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Lecture 4: Diffusion and the Fokker-Planck equation. Outline: intuitive treatment Diffusion as flow down a concentration gradient Drift current and Fokker-Planck equation. Lecture 4: Diffusion and the Fokker-Planck equation. Outline: intuitive treatment
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Lecture 4: Diffusion and the Fokker-Planck equation • Outline: • intuitive treatment • Diffusion as flow down a concentration gradient • Drift current and Fokker-Planck equation
Lecture 4: Diffusion and the Fokker-Planck equation • Outline: • intuitive treatment • Diffusion as flow down a concentration gradient • Drift current and Fokker-Planck equation • examples: • No current: equilibrium, Einstein relation • Constant current, out of equilibrium:
Lecture 4: Diffusion and the Fokker-Planck equation • Outline: • intuitive treatment • Diffusion as flow down a concentration gradient • Drift current and Fokker-Planck equation • examples: • No current: equilibrium, Einstein relation • Constant current, out of equilibrium: • Goldman-Hodgkin-Katz equation • Kramers escape over an energy barrier
Lecture 4: Diffusion and the Fokker-Planck equation • Outline: • intuitive treatment • Diffusion as flow down a concentration gradient • Drift current and Fokker-Planck equation • examples: • No current: equilibrium, Einstein relation • Constant current, out of equilibrium: • Goldman-Hodgkin-Katz equation • Kramers escape over an energy barrier • derivation from master equation
Diffusion Fick’s law:
Diffusion Fick’s law: cf Ohm’s law
Diffusion Fick’s law: cf Ohm’s law conservation:
Diffusion Fick’s law: cf Ohm’s law conservation: =>
Diffusion Fick’s law: cf Ohm’s law conservation: => diffusion equation
Diffusion Fick’s law: cf Ohm’s law conservation: => diffusion equation initial condition
Diffusion Fick’s law: cf Ohm’s law conservation: => diffusion equation initial condition solution:
Diffusion Fick’s law: cf Ohm’s law conservation: => diffusion equation initial condition solution: http://www.nbi.dk/~hertz/noisecourse/gaussspread.m
Drift current and Fokker-Planck equation Drift (convective) current:
Drift current and Fokker-Planck equation Drift (convective) current: Combining drift and diffusion: Fokker-Planck equation:
Drift current and Fokker-Planck equation Drift (convective) current: Combining drift and diffusion: Fokker-Planck equation:
Drift current and Fokker-Planck equation Drift (convective) current: Combining drift and diffusion: Fokker-Planck equation: Slightly more generally, D can depend on x:
Drift current and Fokker-Planck equation Drift (convective) current: Combining drift and diffusion: Fokker-Planck equation: Slightly more generally, D can depend on x: =>
Drift current and Fokker-Planck equation Drift (convective) current: Combining drift and diffusion: Fokker-Planck equation: Slightly more generally, D can depend on x: => First term alone describes probability cloud moving with velocity u(x) Second term alone describes diffusively spreading probability cloud
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries):
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries): Stationary case:
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries): Stationary case: Gas of Brownian particles in gravitational field: u0 = μF = -μmg
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries): Stationary case: Gas of Brownian particles in gravitational field: u0 = μF = -μmg μ =mobility
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries): Stationary case: Gas of Brownian particles in gravitational field: u0 = μF = -μmg μ =mobility Boundary conditions (bottom of container, stationarity):
Examples: constant drift velocity http://www.nbi.dk/~hertz/noisecourse/gaussspreadmove.m Solution (with no boundaries): Stationary case: Gas of Brownian particles in gravitational field: u0 = μF = -μmg μ =mobility Boundary conditions (bottom of container, stationarity): drift and diffusion currents cancel
Einstein relation FP equation:
Einstein relation FP equation: Solution:
Einstein relation FP equation: Solution: But from equilibrium stat mech we know
Einstein relation FP equation: Solution: But from equilibrium stat mech we know So D = μT
Einstein relation FP equation: Solution: But from equilibrium stat mech we know So D = μT Einstein relation
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel Pumps maintain different inside and outside concentrations of ions
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field • Question: At a given Vm, what current flows through the channel?
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field • Question: At a given Vm, what current flows through the channel? x=0 x=d x inside outside
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field • Question: At a given Vm, what current flows through the channel? x=0 x=d Vout= 0 x inside outside V(x) Vm
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field • Question: At a given Vm, what current flows through the channel? x=0 x=d ρout ρin Vout= 0 x inside outside V(x) Vm
Constant current: Goldman-Hodgkin-Katz model of an (open) ion channel • Pumps maintain different inside and outside concentrations of ions • Voltage diff (“membrane potential”) between inside and outside of cell • Can vary membrane potential experimentally by adding external field • Question: At a given Vm, what current flows through the channel? x=0 x=d ρout ? ρin Vout= 0 x inside outside V(x) Vm
Reversal potential If there is no current, equilibrium => ρin/ρout=exp(-βV)
Reversal potential If there is no current, equilibrium => ρin/ρout=exp(-βV) This defines the reversal potential at which J = 0.
Reversal potential If there is no current, equilibrium => ρin/ρout=exp(-βV) This defines the reversal potential at which J = 0. For Ca++, ρout>> ρin => Vr >> 0
GHK model (2) Vm< 0: both diffusive current and drift current flow in x=0 x=d ρout ? ρin Vout= 0 x inside outside V(x) Vm
GHK model (2) Vm< 0: both diffusive current and drift current flow in Vm= 0: diffusive current flows in, no drift current x=0 x=d ρout ? ρin Vout= 0 V(x) x inside outside
GHK model (2) Vm< 0: both diffusive current and drift current flow in Vm= 0: diffusive current flows in, no drift current Vm> 0: diffusive current flows in, drift current flows out x=0 x=d ρout ? Vm V(x) ρin Vout= 0 x inside outside
GHK model (2) Vm< 0: both diffusive current and drift current flow in Vm= 0: diffusive current flows in, no drift current Vm> 0: diffusive current flows in, drift current flows out At Vm= Vr they cancel x=0 x=d ρout ? Vm V(x) ρin Vout= 0 x inside outside
GHK model (2) Vm< 0: both diffusive current and drift current flow in Vm= 0: diffusive current flows in, no drift current Vm> 0: diffusive current flows in, drift current flows out At Vm= Vr they cancel x=0 x=d ρout ? Vm V(x) ρin Vout= 0 x inside outside
Steady-state FP equation Use Einstein relation:
Steady-state FP equation Use Einstein relation: