et al#, p. >?2 7ullock, ,rkand, and Crinnell, p. :>?2 +chmidt. ielsen, pp. ?/1$?10. ( ;urves et al#, pp. >>$>52 7ullock, ,rkand, and Crinnell, p. :>:. Q a b Kagnuson D+, Korassutti DJ, +taines W), Kc7urney KW, Karshall '#. (:DD6 Jan :?". *-n vivo electrophysiological maturation of neurons derived from a multipotent precursor (embryonal carcinoma" cell line*. &rain Res )ev &rain Res# ,- (:": :>0$?:. doi::0.:0:5J0:56.>105(D?"00:55.W. ;K-D //<0<:<. ( Juusola K, 'ouvalainen B, JRrvilehto K, WeckstrSm K. (:DD? +ep". *#ontrast gain, signal.to.noise ratio, and linearity in light.adapted blo%fly photoreceptors*. $ *en !hysiol# .*- (>": 6D>$5<:. doi::0.:016JEgp.:0?.>.6D>. ;K# <<<D<<6. ;K-D /10/05<. http:JJ%%%.pubmedcentral.nih.govJarticlerender.fcgiPtoolGpmcentre3TartidG<<<D<<6. ( @aughlin +7, de Auyter van +teveninck AA, )nderson J# (:DD1 Kay". *The metabolic cost of neural information*. +at +eurosci# . (:": >5$?:. doi::0.:0>1J<>5. ;K-D :0:D6:05.

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D.

[edit] Further reading

)lberts et al. %olecular &iology of the "ell. Carland ;ublishing2 ?th 7kT#dr edition (Karch, <00<". -+7 0.1:6>.><:1.:. Undergraduate level.

Cuyton, )rthur #., John B. Hall. Textbook of medical physiology. W.7. +aunders #ompany2 :0th edition ()ugust :6, <000". -+7 0./<:5.15//.V. Undergraduate level. Hille, 7. ,onic "hannel of Excitable %embranes +inauer )ssociates, +underland, K), U+)2 :st Bdition, :D1?. -+7 0.1/1D>.><<.0 icholls, J.C., Kartin, ).A. and Wallace, 7.C. rom +euron to &rain +inauer )ssociates, -nc. +underland, K), U+) >rd Bdition, :DD<. -+7 0.1/1D>.610.0 ,ve.+ten 'nudsen. &iological %embranes- Theory of Transport. !otentials and Electric ,mpulses. #ambridge University ;ress (+eptember <5, <00<". -+7 0. 6<:.1:0:1.>. Craduate level. ational Kedical +eries for -ndependent +tudy. !hysiology. @ippincott Williams T Wilkins. ;hiladelphia, ;B, U+) ?th Bdition, <00:. -+7 0.5>1.>050>.0

File/Membrane potential ions en0s$g

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KembraneWpotentialWionsWen.svg (+4C file, nominally 660 X ?00 pi&els, file si3e: :16 '7" This image rendered as ; C in other si3es: <00p&, 600p&, :000p&, <000p&.

File/Cell membrane detailed diagram en0s$g

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1escription English/ The cell membrane, also called the plasma membrane or

plasmalemma, is a semipermeable lipid bilayer common to all living cells. -t contains a variety of biological molecules, primarily proteins and lipids, %hich are involved in a vast array of cellular processes. -t also serves as the attachment point for both the intracellular cytoskeleton and, if present, the cell %all.

Ion channel
From Wikipedia, the free encyclopedia

(Aedirected from -on channels" Jump to: navigation, search +ot to be confused /ith- -on Television or -on implantation.

+chematic diagram of an ion channel. . . channel domains (typically four per channel", ) . outer vestibule, 2 . selectivity filter, - . diameter of selectivity filter, 3 . phosphorylation site, 4 . cell membrane. Ion channels are pore.forming proteins that help establish and control the small voltage gradient across the plasma membrane of cells (see cell potential" by allo%ing the flo% of ions do%n their electrochemical gradient.8:9 They are present in the membranes that surround all biological cells. The study of ion channels involves many scientific techni=ues such as voltage clamp electrophysiology (in particular patch clamp", immunohistochemistry, and AT.;#A.

Contents
8hide9

: 7asic features

< 7iological role > Diversity o >.: 7y gating >.:.: 4oltage.gated >.:.< @igand.gated >.:.> ,ther gating o >.< 7y ions o >.> ,ther classifications ? Detailed structure 6 Diseases of ion channels 5 History / The ion channel in fine art 1 +ee also D Aeferences :0 Further reading :: B&ternal links

[edit] 5asic features

-on channels regulate the flo% of ions across the membrane in all cells. -on channels are integral membrane proteins2 or, more typically, an assembly of several proteins. They are present on all membranes of cell (plasma membrane" and intracellular organelles (nucleus, mitochondria, endoplasmic reticulum, golgi apparatus and so on". +uch *multi. subunit* assemblies usually involve a circular arrangement of identical or homologous proteins closely packed around a %ater.filled pore through the plane of the membrane or lipid bilayer.8<98>9 For most voltage.gated ion channels, the pore.forming subunit(s" are called the Z subunit, %hile the au&iliary subunits are denoted [, \, and so on. +ome channels permit the passage of ions based solely on their charge of positive (cation" or negative (anion". Ho%ever, the archetypal channel pore is Eust one or t%o atoms %ide at its narro%est point and is selective for specific species of ion, such as sodium or potassium. These ions move through the channel pore single file nearly as =uickly as the ions move through free fluid. -n some ion channels, passage through the pore is governed by a *gate,* %hich may be opened or closed by chemical or electrical signals, temperature, or mechanical force, depending on the variety of channel.

