Membranes translational motions

Hackenbrock CR, Hochh M, Chan RM. Calorimetric and freeze fractnre analysis of lipid phase transitions and lateral translational motion of intramembrane particles in mitochondrial membranes. Biochim. Biophys. Acta 1976 455 466 84. 

Because diffusion is simply rapid, random, translational motion of atoms and molecules, hydrogen in superpermeable membranes will travel just as quickly from the permeate side of the membranes to the retentate side, as it does in diffusing from the retentate to the permeate side. From the fundamental laws of diffusion, diffusion will tend to eliminate concentration gradients across a membrane, and at equiUbrium the net flux across a membrane will be zero [14]. For superpermeable membranes, equilibrium will be very quickly approached, unless hydrogen is removed extremely rapidly from the permeate side of the membrane. 

The currently accepted structure of B. is the fluid mosaic model. Lipid molecules and membrane proteins are free to diffuse laterally and to spin within the bilayer in which they are located. However, a flip-flop motion from the inner to the outer surface, or vice versa, is energetically unfavorable, because it would require movement of hydrophilic substituents through the hydrophobic phase. Hence this type of motion is almost never displayed by proteins, and it occurs much less readily than translational motion in the case of lipids. Since there is little movement of material between the inner and outer layers of the bilayer, the two faces of the B. can have different compositions. For membrane proteins, this asymmetry is absolute, and, at least in the plasma membrane, different proportions of lipid classes exist in the two monolayers. Attached carbohydrate residues appear to be located only on the noncytosolic surface. Carbohydrate groups extending from the B. participate in cell recognition, cell adhesion, possibly in intercellular communication, and they also contribute to the distinct immunological character of the cell. 

The membrane-bound bacteriorhodopsin system also provides an example of the illustration of electrochemical aspects of enzymic machineries which were not discussed in the context of this symposium. Membrane-bound enzyme systems are of special interest because they are oriented yielding a vectorial property which was stressed by Peter Mitchell already in the early sixties. The structural mobility of intramembrane molecules is restricted due to the lipid protein interaction. There might be controlled translational motion or rotational motion or in cases such as the purple membrane no gross motion at all. 

Diffusion, Flow and Permeation. Diffusion-weighted NMR imaging (DWI) is sensitive to the random translational motion of water molecules due to Brownian motion. Although the mechanism is still not completely understood, the cellular swelling that accompanies cell membrane depolarization results in a reduction in the net displacement of diffusing water molecules and... 

Under normal conditions the membrane bilayer is in a fluid state. Membrane proteins can migrate within the plane of the membrane with diffusion coefficients of about 10 cm sec while lipids diffuse with coefficients of about 10 cm sec . Overall behaviour might be considered, therefore, in thermodynamic terms. But generalized deductions relating fluidity of the membrane to enzyme activity are difficult to make for several reasons. For example, motion in a given lipid molecule may include rapid rotations but slow lateral movement. Also, increased disorder in a bilayer may not correlate with increased translational motion. Moreover, all membranes so far examined have shown transbilayer asymmetry while there is evidence in several cases for at least small areas of concentration of certain lipids, i.e. micro-lateral heterogeneity. These sorts of consideration complicate the interpretation of experiments designed to show how the bilayer lipids affect membrane enzyme activities at a molecular level. 

The most likely way for pardaxin molecules to insert across the membrane in an antiparallel manner is for them to form antiparallel aggregates on the membrane surface that then insert across the membrane. We developed a "raft"model (data not shown) that is similar to the channel model except that adjacent dimers are related to each other by a linear translation instead of a 60 rotation about a channel axis. All of the large hydrophobic side chains of the C-helices are on one side of the "raft" and all hydrophilic side chains are on the other side. We postulate that these "rafts" displace the lipid molecules on one side of the bilayer. When two or more "rafts" meet they can insert across the membrane to form a channel in a way that never exposes the hydrophilic side chains to the lipid alkyl chains. The conformational change from the "raft" to the channel structure primarily involves a pivoting motion about the "ridge" of side chains formed by Thr-17, Ala-21, Ala-25, and Ser-29. These small side chains present few steric barriers for the postulated conformational change. 

