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Electrochemical Potential. Δ E i (ek) = E m – E i ( electrochemical potential ) E.g. for Na + ; -80-(+60)=-140 mV For Ca 2+ ; -80-(+129)=-209 mV For K + ; -80-(-94)=+14 mV. GIBBS-DONNAN EQUILIBRIUM POTENTIALS . GIBBS-DONNAN POTENTIAL.
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ElectrochemicalPotential • ΔEi(ek) = Em – Ei(electrochemicalpotential) • E.g. for Na+ ; -80-(+60)=-140 mV • For Ca2+ ; -80-(+129)=-209 mV • For K+ ; -80-(-94)=+14 mV
GIBBS-DONNAN POTENTIAL • is passive; that is, energy is not necessarytoitsestablishment. • Incontrast, therestingpotential is activelygeneratedbytheaction of theNa+-K+pump.
Membrane Membrane Equilibrium Initial The fluxcontinuetobecomepartialmolarfreeenergies at equilibriumforNa+andfor Cl- in bothtwosides.
Inequilibrium • G0Na + RT ln[Na+]1 + FV1 = G0Na + RT ln[Na+]2 + FV2 • G0Cl + RT ln[Cl-]1 - FV1 = G0Cl + RT ln[Cl-]2 - FV2 • [Na+]1/[Na+]2 = [Cl-]2/[Cl-]1 = r • (Donnanequation, r: Donnanratio)
Because of electroneutrality • [Na+]1 = [Cl-]1 = c1 • [Na+]2 = [Cl-]2+ [A-]
Thepotentialdifferences at equilibrium; • V = V2 – V1 =RT/zFln([Na+]1/[Na+]2) =-RT/zFln([Cl-]2/[Cl-]1) =-RT/zFlnr
At 37 °C RT/zF = -61 mV [Na+2] = [Cl-2]
[Na+]1 = 100mM + x • [Cl-]1 = x • [Na+]2 = 100mM – x • [Cl-]2 = 100mM – x ise; • [Na+]1[Cl-]1 = [Na+]2[Cl-]2 • There is a flux as itsquantity is “x”fromside 1 toside 2 andfromside 2 toside 1;
As a result; • Donnanequilibriumshowsthatthepotentialdifferencesmayarisealsobypassivemechanisms (orforces).
DiffusionPotential • It is similartoNernstPotential. • The restingpotentialarisesfromthediffusionpotential of K+.
It is alsoexplainedbyusingterm of mobility. • Mobility(μ)is a movement of an ionthrough a cellmembrane.
IonMobility(μ): • When an electricalfield is appliedacross a cellmembrane (E), ionsgain a velocitytomovethroughthiselectricalfield.
Mobilitydepends on temperatureandelectricalfield. • Furthermore, thecharactheristics of environmentandconcentration.
There is a potentialdifferencesbetweentwosides of membranebecause of thedifferences of ion’smobilities. • E.g.; …
Infigure, themobility of Cl-is higherthanthemobility of Na+. • Therefore, rightside of membrane is morenegativethanleftside of membrane.
Incase, thePotentialdifference is; RT(μNa – μCl) (c1) F (μNa + μCl) (c2) • (volt) • c1: concentration of leftside • c2: concentration of rightside ln V =
By far the most commonly used formalism for describing ion permeability and selectivity of membranes has been described by Goldman (1943) and Hodgkin & Katz (1949). • Its name is ‘constant field theory’.
Two important assumptions are that • ions cross the membrane independently (without interaction with each other), • and that the electric field in the membrane is constant (the potential drops linearly across the membrane).
Thesesimplifyingassumptionsleadtotwowidelyusedexpressions: • 1. GHK currentequation, • 2. GHK voltageequation.
1. GHK currentequation • says that the current carried by ‘ion S’ is equal to the permeability PS multiplied by a nonlinear function of voltage. • This equationallowsonetocalculatetheabsoluteionpermeabilityfrom a currentmeasurementifconcentrationandmembranepotentialareknown.
2. GHK voltageequation • Itgivesthemembranepotential at whichno net currentflows. • Forexample, it givestherestingmembranepotential in a cellwithoutelectrogenicpumps.
Thisequation is a functionthat • it contains ion fluxes, membrane potential and the ion concentrations of intracellular and extracellular fluid.
Goldman-Hodgkin-Katz, • Firstly thought that at least three ions (Na+, K+, Cl-) contribute to the membrane potential. • Then, they admit the electrical field of cell membrane as a linear and constant.
By the way, • they consider the resting membrane potential as a diffusion potential of three ions. • Instead of Mobility, • Pi = Di / δ = μi RT / zi Fδ
RT [(PK[K+]out+PNa[Na+]out+PCl[Cl-]in)] F [(PK[K+]in+PNa[Na+]in+PCl[Cl-]out)] (Goldman-Hodgkin-KatzEquation) ln EM=ERev=
If the permeability of cell membrane to one ion is higher than the others, the GHK equation is reduced to Nernst equilibrium potential for this ion.
Forexample, • the permeability of cell membrane to potassium is higher than sodium and chlor. So, • Em~ EK
As a result, • Themembranepotential of anycell is neartheNernstpotential of anyionorionswhich has orhavehigherconcentrationorhigherpermeabilityformembrane.
E.g., • Thepermeability of cellmembrane’s of squidaxonforions is: • PK:1 PNa:0.04 PCl:0.45
REVERSAL POTENTIAL • Zero-currentpotential • EM in GHK equation is alsoErev.
Also, it equals Nernst potential for one ion. • For more than one ion, it equals the average of Nernst Potentials of ions.
But, • If only one permeant ion is on each side of the membrane and both ions have the valance z, Erev is given by the simple biionicequation: RT (PA [A]0) zF (PB [B]i) Erev= ln
REFERENCES • Pehlivan F. (5. Baskı). Biyofizik. Pelikan Kitabevi. • Sperelakis N. (1995). Cell Physiology. AcademicPress. Inc. • Hille B. (2001). IonChannels of ExcitableMembranes.SinauerAssociates. Inc.