[edit] 5iological role

7ecause *voltage.activated* channels underlie the nerve impulse and because *transmitter.activated* channels mediate conduction across the synapses, channels are especially prominent components of the nervous system. -ndeed, most of the offensive and defensive to&ins that organisms have evolved for shutting do%n the nervous systems of predators and prey (e.g., the venoms produced by spiders, scorpions, snakes, fish, bees, sea snails and others" %ork by modulating ion channel conductance andJor kinetics. -n addition, ion channels are key components in a %ide variety of biological processes that

involve rapid changes in cells, such as cardiac, skeletal, and smooth muscle contraction, epithelial transport of nutrients and ions, T.cell activation and pancreatic beta.cell insulin release. -n the search for ne% drugs, ion channels are a fre=uent target.8?9869859

[edit] 1i$ersity
There are over >00 types of ion channels in a living cell.8/9 -on channels may be classified by the nature of their gating, the species of ions passing through those gates, the number of gates (pores" and locali3ation of proteins. Further heterogeneity of ion channels arises %hen channels %ith different constitutive subunits give rise to a specific kind of current.819 )bsence or mutation of one or more of the contributing types of channel subunits can result in loss of function and, potentially, underly neurologic diseases.

[edit] 5y gating
-on channels may be classified by gating, i.e. %hat opens and closes the channels. 4oltage.gated ion channels open or close depending on the voltage gradient across the plasma membrane, %hile ligand.gated ion channels open or close depending on binding of ligands to the channel. [edit] Voltage"gated Kain article: 4oltage.gated ion channel 4oltage.gated ion channels open and close in response to membrane potential.

4oltage.gated sodium channels: This family contains at least D members and is largely responsible for action potential creation and propagation. The pore. forming Z subunits are very large (up to ?,000 amino acids" and consist of four homologous repeat domains (-.-4" each comprising si& transmembrane segments (+:.+5" for a total of <? transmembrane segments. The members of this family also coassemble %ith au&iliary [ subunits, each spanning the membrane once. 7oth Z and [ subunits are e&tensively glycosylated. 4oltage.gated calcium channels: This family contains :0 members, though these members are kno%n to coassemble %ith Z<], [, and \ subunits. These channels play an important role in both linking muscle e&citation %ith contraction as %ell as neuronal e&citation %ith transmitter release. The Z subunits have an overall structural resemblance to those of the sodium channels and are e=ually large. o #ation channels of sperm: This small family of channels, normally referred to as #atsper channels, is related to the t%o.pore channels and distantly related to TA; channels.

4oltage.gated potassium channels ('4": This family contains almost ?0 members, %hich are further divided into :< subfamilies. These channels are kno%n mainly for their role in repolari3ing the cell membrane follo%ing action potentials. The Z subunits have si& transmembrane segments, homologous to a single domain of the sodium channels. #orrespondingly, they assemble as tetramers to produce a functioning channel. +ome transient receptor potential channels: This group of channels, normally referred to simply as TA; channels, is named after their role in Drosophila phototransduction. This family, containing at least <1 members, is incredibly diverse in its method of activation. +ome TA; channels seem to be constitutively open, %hile others are gated by voltage, intracellular #a<!, pH, redo& state, osmolarity, and mechanical stretch. These channels also vary according to the ion(s" they pass, some being selective for #a<! %hile others are less selective, acting as cation channels. This family is subdivided into 5 subfamilies based on homology: classical (TA;#", vanilloid receptors (TA;4", melastatin (TA;K", polycystins (TA;;", mucolipins (TA;K@", and ankyrin transmembrane protein : (TA;)". Hyperpolari3ation.activated cyclic nucleotide.gated channels: The opening of these channels is due to hyperpolari3ation rather than the depolari3ation re=uired for other cyclic nucleotide.gated channels. These channels are also sensitive to the cyclic nucleotides c)K; and cCK;, %hich alter the voltage sensitivity of the channel^s opening. These channels are permeable to the monovalent cations '! and a!. There are ? members of this family, all of %hich form tetramers of si&. transmembrane Z subunits. )s these channels open under hyperpolari3ing conditions, they function as pacemaking channels in the heart, particularly the +) node. 4oltage.gated proton channels: 4oltage.gated proton channels open %ith depolari3ation, but in a strongly pH.sensitive manner. The result is that these channels open only %hen the electrochemical gradient is out%ard, such that their opening %ill only allo% protons to leave cells. Their function thus appears to be acid e&trusion from cells. )nother important function occurs in phagocytes (e.g. eosinophils, neutrophils, macrophages" during the *respiratory burst.* When bacteria or other microbes are engulfed by phagocytes, the en3yme )D;H o&idase assembles in the membrane and begins to produce reactive o&ygen species (A,+" that help kill bacteria. )D;H o&idase is electrogenic, moving electrons across the membrane, and proton channels open to allo% proton flu& to balance the electron movement electrically.

[edit] igand"gated Kain article: @igand.gated ion channel

)lso kno%n as ionotropic receptors, this group of channels open in response to specific ligand molecules binding to the e&tracellular domain of the receptor protein. @igand binding causes a conformational change in the structure of the channel protein that ultimately leads to the opening of the channel gate and subse=uent ion flu& across the plasma membrane. B&amples of such channels include the cation.permeable *nicotinic* )cetylcholine receptor, ionotropic glutamate.gated receptors and )T;.gated ;<V receptors, and the anion.permeable \.aminobutyric acid.gated C)7)) receptor. -on channels activated by second messengers may also be categori3ed in this group, although ligands and second messengers are other%ise distinguished from each other. [edit] 6ther gating ,ther gating include activationJinactivation by e.g. second messengers from the inside of the cell membrane, rather as from outside, as in the case for ligands. -ons may count to such second messengers, and then causes direct activation, rather than indirect, as in the case %ere the electric potential of ions cause activationJinactivation of voltage.gated ion channels.

+ome potassium channels o -n%ard.rectifier potassium channels: These channels allo% potassium to flo% into the cell in an in%ardly rectifying manner, i.e., potassium flo%s effectively into, but not out of, the cell. This family is composed of :6 official and : unofficial members and is further subdivided into / subfamilies based on homology. These channels are affected by intracellular )T;, ;-;<, and C.protein [\ subunits. They are involved in important physiological processes such as the pacemaker activity in the heart, insulin release, and potassium uptake in glial cells. They contain only t%o transmembrane segments, corresponding to the core pore. forming segments of the '4 and '#a channels. Their Z subunits form tetramers. o #alcium.activated potassium channels: This family of channels is, for the most part, activated by intracellular #a<! and contains 1 members. o T%o.pore.domain potassium channels: This family of :6 members form %hat is kno%n as leak channels, and they follo% Coldman.Hodgkin.'at3 (open" rectification. @ight.gated channels like channelrhodopsin are directly opened by the action of light. Kechanosensitive ion channels are opening under the influence of stretch, pressure, shear, displacement. #yclic nucleotide.gated channels: This superfamily of channels contains t%o families: the cyclic nucleotide.gated (# C" channels and the hyperpolari3ation.

activated, cyclic nucleotide.gated (H# " channels. -t should be noted that this grouping is functional rather than evolutionary. o #yclic nucleotide.gated channels: This family of channels is characteri3ed by activation due to the binding of intracellular c)K; or cCK;, %ith specificity varying by member. These channels are primarily permeable to monovalent cations such as '! and a!. They are also permeable to #a<!, though it acts to close them. There are 5 members of this family, %hich is divided into < subfamilies. o Hyperpolari3ation.activated cyclic nucleotide.gated channels

Temperature Cated #hannels: Kembers of the Transient Aeceptor ;otential ion channel superfamily, such as TA;4: or TA;K1 are opened either by hot or cold temperatures.

[edit] 5y ions

#hloride channels: This superfamily of poorly.understood channels consists of appro&imately :> members. They include #l#s, #@-#s, 7estrophins and #FTAs. These channels are non.selective for small anions2 ho%ever chloride is the most abundant anion, and hence they are kno%n as chloride channels. ;otassium channels o 4oltage.gated potassium channels e.g., 'vs, 'irs etc. o #alcium.activated potassium channels e.g., 7'#a or Ka&i', +', etc. o -n%ard.rectifier potassium channels o T%o.pore.domain potassium channels: This family of :6 members form %hat is kno%n as leak channels, and they follo% Coldman.Hodgkin.'at3 (open" rectification. +odium channels o voltage.gated sodium channels a4s o epithelial sodium channels (B a#" #alcium channels #a4s ;roton channels o 4oltage.gated proton channels +on-selective cation channels: These let many types of cations, mainly a!, '! and #a<! through the channel. o Kost Transient receptor potential channels

[edit] 6ther classifications

There are other types of ion channel classifications that are based on less normal characteristics, e.g. multiple pores and transient potentials. )lmost all ion channels have one single pore. Ho%ever, there are also those %ith t%o:

T%o.pore channels: This small family of < members putatively forms cation. selective ion channels. They are predicted to contain t%o '4.style si&. transmembrane domains, suggesting they form a dimer in the membrane. These channels are related to catsper channels channels and, more distantly, TA; channels.

There are channels that are classified by the duration of the response to stimuli:

Transient receptor potential channels: This group of channels, normally referred to simply as TA; channels, is named after their role in Drosophila phototransduction. This family, containing at least <1 members, is incredibly diverse in its method of activation. +ome TA; channels seem to be constitutively open, %hile others are gated by voltage, intracellular #a<!, pH, redo& state, osmolarity, and mechanical stretch. These channels also vary according to the ion(s" they pass, some being selective for #a<! %hile others are less selective, acting as cation channels. This family is subdivided into 5 subfamilies based on homology: canonical (TA;#", vanilloid receptors (TA;4", melastatin (TA;K", polycystins (TA;;", mucolipins (TA;K@", and ankyrin transmembrane protein : (TA;)".

