Lamellar phase

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Lamellar phase refers generally to packing of polar-headed long chain nonpolar-tail molecules in an environment of bulk polar liquid, as sheets of bilayers separated by bulk liquid. In biophysics, polar lipids (mostly, phospholipids, and rarely, glycolipids) pack as a liquid crystalline bilayer, with hydrophobic fatty acyl long chains directed inwardly and polar headgroups of lipids aligned on the outside in contact with water, as a 2-dimensional flat sheet surface. Under transmission electron microscope (TEM), after staining with polar headgroup reactive chemical osmium tetroxide, lamellar lipid phase appears as two thin parallel dark staining lines/sheets, constituted by aligned polar headgroups of lipids. 'Sandwiched' between these two parallel lines, there exists one thicker line/sheet of non-staining closely packed layer of long lipid fatty acyl chains. This TEM-appearance became famous as Robertson's unit membrane - the basis of all biological membranes, and structure of lipid bilayer in unilamellar liposomes. In multilamellar liposomes, many such lipid bilayer sheets are layered concentrically with water layers in between.

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Figure 1 Multi-lamellar phase of aqueous lipid dispersions, each white lamella represents a lipid bilayer organization in liposome made by vortex-mixing of dried total lipid extract of spinach thylakoid membranes with distilled water. Phosphotungstic acid negative stained sample viewed with transmission electron microscopy technique. Lamellar phase lipids (original research work of Dr. R.C. YashRoy).jpg
Figure 1 Multi-lamellar phase of aqueous lipid dispersions, each white lamella represents a lipid bilayer organization in liposome made by vortex-mixing of dried total lipid extract of spinach thylakoid membranes with distilled water. Phosphotungstic acid negative stained sample viewed with transmission electron microscopy technique.

In lamellar lipid bilayers, polar headgroups of lipids align together at the interface of water and hydrophobic fatty-acid acyl chains align parallel to one another 'hiding away' from water. The lipid head groups are somewhat more 'tightly' packed than relatively 'fluid' hydrocarbon fatty acyl long chains. The lamellar lipid bilayer organization, thus reveals a 'flexibility gradient' of increasing freedom of motions from near the head-groups towards the terminal fatty-acyl chain methyl groups. Existence of such a dynamic organization of lamellar phase in liposomes as well as biological membranes can be confirmed by spin label electron paramagnetic resonance and high resolution nuclear magnetic resonance spectroscopy studies of biological membranes and liposomes. [1]

In 'soft matter science', where physics and chemistry meet biological science, a bilayer lamellar phase has been recently created from fluorinated silica, and it has been projected for use as a shear-thinning lubricant. [2]

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Lipid Substance of biological origin that is soluble in nonpolar solvents

In biology and biochemistry, a lipid is a macro biomolecule that is soluble in nonpolar solvents. Non-polar solvents are typically hydrocarbons used to dissolve other naturally occurring hydrocarbon lipid molecules that do not dissolve in water, including fatty acids, waxes, sterols, fat-soluble vitamins, monoglycerides, diglycerides, triglycerides, and phospholipids.

Phospholipid Class of lipids

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Lipid bilayer Thin polar membrane made of two layers of lipid molecules

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Liposome Composite structures made of phospholipids and may contain small amounts of other molecules

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Glycerophospholipid

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Amphiphile Hydrophilic and lipophilic chemical compound

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Dipalmitoylphosphatidylcholine Chemical compound

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Membrane lipid Lipid molecules on cell membrane

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Lipid polymorphism

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Lyotropic liquid crystal

A liquid crystalline mesophase is called lyotropic if formed by dissolving an amphiphilic mesogen in a suitable solvent, under appropriate conditions of concentration, temperature and pressure. A mixture of soap and water is an everyday example of a lyotropic liquid crystal.

One property of a lipid bilayer is the relative mobility (fluidity) of the individual lipid molecules and how this mobility changes with temperature. This response is known as the phase behavior of the bilayer. Broadly, at a given temperature a lipid bilayer can exist in either a liquid or a solid phase. The solid phase is commonly referred to as a “gel” phase. All lipids have a characteristic temperature at which they undergo a transition (melt) from the gel to liquid phase. In both phases the lipid molecules are constrained to the two dimensional plane of the membrane, but in liquid phase bilayers the molecules diffuse freely within this plane. Thus, in a liquid bilayer a given lipid will rapidly exchange locations with its neighbor millions of times a second and will, through the process of a random walk, migrate over long distances.

