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The '''Saffman–Delbrück model''' describes a lipid membrane as a thin layer of viscous fluid, surrounded by a less viscous bulk liquid.  This picture was originally proposed to determine the diffusion coefficient of membrane proteins, but has also been used to describe the dynamics of fluid domains within lipid membranes. The Saffman–Delbrück formula is often applied to determine the size of an object embedded in a membrane from its observed [[Brownian motion|diffusion coefficient]], and is characterized by the weak logarithmic dependence of diffusion constant on object radius.
[[Image:sd-illus-wiki.png|frame|Embedded cylindrical object of radius <math>a</math> in a membrane with viscosity <math>\eta_m</math>, height <math>h</math>, surrounded by bulk fluid with viscosity <math>\eta_f</math>|right]]
 
==Origin==
In a three-dimensional highly viscous liquid, a spherical object of radius ''a'' has diffusion coefficient
 
: <math>
D_{3D} = \frac{k_B T}{6 \pi \eta a}
</math>
 
by the well-known [[Stokes–Einstein relation]]. By contrast, the diffusion coefficient of a circular object embedded in a two-dimensional fluid diverges; this is [[Stokes' paradox]]. In a real lipid membrane, the diffusion coefficient may be limited by:
 
# the size of the membrane
# the inertia of the membrane (finite [[Reynolds number]])
# the effect of the liquid surrounding the membrane
 
[[Philip Saffman]] and [[Max Delbrück]] calculated the diffusion coefficient for these three cases, and showed that Case 3 was the relevant effect.<ref name="sd">[http://www.pnas.org/content/72/8/3111.abstract P. G. Saffman and M. Delbrück, ''Brownian motion in biological membranes'', Proc. Nat. Acad. Sci., USA, vol. 72 p. 3111–3113 1975]</ref>
 
==Saffman–Delbrück formula==
The diffusion coefficient of a cylindrical inclusion of radius <math>a</math> in a membrane with thickness <math>h</math> and [[viscosity]] <math>\eta_m</math>, surrounded by bulk fluid with viscosity <math>\eta_f</math> is:
 
: <math>
D_{sd} = \frac{k_B T}{4 \pi \eta_m h} \left[\ln(2 L_{sd} / a) - \gamma\right]
</math>
 
where the Saffman–Delbrück length <math>L_{sd} = \frac{h \eta_m}{2 \eta_f}</math> and <math>\gamma\approx 0.577</math> is the [[Euler–Mascheroni constant]]. Typical values of <math>L_{sd}</math> are 0.1 to 10 micrometres.<ref name="petrovschwille">[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2242757/ E.P. Petrov and P. Schwille, ''Translational Diffusion in Lipid Membranes beyond the Saffman–Delbrück Approximation'', Biophys. J. vol. 94, pL41–L43 2008]</ref> This result is an approximation applicable for radii <math>a \ll L_{sd}</math>, which is appropriate for proteins (<math>a\approx</math> nm), but not for micrometre-scale lipid domains.
 
The Saffman–Delbrück formula predicts that diffusion coefficients <math>D_{sd}</math> will only depend weakly on the size of the embedded object; for example, if <math>L_{sd} = 1 \mu m</math>, changing <math>a</math> from 1&nbsp;nm to 10&nbsp;nm only reduces the diffusion coefficient <math>D_{sd}</math> by&nbsp;30%.
 
==Beyond the Saffman–Delbrück length==
Hughes, Pailthorpe, and White extended the theory of Saffman and Delbrück to inclusions with any radii <math>a</math>;<ref name="hpw">[http://dx.doi.org/10.1017/S0022112081000785 B.D. Hughes, B.A. Pailthorpe, and L.R. White, ''The translational and rotational drag on a cylinder moving in a membrane'', J. Fluid Mech. vol. 110, p. 349–372 1981]</ref> for <math>a \gg L_{sd}</math>,
 
: <math>
D \to \frac{k_B T}{8 \eta_m h a} \frac{L_{sd}}{a}
</math>
 
A useful formula that produces the correct diffusion coefficients between these two limits is <ref name="petrovschwille"/>
 
: <math>
D = \frac{k_B T}{4 \pi \eta_m h} \left[\ln(2/\epsilon) - \gamma + 4\epsilon/\pi - (\epsilon^2/2)\ln(2/\epsilon)\right] \left[1 - (\epsilon^3/\pi) \ln(2/\epsilon) + c_1 \epsilon^{b_1} / (1 + c_2 \epsilon^{b_2}) \right]^{-1}
</math>
 
where <math>\epsilon = a / L_{sd}</math>, <math>b_1 = 2.74819</math>, <math>b_2 = 0.51465</math>, <math>c_1 = 0.73761</math>, and <math>c_2 = 0.52119</math>.
 