[edit] 1etailed structure

#hannels differ %ith respect to the ion they let pass (for e&ample, a!, '!, #l(", the %ays in %hich they may be regulated, the number of subunits of %hich they are composed and other aspects of structure. #hannels belonging to the largest class, %hich includes the voltage.gated channels that underlie the nerve impulse, consists of four subunits %ith si& transmembrane helices each. ,n activation, these helices move about and open the pore. T%o of these si& helices are separated by a loop that lines the pore and is the primary determinant of ion selectivity and conductance in this channel class and some others. The e&istence and mechanism for ion selectivity %as first postulated in the :D50s by #lay )rmstrong.8D9 He suggested that the pore lining could efficiently replace the %ater molecules that normally shield potassium ions, but that sodium ions %ere too small to allo% such shielding, and therefore could not pass through. This mechanism %as finally confirmed %hen the structure of the channel %as elucidated. The channel subunits of one such other class, for e&ample, consist of Eust this *;* loop and t%o transmembrane helices. The determination of their molecular structure by Aoderick Kac'innon using V. ray crystallography %on a share of the <00> obel ;ri3e in #hemistry. 7ecause of their small si3e and the difficulty of crystalli3ing integral membrane proteins for V.ray analysis, it is only very recently that scientists have been able to directly e&amine %hat channels *look like.* ;articularly in cases %here the crystallography

re=uired removing channels from their membranes %ith detergent, many researchers regard images that have been obtained as tentative. )n e&ample is the long.a%aited crystal structure of a voltage.gated potassium channel, %hich %as reported in Kay <00>. 8:098::9 ,ne inevitable ambiguity about these structures relates to the strong evidence that channels change conformation as they operate (they open and close, for e&ample", such that the structure in the crystal could represent any one of these operational states. Kost of %hat researchers have deduced about channel operation so far they have established through electrophysiology, biochemistry, gene se=uence comparison and mutagenesis. #hannels can have single (#@-#s" to multiple transmembrane (' channels, ;<V receptors, a channels" domains %hich span plasma membrane to form pores. ;ore can determine the selectivity of the channel. Cate can be formed either inside or outside the pore region.

[edit] 1iseases of ion channels

There are a number of chemicals and genetic disorders %hich disrupt normal functioning of ion channels and have disastrous conse=uences for the organism. Cenetic disorders of ion channels and their modifiers are kno%n as #hannelopathies. +ee #ategory:#hannelopathy for a full list. Chemicals

Tetrodoto&in (TTV", used by puffer fish and some types of ne%ts for defense. -t blocks sodium channels. +a&ito&in, is produced by a dinoflagellate also kno%n as *red tide*. -t blocks voltage dependent sodium channels. #onoto&in, is used by cone snails to hunt prey. @idocaine and ovocaine belong to a class of local anesthetics %hich block sodium ion channels. Dendroto&in is produced by mamba snakes, and blocks potassium channels. -berioto&in is produced by the &uthus tamulus and blocks potassium channels. Heteropodato&in is produced by Heteropoda venatoria and blocks potassium channels.

%enetic

+haker gene mutations cause a defect in the voltage gated ion channels, slo%ing do%n the repolari3ation of the cell. B=uine hyperkalaemic periodic paralysis as %ell as Human hyperkalaemic periodic paralysis (Hyper;;" are caused by a defect in voltage dependent sodium channels. ;aramyotonia congenita (;#" and potassium aggravated myotonias (;)K" Cenerali3ed epilepsy %ith febrile sei3ures plus (CBF+!"

Bpisodic )ta&ia (B)", characteri3ed by sporadic bouts of severe discoordination %ith or %ithout myokymia, and can be provoked by stress, startle, or heavy e&ertion such as e&ercise. Familial hemiplegic migraine (FHK" +pinocerebellar ata&ia type :> @ong `T syndrome is a ventricular arrhythmia syndrome caused by mutations in one or more of presently ten different genes, most of %hich are potassium channels and all of %hich affect cardiac repolari3ation. 7rugada syndrome is another ventricular arrhythmia caused by voltage.gated sodium channel gene mutations. #ystic fibrosis is caused by mutations in the #FTA gene, %hich is a chloride channel. Kucolipidosis type -4 is caused by mutations in the gene encoding the TA;K@: channel

[edit] 7istory
The fundamental properties of currents mediated by ion channels %ere analy3ed by the 7ritish biophysicists )lan Hodgkin and )ndre% Hu&ley as part of their obel ;ri3e. %inning research on the action potential, published in :D6<. They built on the %ork of other physiologists, such as #ole and 7akerOs research into voltage.gated membrane pores from :D?:.8:<98:>9 The e&istence of ion channels %as confirmed in the :D/0s by 7ernard 'at3 and Aicardo Kiledi using noise analysis. -t %as then sho%n more directly %ith an electrical recording techni=ue kno%n as the *patch clamp*, %hich led to a obel ;ri3e to Br%in eher and 7ert +akmann, the techni=ueOs inventors. Hundreds if not thousands of researchers continue to pursue a more detailed understanding of ho% these proteins %ork. -n recent years the development of automated patch clamp devices helped to increase significantly the throughput in ion channel screening. The obel ;ri3e in #hemistry for <00> %as a%arded to t%o )merican scientists: Aoderick Kac'innon for his studies on the physico.chemical properties of ion channel structure and function, including &.ray crystallographic structure studies, and ;eter )gre for his similar %ork on a=uaporins.8:?9

[edit] 8he ion channel in fine art

&irth of an ,dea (<00/" by Julian 4oss.)ndreae. The sculpture %as commissioned by Aoderick Kac'innon based on the moleculeOs atomic coordinates that %ere determined by Kac'innonOs group in <00:. Aoderick Kac'innon commissioned &irth of an ,dea, a 6O (:.60 m" tall sculpture based on the 'cs) potassium channel.8:69 The art%ork contains a %ire obEect representing the pore liner %ith a blo%n glass obEect representing the main cavity of the channel structure.