Ethanol-induced non-lamellar phases in phospholipids

The presence of ethanol can lead to the formations of non-lamellar phases also known as non-bilayer phases. Ethanol has been recognized as being an excellent solvent in an aqueous solution for inducing non-lamellar phases in phospholipids. The formation of non-lamellar phases in phospholipids is not completely understood, but it is significant that this amphiphilic molecule is capable of doing so. The formation of non-lamellar phases is significant in biomedical studies which include drug delivery, the transport of polar and non-polar ions using solvents capable of penetrating the biomembrane, increasing the elasticity of the biomembrane when it is being disrupted by unwanted substances and functioning as a channel or transporter of biomaterial.

Cell membrane Biological membrane that separates the interior of a cell from its outside environment

The cell membrane is a biological membrane that separates the interior of all cells from the outside environment and protects the cell from its environment. The cell membrane consists of a lipid bilayer, made up of two layers of phospholipids with cholesterols interspersed between them, maintaining appropriate membrane fluidity at various temperatures. The membrane also contains membrane proteins, including integral proteins that span the membrane and serve as membrane transporters, and peripheral proteins that loosely attach to the outer (peripheral) side of the cell membrane, acting as enzymes to facilitate interaction with the cell's environment. Glycolipids embedded in the outer lipid layer serve a similar purpose. The cell membrane controls the movement of substances in and out of cells and organelles, being selectively permeable to ions and organic molecules. In addition, cell membranes are involved in a variety of cellular processes such as cell adhesion, ion conductivity and cell signalling and serve as the attachment surface for several extracellular structures, including the cell wall and the carbohydrate layer called the glycocalyx, as well as the intracellular network of protein fibers called the cytoskeleton. In the field of synthetic biology, cell membranes can be artificially reassembled.

Annular lipids are a set of lipids or lipidic molecules which preferentially bind or stick to the surface of membrane proteins in biological cells. They constitute a layer, or an annulus/ shell, of lipids which are partially immobilized due to the existence of lipid-protein interactions. Polar headgroups of these lipids bind to the hydrophilic part of the membrane protein(s) at the inner and outer surfaces of lipid bilayer membrane. The hydrophobic surface of the membrane proteins is bound to the apposed lipid fatty acid chains of the membrane bilayer. For integral membrane proteins spanning the thickness of the membrane bilayer, these annular/shell lipids may act like a lubricating layer on the proteins' surfaces, thereby facilitating almost free rotation and lateral diffusion of membrane proteins within the 2-dimensional expanse of the biological membrane(s). Outside the layer of shell/annular lipids, lipids are not tied down to protein molecules. However, they may be slightly restricted in their segmental motion freedom due to mild peer pressure of protein molecules, if present in high concentration, which arises from extended influence of protein-lipid interaction. Membrane areas away from protein molecules contain lamellar phase bulk lipids, which are largely free from any restraining effects due to protein-lipid interactions. Thermal denaturation of membrane proteins may destroy the secondary and tertiary structure of membrane proteins, exposing newer surfaces to membrane lipids and therefore increasing the number of lipids molecules in the annulus/shell layer. This phenomenon can be studied by the spin label electron paramagnetic resonance technique. The protein-lipid binding are dependent on OmpF pH levels and their structural features and location of the membranes. When said lipids bind to OmpF it is sensitive to changes that may occur in the electrospray polarity.

A unilamellar liposome is a spherical chamber/vesicle, bounded by a single bilayer of an amphiphilic lipid or a mixture of such lipids, containing aqueous solution inside the chamber. Unilamellar liposomes are used to study biological systems and to mimic cell membranes, and are classified into three groups based on their size: small unilamellar liposomes/vesicles (SUVs) that with a size range of 20–100 nm, large unilamellar liposomes/vesicles (LUVs) with a size range of 100–1000 nm and giant unilamellar liposomes/vesicles (GUVs) with a size range of 1-200 µm. GUVs are mostly used as models for biological membranes in research work. Animal cells are 10–30 µm and plant cells are typically 10–100 µm. Even smaller cell organelles such as mitochondria are typically 1-2 µm. Therefore, a proper model should account for the size of the specimen being studied. In addition, the size of vesicles dictates their membrane curvature which is an important factor in studying fusion proteins. SUVs have a higher membrane curvature and vesicles with high membrane curvature can promote membrane fusion faster than vesicles with lower membrane curvature such as GUVs.

References

  1. YashRoy R C (1990) Magnetic resonance studies of dynamic organisation of lipids in chloroplast membranes. Journal of Biosciences, vol. 15 (No.4), pp. 281-288.
  2. Pottage M J, Kasuma T, Grillo I, Gravey C J, Stickland A D and Tabor R F (2014) Fluorinated lamellar phases: Structural characterization and use as templates for highly ordered silica materials. Soft Matter, vol. 10 (No. 27), pp. 4902-4912.