==Experimental studies==
Though the Saffman–Delbruck formula is commonly used to infer the sizes of nanometer-scale objects, recent experiments on proteins have suggested that the diffusion coefficient's dependence on radius <math>a</math> should be <math>a^{-1}</math> instead of <math>\ln(a)</math>.<ref name="gambin">[http://dx.doi.org/10.1073/pnas.0511026103  Y. Gambin et al., ''Lateral mobility of proteins in liquid membranes revisited'', Proc. Nat. Acad. Sci., USA, vol. 103, pp. 2098–2102, 2006]</ref> However, for larger objects (such as micrometre-scale [[lipid rafts|lipid domains]]), the Saffman–Delbruck model (with the extensions above) is well-established <ref name = "petrovschwille"/><ref name = "klingler">[http://dx.doi.org/10.1021/j100124a052 J.F. Klingler and H.M. McConnell 1993, ''Brownian motion and fluid mechanics of lipid monolayer domains'' J. Phys. Chem. vol. 93, p. 6096–6100, 1993]</ref><ref name = "cicuta">[http://dx.doi.org/10.1021/jp0702088/ P. Cicuta, S.L. Veatch, and S.L. Keller, ''Diffusion of Liquid Domains in Lipid Bilayer Membranes'' J. Phys. Chem. B, vol. 111, p. 3328–3331, 2007]</ref>
 
==References==
{{reflist}}
 
{{DEFAULTSORT:Saffman-Delbruck Model}}
[[Category:Biophysics]]
[[Category:Proteins]]
[[Category:Membrane biology]]

Revision as of 01:34, 1 February 2014

The Saffman–Delbrück model describes a lipid membrane as a thin layer of viscous fluid, surrounded by a less viscous bulk liquid. This picture was originally proposed to determine the diffusion coefficient of membrane proteins, but has also been used to describe the dynamics of fluid domains within lipid membranes. The Saffman–Delbrück formula is often applied to determine the size of an object embedded in a membrane from its observed diffusion coefficient, and is characterized by the weak logarithmic dependence of diffusion constant on object radius.

Embedded cylindrical object of radius a in a membrane with viscosity ηm, height h, surrounded by bulk fluid with viscosity ηf

Origin

In a three-dimensional highly viscous liquid, a spherical object of radius a has diffusion coefficient

D3D=kBT6πηa

by the well-known Stokes–Einstein relation. By contrast, the diffusion coefficient of a circular object embedded in a two-dimensional fluid diverges; this is Stokes' paradox. In a real lipid membrane, the diffusion coefficient may be limited by:

  1. the size of the membrane
  2. the inertia of the membrane (finite Reynolds number)
  3. the effect of the liquid surrounding the membrane

Philip Saffman and Max Delbrück calculated the diffusion coefficient for these three cases, and showed that Case 3 was the relevant effect.[1]

Saffman–Delbrück formula

The diffusion coefficient of a cylindrical inclusion of radius a in a membrane with thickness h and viscosity ηm, surrounded by bulk fluid with viscosity ηf is:

Dsd=kBT4πηmh[ln(2Lsd/a)γ]

where the Saffman–Delbrück length Lsd=hηm2ηf and γ0.577 is the Euler–Mascheroni constant. Typical values of Lsd are 0.1 to 10 micrometres.[2] This result is an approximation applicable for radii aLsd, which is appropriate for proteins (a nm), but not for micrometre-scale lipid domains.

The Saffman–Delbrück formula predicts that diffusion coefficients Dsd will only depend weakly on the size of the embedded object; for example, if Lsd=1μm, changing a from 1 nm to 10 nm only reduces the diffusion coefficient Dsd by 30%.

Beyond the Saffman–Delbrück length

Hughes, Pailthorpe, and White extended the theory of Saffman and Delbrück to inclusions with any radii a;[3] for aLsd,

DkBT8ηmhaLsda

A useful formula that produces the correct diffusion coefficients between these two limits is [2]

D=kBT4πηmh[ln(2/ϵ)γ+4ϵ/π(ϵ2/2)ln(2/ϵ)][1(ϵ3/π)ln(2/ϵ)+c1ϵb1/(1+c2ϵb2)]1

where ϵ=a/Lsd, b1=2.74819, b2=0.51465, c1=0.73761, and c2=0.52119.

Experimental studies

Though the Saffman–Delbruck formula is commonly used to infer the sizes of nanometer-scale objects, recent experiments on proteins have suggested that the diffusion coefficient's dependence on radius a should be a1 instead of ln(a).[4] However, for larger objects (such as micrometre-scale lipid domains), the Saffman–Delbruck model (with the extensions above) is well-established [2][5][6]

References

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