Saffman–Delbrück Model
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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 diffusion coefficient, and is characterized by the weak logarithmic dependence of diffusion constant on object radius.


Origin

In a three-dimensional highly viscous liquid, a spherical object of radius ''a'' has diffusion coefficient : D_ = \frac by the well-known
Stokes–Einstein relation In physics (specifically, the kinetic theory of gases), the Einstein relation is a previously unexpected connection revealed independently by William Sutherland in 1904, Albert Einstein in 1905, and by Marian Smoluchowski in 1906 in their works on ...
. By contrast, the diffusion coefficient of a circular object embedded in a two-dimensional fluid diverges; this is
Stokes' paradox In the science of fluid flow, Stokes' paradox is the phenomenon that there can be no creeping flow of a fluid around a disk in two dimensions; or, equivalently, the fact there is no non-trivial steady-state solution for the Stokes equations arou ...
. 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 In fluid mechanics, the Reynolds number () is a dimensionless quantity that helps predict fluid flow patterns in different situations by measuring the ratio between inertial and viscous forces. At low Reynolds numbers, flows tend to be domi ...
) # 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.P. G. Saffman and M. Delbrück, ''Brownian motion in biological membranes'', Proc. Natl. Acad. Sci. USA, vol. 72 p. 3111–3113 1975
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Saffman–Delbrück formula

The diffusion coefficient of a cylindrical inclusion of radius a in a membrane with thickness h and viscosity \eta_m, surrounded by bulk fluid with viscosity \eta_f is: : D_ = \frac \left ln(2 L_ / a) - \gamma\right where the Saffman–Delbrück length L_ = \frac and \gamma\approx 0.577 is the Euler–Mascheroni constant. Typical values of L_ are 0.1 to 10 micrometres. This result is an approximation applicable for radii a \ll L_, which is appropriate for proteins (a\approx nm), but not for micrometre-scale lipid domains. The Saffman–Delbrück formula predicts that diffusion coefficients D_ will only depend weakly on the size of the embedded object; for example, if L_ = 1 \mu m, changing a from 1 nm to 10 nm only reduces the diffusion coefficient D_ by 30%.


Beyond the Saffman–Delbrück length

Hughes, Pailthorpe, and White extended the theory of Saffman and Delbrück to inclusions with any radii a; for a \gg L_, : D \to \frac \frac = \frac A useful formula that produces the correct diffusion coefficients between these two limits is : D = \frac \left ln(2/\epsilon) - \gamma + 4\epsilon/\pi - (\epsilon^2/2)\ln(2/\epsilon)\right\left - (\epsilon^3/\pi) \ln(2/\epsilon) + c_1 \epsilon^ / (1 + c_2 \epsilon^) \right where \epsilon = a / L_, b_1 = 2.74819, b_2 = 0.51465, c_1 = 0.73761, and c_2 = 0.52119. Please note that the original version of has a typo in b_2; the value in the correction to that article should be used.


Experimental studies

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


Extending Saffman–Delbrück for Hydrodynamic Coupling of Proteins within Curved Lipid Bilayer Membranes

The Saffman–Delbrück approach has also been extended in recent works for modeling hydrodynamic interactions between proteins embedded within curved lipid bilayer membranes, such as in vesicles and other structures. These works use related formulations to study the roles of the membrane hydrodynamic coupling and curvature in the collective drift-diffusion dynamics of proteins within bilayer membranes. Various models of the protein inclusions within curved membranes have been developed, including models based on series truncations, immersed boundary methods, and fluctuating hydrodynamics.


References

{{DEFAULTSORT:Saffman-Delbruck Model Biophysics Proteins Membrane biology