[edit] See also

)ction potential )ctive transport #hannelome #hannelomics #hannelopathy Cating (electrophysiology" -on channel family as defined in ;fam and -nter;ro 'i Database Kagnesium transport euroto&in ;assive transport +ynthetic ion channels Transmembrane receptor Ke+H entry for -on channels

[edit] #eferences

:. <.

>.

?. 6. 5. /.

1. D.

:0. ::. :<.

:>.

( Hille, 7ertil (<00:". ,on channels of excitable membranes (third ed.". +underland, Kass: +inauer )ssociates. -+7 0.1/1D>.><:.<. ( ;urves, (<00:". *#hapter ?: #hannels and Transporters*. -n Dale ;urves, Ceorge J. )ugustine, David Fit3patrick, @a%rence. #. 'at3, )nthony.+amuel @aKantia, James ,. Kc amara, +. Kark Williams, editors. +euroscience (<nd ed.". +inauer )ssociates -nc.. -+7 0.1/1D>./?:.<. http:JJ%%%.ncbi.nlm.nih.govJbooksJbv.fcgiP ridGneurosci.chapter.<</. ( Hille 7, #atterall, W) (:DDD". *#hapter 5: Blectrical B&citability and -on #hannels*. -n Ceorge J +iegel, 7ernard W )granoff, A. W )lbers, +tephen ' Fisher and Kichael D Uhler. &asic neurochemistry- molecular. cellular. and medical aspects . ;hiladelphia: @ippincott.Aaven. -+7 0.>D/.6:1<0.V. http:JJ%%%.ncbi.nlm.nih.govJbooksJbv.fcgiPridGbnchm.chapter.?<:. ( #amerino D#, Tricarico D, Desaphy JF ()pril <00/". *-on channel pharmacology*. +eurotherapeutics - (<": :1?$D1. doi::0.:0:5JE.nurt.<00/.0:.0:>. ;K-D :/>D6:<1. ( 4erkman )+, Calietta @J (February <00D". *#hloride channels as drug targets*. +at Rev )rug )iscov , (<": :6>$/:. doi::0.:0>1Jnrd</10. ;K-D :D:6>661. ( #amerino D#, Desaphy JF, Tricarico D, ;ierno +, @iantonio ) (<001". *Therapeutic approaches to ion channel diseases*. 'dv# *enet# 4-: 1:$:?6. doi::0.:0:5J+0056.<550(01"0010?.>. ;K-D :D:5:1>>. ( Cabashvili -+, +okolo%ski 7H, Korton ##, Ciersch )7 (+eptember <00/". *-on channel gene e&pression in the inner ear*. $# 'ssoc# Res# 0tolaryngol# , (>": >06$<1. doi::0.:00/Js:0:5<.00/.001<.y. ;K# <6>1?>/. ;K-D :/6?:/5D. http:JJ%%%.pubmedcentral.nih.govJarticlerender.fcgiPtoolGpmcentre3TartidG<6>1?>/. ( 4icini + ()pril :DDD". * e% perspectives in the functional role of C)7)()" channel heterogeneity*. %ol# +eurobiol# .+ (<": D/$::0. doi::0.:00/J7F0</?>565. ;K-D :0>/:?56. ( 7e3anilla F, )rmstrong #K ( ovember :D/<". * egative conductance caused by entry of sodium and cesium ions into the potassium channels of s=uid a&ons*. $# *en# !hysiol# 4* (6": 611$501. doi::0.:016JEgp.50.6.611. ;K# <<<50D:. ;K-D ?5??></. http:JJ%%%.pubmedcentral.nih.govJarticlerender.fcgiPtoolGpmcentre3TartidG<<<50D:. ( Jiang Y, @ee ), #hen J, Auta 4, #adene K, #hait 7T, Kac'innon A (Kay <00>". *V.ray structure of a voltage.dependent '! channel*. +ature -)2 (5D>6": >>$?:. doi::0.:0>1Jnature0:610. ;K-D :</<:5:1. ( The detailed >D structure of the magnesium channel from bacteria can be seen here 8:9. ( Aonald ;ethig and Douglas 7 'ell (:D1/". *The passive electrical properties of biological systems: their significance in physiology, biophysics and biotechnology.*. !hys# %ed# &iol# 2) (1": D>>$D/0. doi::0.:011J00>:.D:66J><J1J00:. ;K-D >>05/<:. http:JJdbkgroup.orgJ;apersJpethigWkellWpmb1/.pdf. 8)n e&pansive revie% of bioelectrical characteristics from :D1/. @ay summary9. *... the observation of an inductance (negative capacitance" by #ole and 7aker (:D?:" during measurements of the )# electrical properties of s=uid a&ons led directly to the concept of voltage.gated membrane pores, as embodied in the celebrated Hodgkin.Hu&ley (:D6<" treatment (#ole :D/<, Jack er a: :D/6", as the crucial mechanism of neurotransmission.*. ( 'enneth +. #ole and Aichard F. 7aker (February :D?:". *@ongitudinal impedance of the s=uid giant a&on.*. The $ournal of *eneral !hysiology (The Aockefeller University ;ress" )- (5": //:. doi::0.:016JEgp.<?.5.//:. ;K# <<>100/. ;K-D :D1/><6<. http:JJEgp.rupress.orgJcgiJreprintJ<?J5J//:. 8Describes %hat happens %hen you stick a giant s=uid a&on %ith electrodes and pass through an alternating current, and then notice that sometimes the voltage rises %ith time, and sometimes it