Studies of the effect of permeant s size on the translational diffusion in membranes suggest that a free-volume model is appropriate for the description of diffusion processes in the bilayers [93]. The dynamic motion of the chains of the membrane lipids and proteins may result in the formation of transient pockets of free volume or cavities into which a permeant molecule can enter. Diffusion occurs when a permeant jumps from a donor to an acceptor cavity. Results from recent molecular dynamics simulations suggest that the free volume transport mechanism is more likely to be operative in the core of the bilayer [84]. In the more ordered region of the bilayer, a kink shift diffusion mechanism is more likely to occur [84,94]. Kinks may be pictured as dynamic structural defects representing small, mobile free volumes in the hydrocarbon phase of the membrane, i.e., conformational kink g tg ) isomers of the hydrocarbon chains resulting from thermal motion [52] (Fig. 8). Small molecules can enter the small free volumes of the kinks and migrate across the membrane together with the kinks. 

We would like to point out that an order parameter indicates the static property of the lipid bilayer, whereas the rotational motion, the oxygen transport parameter (Section 4.1), and the chain bending (Section 4.4) characterize membrane dynamics (membrane fluidity) that report on rotational diffusion of alkyl chains, translational diffusion of oxygen molecules, and frequency of alkyl chain bending, respectively. The EPR spin-labeling approach also makes it possible to monitor another bulk property of lipid bilayer membranes, namely local membrane hydrophobicity. 

Pulsed field gradient (PFG)-NMR experiments have been employed in the groups of Zawodzinski and Kreuer to measure the self-diffusivity of water in the membrane as a function of the water content. From QENS, the typical time and length scales of the molecular motions can be evaluated. It was observed that water mobility increases with water content up to almost bulk-like values above T 10, where the water content A = nn o/ nsojH is defined as the ratio of the number of moles of water molecules per moles of acid head groups (-SO3H). In Perrin et al., QENS data for hydrated Nation were analyzed with a Gaussian model for localized translational diffusion. Typical sizes of confining domains and diffusion coefficients, as well as characteristic times for the elementary jump processes, were obtained as functions of A the results were discussed with respect to membrane structure and sorption characteristics. ... 

Fig. 61. Schematics of pressure-induced and applied-potential-induced BLM deformations. Application of hydrostatic pressure (by lowering a piston into the aqueous solution bathing the cis side of the BLM) displaces the BLM from position 1 to position 2. The displacement involves both translational (lateral) motion (Ft) and curvature increase (Fc). As indicated, deformation of the BLM is accompanied by a change in its torus (Plateau-Gibbs border). 2R and 2Rm represent the diameters of the aperture of the pinhole in the Tefzel film and that of the membrane (excluding the torus). The object laser beam, incident upon the trans side of the BLM and reflected by it at 45° at a shortened wavelength produces concentric optical interference fringes with the reference laser beam. Ag/AgCl electrodes, placed in the cis and trans sides of the BLM, allow for continuous electrical measurements [413]...

In the popular fluid mosaic model for biomembranes, membrane proteins and other membrane-embedded molecules are in a two-dimensional fluid formed by the phospholipids. Such a fluid state allows free motion of constituents within the membrane bilayer and is extremely important for membrane function. The term "membrane fluidity" is a general concept, which refers to the ease of motion for molecules in the highly anisotropic membrane environment. We give a brief description of physical parameters associated with membrane fluidity, such as rotational and translational diffusion rates, order parameters etc., and review physical methods used for their determination. We also show limitations of the fluid mosaic model and discuss recent developments in membrane science that pertain to fluidity, such as evidence for compartmentalization of the biomembrane by the cell cytoskeleton. 

Diffusion is the random movement of a particle because of an exchange of thermal energy with its environment. Membrane lipids and proteins participate in highly anisotropic translational and rotational diffusion motion. Translational diffusion in the plane of the membrane is described by the mean square lateral displacement after a time At (r ) = TD At. Lipid lateral diffusion coefficients in fluid phase bilayers are typically in the range Dj 10 to 10 cm /s (3). 

Although biologic membranes serve as physical barriers, it is important to recognize that their molecular constituents are in a constant state of motional flux. Many different types of molecular motion are present in biological membranes including rotation, translation and libration, each of which contributes in important ways to the physical properties of cellular membranes. Since alterations in membrane physical properties have profound effects on the kinetics of many transmembrane enzymes and modulate the rates and types of interactions between proteins, it comes as no surprise that the molecular dynamics of a cell membrane is an important modulator of signal transduction (e.g., Lenaz, 1987). Thus, biological... 

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