decreases. @ay summary9. *The inductive reactance is a property of the a&on and re=uires that it contain an inductive structure. The variation of the impedance %ith interpolar distance indicates that the inductance is in the membrane*. :?. ( The Aoyal +%edish )cademy of +cience (<00>.:0.01". *The obel ;ri3e in #hemistry <00>*. ;ress release. http:JJnobelpri3e.orgJnobelWpri3esJchemistryJlaureatesJ<00>Jpress.html. Aetrieved <0:0. 0:.:1. :6. ( 7all, ;hilip (Karch <001". *The crucible: )rt inspired by science should be more than Eust a pretty picture*. "hemistry 1orld 3 (>": ?<$?>. http:JJ%%%.rsc.orgJchemistry%orldJ-ssuesJ<001JKarchJ#olumnThecrucible.asp. Aetrieved <00D.0:.:<.

[edit] Further reading

)n -ntervie% %ith Aoderick Kac'innon Freevie% video by the 4ega +cience Trust. Doyle D), Korais #abral J, ;fuet3ner A), 'uo ), Culbis JK, #ohen +@, #hait 7T, Kac'innon A ()pril :DD1". *The structure of the potassium channel: molecular basis of '! conduction and selectivity*. (cience ),* (6>50": 5D$//. doi::0.::<5Jscience.<10.6>50.5D. ;K-D D6<616D. http:JJ%%%.cellbio.%ustl.eduJfacultyJhuettnerJ5D.pdf. 7e3anilla F ()pril <000". *The voltage sensor in voltage.dependent ion channels*. !hysiol# Rev# ,* (<": 666$D<. ;K-D :0/?/<0:. http:JJphysrev.physiology.orgJcgiJcontentJabstractJ10J<J666. +ingh H (<0:0". *T%o decades %ith dimorphic #hloride -ntracellular #hannels (#@-#s".*. ebs letters# 3,- (:0": <::<$<:. doi::0.:0:5JE.febslet.<0:0.0>.0:>. ;K-D <0<<5/1>. #atterall W) (<0:0". *-on channel voltage sensors: structure, function, and pathophysiology.*. +euron 49 (5": D:6$D<1. doi::0.:0:5JE.neuron.<0:0.01.0<:. ;K# <D601<D. ;K-D <015D6D0. http:JJ%%%.pubmedcentral.nih.govJarticlerender.fcgiP toolGpmcentre3TartidG<D601<D. 'athy @is3e%ski (<00/.05.:6". *,pening the Cates on -on #hannel Drugs*. *enetic Engineering 2 &iotechnology +e/s (Kary )nn @iebert, -nc.": pp. :1, <0. http:JJ%%%.genengne%s.comJarticlesJchitem.asp&PaidG<:6D. Aetrieved <001.0/.05. *#onditions targeted include diabetes, cystic fibrosis, hypertension, pain and cancer* +hahidul, -slam (January <0::". Transient Receptor !otential "hannels. )dvances in B&perimental Kedicine and 7iology. 9*-. 7erlin: +pringer. p. /00. -+7 D/1.D?.00/. 0<5?.5.

[edit] E:ternal lin!s

Wikiversity has learning materials about PoissonBoltzmann profile for an ion channel *4oltage.Cated -on #hannels*. ,3!H'R )atabase of Receptors and ,on "hannels. -nternational Union of 7asic and #linical ;harmacology. http:JJ%%%.iuphar.db.orgJD)T)7)+BJAeceptorFamiliesFor%ardPtypeG-#. *TA-; Database*. a manually curated database of protein-protein interactions for mammalian TR! channels. http:JJ%%%.trpchannel.org.

File/8urgor pressure on plant cells diagram0s$g

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1escription -n biology, turgor pressure or turgidity is the pressure of the cell contents against the cell %all, in plant cells, determined by the %ater content of the vacuole, resulting from osmotic pressure.

Plasmolysis
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7efore plasmolysis, and after. Plasmolysis is the process in plant cells %here the plasma membrane pulls a%ay from the cell %all due to the loss of %ater through osmosis. The reverse process, cytolysis, can occur if the cell is in a hypotonic solution resulting in a higher e&ternal osmotic pressure and a net flo% of %ater into the cell. Through observation of plasmolysis and deplasmolysis it is possible to determine the tonicity of the cellOs environment as %ell as the rate solute molecules cross the cellular membrane.

Contents
8hide9

: Turgidity < ;lasmolysis > +ee also ? Aeferences 6 B&ternal links

[edit] 8urgidity
Kain article: Turgor ;ressure ) plant cell in a more hypotonic solution %ill absorb %ater by endosmosis, so that the increased volume of %ater in the cell %ill increase pressure, making the protoplasm push against the cell %all, a condition kno%n as turgor. Turgor makes plant cells push against each other in the same %ay and is the main line method of support in non.%oody plant

tissue. ;lant cell %alls resist further %ater entry after a certain point, kno%n as full turgor, %hich stops plant cells from bursting as animal cells do in the same conditions. This is also the reason that plants stand upright. Without the stiffness of the plant cells the plant %ould fall under its o%n %eight. This is the same reason that %hen %e pour %ater into a potted plant it perks up.

[edit] Plasmolysis

;lant cell under different environments -f a plant cell is placed in a hypertonic solution, the plant cell loses %ater and hence turgor pressure, making the plant cell flaccid. ;lants %ith cells in this condition %ilt. Further %ater loss causes plasmolysis: pressure decreases to the point %here the protoplasm of the cell peels a%ay from the cell %all, leaving gaps bet%een the cell %all and the membrane. Bventually cytorrhysis $ the complete collapse of the cell %all $ can occur. There are some mechanisms in plants to prevent e&cess %ater loss in the same %ay as e&cess %ater gain, but plasmolysis can be reversed if the cell is placed in a %eaker solution (hypotonic solution". +tomata help keep %ater in the plant so it does not dry out. Wa& also keeps %ater in the plant. The e=uivalent process in animal cells is called crenation. The li=uid content of the cell leaks out due to diffusion. The cell collapse and cell membrane pulls a%ay from the cell %all(in plants". Kost animal cells consist of only a phospholipid bilayer and not a cell %all, therefore shrinking up under such conditions. ;lasmolysis only occurs in e&treme conditions and rarely happens in nature. -t is induced in the laboratory by immersing cells in strong saline or sugar solutions to cause e&osmosis, often using Blodea plants or onion epidermal cells, %hich have coloured cell sap so that the process is clearly visible. ;lasmolysis can be of t%o types. -t can be either concave plasmolysis or conve& plasmolysis. #onve& plasmolysis is al%ays irreversible %hile concave plasmolysis is usually reversible.8citation needed9

[edit] See also

#renation #ytolysis, %here the cell bursts rather than shrinks.

Cytolysis
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blood cells under different solutions Cytolysis, or osmotic lysis, occurs %hen a cell bursts due to an osmotic imbalance that has caused e&cess %ater to move into the cell. -t occurs in a hypotonic environment, %here %ater diffuses into the cell and causes its volume to increase. -f the volume of %ater e&ceeds the cell membraneOs capacity then the cell %ill burst.The cell %ill only burst if the cell is an animal cell and it %ill only e&pand if it is a plant cell.

Contents
8hide9

: -n ;lants < -n Kammals > -n 7acteria ? ;revention 6 +ee also 5 Aeferences

[edit] In Plants
#ytolysis does not occur in plant cells because plant cells have a strong bond cell %all that contains the osmotic pressure, or turgor pressure, %hich %ould cause cytolysis not to occur. #ontrary to organisms %ithout a cell %all, plant cells must be in a hypotonic environment in order to have this turgor pressure, %hich provides the cells more structural support, preventing the plant from %ilting. -n a hypertonic environment, plasmolysis occurs, %hich is nearly the complete opposite of cytolysis: -nstead of e&panding, the cytoplasm of the plant cell retracts from the cell %all, causing the plant to %ilt.

[edit] In Mammals
,smotic lysis is often one result of a stroke, because improper nutrient perfusion and %aste removal alter cell metabolism. +uch malfunction results in an inflo% of e&tracellular fluid into the cells.

[edit] In 5acteria
,smotic lysis %ould be e&pected to occur %hen bacterial cells are treated %ith a hypotonic solution %ith added lyso3yme.

[edit] Pre$ention
Different cells and organisms have adapted different %ays of preventing cytolysis from occurring. For e&ample, the paramecium uses a contractile vacuole, %hich rapidly pumps out e&cessive %ater to prevent the build.up of %ater and the other%ise subse=uent lysis. ,ther organisms pump solutes out of their cytosol, %hich brings the solute concentration closer to that of their environment and slo%s do%n the process of %aterOs diffusion into the cell, preventing cytolysis. -f the cell can pump out enough solutes so that an isotonic environment can be achieved, there %ill be no net movement of %ater.

[edit] See also

#ell disruption @ysis ,smotic pressure ;lasmolysis #renation Water into&ication

[edit] #eferences

Kc#lendon, Jesse Francis (:D:/". !hysical "hemistry of Vital !henomena-. ,riginal from the University of #alifornia: ;rinceton University ;ress. p. <?0 pages. http:JJbooks.google.comJbooksP idG7aUa))))-))JTpgG;):?<Td=G#ytolysis!. %ikipediaTasWbrrG:TclientGfirefo&.a.

6smotic pressure
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9%orse equation9 redirects here# or the potential energy of a diatomic molecule. see %orse potential# or the functions in differential topology. see %orse theory#

,smotic pressure on red blood cells 6smotic pressure is the pressure %hich needs to be applied to a solution to prevent the in%ard flo% of %ater across a semipermeable membrane.8:9 The phenomenon of osmotic pressure arises from the tendency of a pure solvent to move through a semi.permeable membrane and into a solution containing a solute to %hich the membrane is impermeable. This process is of vital importance in biology as the cellOs membrane is selective to%ards many of the solutes found in living organisms. -n order to visuali3e this effect, imagine a U shaped clear tube %ith e=ual amounts of %ater on each side, separated by a membrane at its base that is impermeable to the sugar molecules (made from dialysis tubing". +ugar has been added to the %ater on one side. The height of the %ater on each side %ill change proportional to the pressure of the solutions. ,smotic pressure causes the height of the %ater in the compartment containing the sugar to rise, due to movement of the pure %ater from its compartment into the compartment containing the sugar %ater. This process %ill stop once the pressures of the %ater and sugar %ater to%ard both sides of the membrane are e=uated. (+ee 0smosis". Jacobus Henricus van Ot Hoff first proposed a formula for calculating the osmotic pressure, but this %as later improved upon by Harmon orthrop Korse.8citation needed9 6smotic potential is the opposite of %ater potential, %hich is the degree to %hich a solvent tends to stay in a li=uid.

Contents
8hide9

: Thermodynamic e&planation < Derivation of osmotic pressure > Korse e=uation ? )pplications o ?.: ;otential osmotic pressure 6 +ee also

5 Aeferences / B&ternal links

[edit] 8hermodynamic e:planation

#onsider the system at the point it has reached e=uilibrium. The condition for this is that the chemical potential of the solvent (since only it is free to flo% to%ards e=uilibrium" on both sides of the membrane is e=ual. The compartment containing the pure solvent has a chemical potential of b0(l,p". ,n the other side, the compartment containing the solute has an additional contribution from the solute (factored as the mole fraction of the solute, cs d :" but there also appears an addition in pressure. The balance is therefore:

%here p denotes the e&ternal pressure, l the solvent, cs the mole fraction of the solvent and e the osmotic pressure e&erted by the solutes. The addition of solute decreases the chemical potential (an entropic effect", %hile the pressure increases the chemical potential, and thus a balance is reached. ote that the presence of the solute decreases the potential due to cs being smaller than :.

[edit] 1eri$ation of osmotic pressure

-n order to find e, the osmotic pressure, %e can %rite the chemical potentials e&plicitly:

The negative e&pression on the left is a result of the increase in available states, causing an increase in entropy and a reduction of the chemical potential. The addition to the pressure is e&pressed through the e&pression for the energy of e&pansion:

-nserting the e&pression presented above into the chemical pontential e=uation for the entire system and rearranging %ill arrive at:

[edit] Morse equation

The osmotic pressure e of a dilute solution can be appro&imated using the Korse e=uation (named after Harmon orthrop Korse":8<9

e G i%RT,
%here i is the dimensionless van Ot Hoff factor % is the molarity RG1.>:??/< @ L k;a L mol.: L '.: is the gas constant T is the thermodynamic (absolute" temperature This e=uation gives the pressure on one side of the membrane2 the total pressure on the membrane is given by the difference bet%een the pressures on the t%o sides. ote the similarity of the above formula to the ideal gas la% and also that osmotic pressure is not dependent on particle charge. This e=uation %as derived by van Ot Hoff. ,smotic pressure is an important factor affecting cells. ,smoregulation is the homeostasis mechanism of an organism to reach balance in osmotic pressure.

Hypertonicity is the presence of a solution that causes cells to shrink. Hypotonicity is the presence of a solution that causes cells to s%ell. -sotonic is the presence of a solution that produces no change in cell volume.

When a biological cell is in a hypotonic environment, the cell interior accumulates %ater, %ater flo%s across the cell membrane into the cell, causing it to e&pand. -n plant cells, the cell %all restricts the e&pansion, resulting in pressure on the cell %all from %ithin called turgor pressure.

[edit] &pplications
,smotic pressure is the basis of filtering (*reverse osmosis*", a process commonly used to purify %ater. The %ater to be purified is placed in a chamber and put under an amount of pressure greater than the osmotic pressure e&erted by the %ater and the solutes dissolved in it. ;art of the chamber opens to a differentially permeable membrane that lets %ater molecules through, but not the solute particles. The osmotic pressure of ocean %ater is about </ atm. Aeverse osmosis desalinates fresh %ater from ocean salt %ater. ,smotic pressure is necessary for many plant functions. -t is the resulting turgor pressure on the cell %all that allo%s herbaceous plants to stand upright, and ho% plants regulate the aperture of their stomata. -n animal cells %hich lack a cell %all ho%ever, e&cessive osmotic pressure can result in cytolysis.

#ell %all #ytolysis

Cibbs.Donnan effect ,smosis ;feffer cell ;lasmolysis Turgor pressure

For the calculation of molecular %eight by using colligative properties, osmotic pressure is the most preferred property.

[edit] Potential osmotic pressure

;otential osmotic pressure is the ma&imum osmotic pressure that could develop in a solution if it %ere separated from distilled %ater by a selectively permeable membrane. -t is the number of solute particles in a unit volume of the solution that directly determines its potential osmotic pressure. -f one %aits for e=uilibrium, osmotic pressure reaches potential osmotic pressure.

[edit] See also

Jacobus Henricus van Ot Hoff #olligative properties

[edit] #eferences
:. ( 4oet, Donald2 Judith C. 4oet, #harlotte W. ;ratt (<00:". undamentals of &iochemistry (Aev. ed.". e% York: Wiley. p. >0. -+7 D/10?/:?:/6D0. <. ( Kansoor K. )miEi, 7everly J. +andmann (<00<". 'pplied !hysical !harmacy. KcCra%.Hill ;rofessional. pp. 6?$6/. -+7 00/:>60/5?. http:JJbooks.google.comJbooksPidG`. 4yaWi7Dcc#TpgG;)65Td=GKorse!e=uationTasWbrrG>f;;)6/,K:.

[edit] E:ternal lin!s

,smotic pressure and potential ,smotic pressure obel ;ri3e lecture on osmotic pressure and chemical e=uilibrium (pdf"