Cell membrane

Editor-In-Chief: C. Michael Gibson, M.S., M.D. [1]

The cell membrane (also called the plasma membrane, plasmalemma or "phospholipid bilayer") is a semipermeable lipid bilayer found in all cells.^[1] It contains a wide variety of biological molecules, primarily proteins and lipids, which are involved in a vast array of cellular processes, and also serves as the attachment point for both the intracellular cytoskeleton and, if present, the cell wall.

Function

The cell membrane surrounds the cytoplasm of a cell and, in animal cells, physically separates the intracellular components from the extracellular environment, thereby serving a function similar to that of skin. In fungi, bacteria, and plants an additional cell wall forms the outermost boundary, however, the cell wall plays mostly a mechanical support role rather than a role as a selective boundary. The cell membrane also plays a role in anchoring the cytoskeleton to provide shape to the cell, and in attaching to the extracellular matrix to help group cells together in the formation of tissues.

The barrier is selectively permeable and able to regulate what enters and exits the cell, thus facilitating the transport of materials needed for survival. The movement of substances across the membrane can be either passive, occurring without the input of cellular energy, or active, requiring the cell to expend energy in moving it. The membrane also maintains the cell potential.

Specific proteins embedded in the cell membrane can act as molecular signals which allow cells to communicate with each other. Protein receptors are found ubiquitously and function to receive signals from both the environment and other cells. These signals are transduced into a form which the cell can use to directly effect a response. Other proteins on the surface of the cell membrane serve as "markers" which identify a cell to other cells. The interaction of these markers with their respective receptors forms the basis of cell-cell interaction in the immune system.

Structure

Lipid bilayer

The cell membrane consists of a thin layer of amphipathic lipids which spontaneously arrange so that the hydrophobic "tail" regions are shielded from the surrounding polar fluid, causing the more hydrophilic "head" regions to associate with the cytosolic and extracellular faces of the resulting bilayer. This forms a continuous, spherical lipid bilayer containing the cellular components approximately 7 nm thick, barely discernible with a transmission electron microscope.^[1]

The arrangement of hydrophilic and hydrophobic heads of the lipid bilayer prevents hydrophilic solutes from passively diffusing across the band of hydrophobic tail groups, allowing the cell to control the movement of these substances via transmembrane protein complexes such as pores and gates.

Flippases and Scramblases concentrate phosphatidyl serine, which carries a negative charge, on the inner membrane. Along with NANA, this creates an extra barrier to charged Moities moving through the membrane.

Membranes serve diverse functions in eukaryotic and prokaryotic cells. One important role is to regulate the movement of materials into and out of cells. The phospholipid bilayer structure (fluid mosaic model) with specific membrane proteins accounts for the selective permeability of the membrane and passive and active transport mechanisms. In addition, membranes in prokaryotes and in the mitochondria and chloroplasts of eukaryotes facilitate the synthesis of ATP through chemiosmosis.

Integral membrane proteins

The cell membrane contains many integral membrane proteins which pepper the entire surface. These structures, which can be visualized by electron microscopy or fluorescence microscopy, can be found on the inside of the membrane, the outside, or through-and-through. They include synapses, desmosomes, clathrin-coated pits, caveolaes, and different structures involved in cell adhesion.

Membrane skeleton

The cytoskeleton is found underlying the cell membrane in the cytoplasm and provides a scaffolding for membrane proteins to anchor to, as well as forming organelles which extend from the cell. Anchoring proteins restricts them to a particular cell surface — for example, the apical surface of epithelial cells that line the vertebrate gut — and limits how far they may diffuse within the bilayer. The cytoskeleton is able to form appendage-like organelles, such as cilia, which are covered by the cell membrane and project from the surface of the cell. The apical surfaces of the aforementioned epithelial cells are dense with finger-like projections, called microvilli, which increase cell surface area and thereby increase the absorption rate of nutrients. The cell membrane acts as a protecting body.

Structure and the Fluid mosaic model

According to the fluid mosaic model of S. Jonathan Singer and Garth Nicolson, the biological membranes can be considered as a two-dimensional liquid where all lipid and protein molecules diffuse more or less freely^[2]. This picture may be valid in the space scale of 10 nm. However, the plasma membranes contain different structures or domains that can be classified as (a) protein-protein complexes; (b) lipid rafts, (c) pickets and fences formed by the actin-based cytoskeleton; and (d) large stable structures, such as synapses or desmosomes.

The fluid mosaic model can be seen when the membrane proteins of two cells (e.g., a human cell and a mouse cell) are tagged with different-coloured fluorescent labels. When the two cells are fused, the two colours intermix, indicating that the proteins are free to move in the 2D plane.

Composition

Cell membranes contain a variety of biological molecules, notable lipids and proteins. Material is incorporated into the membrane, or deleted from it, by a variety of mechanisms:

Fusion of intracellular vesicles with the membrane not only excretes the contents of the vesicle, but also incorporates the vesicle membrane's components into the cell membrane. The membrane may form blebs that pinch off to become vesicles.
If a membrane is continuous with a tubular structure made of membrane material, then material from the tube can be drawn into the membrane continuously.
Although the concentration of membrane components in the aqueous phase is low (stable membrane components have low solubility in water), exchange of molecules with this small reservoir is possible.

In all cases, the mechanical tension in the membrane has an effect on the rate of exchange. In some cells, usually having a smooth shape, the membrane tension and area are interrelated by elastic and dynamical mechanical properties, and the time-dependent interrelation is sometimes called homeostasis, area regulation or tension regulation.

Lipids

The cell membrane consists of three classes of amphipathic lipids: phospholipids, glycolipids, and steroids. The relative composition of each depends upon the type of cell, but in the majority of cases phospholipids are the most abundant.^[3] In RBC studies, 30% of the plasma membrane is lipid.

The fatty chains in phospholipids and glycolipids usually contain an even number of carbon atoms, typically between 14 and 24. The 16- and 18-carbon fatty acids are the most common. Fatty acids may be saturated or unsaturated, with the configuration of the double bonds nearly always cis. The length and the degree of unsaturation of fatty acids chains have a profound effect on membranes fluidity^[4].

The entire membrane is held together via non-covalent interaction of hydrophobic tails, however the structure is quite fluid and not fixed rigidly in place. Phospholipid molecules in the cell membrane are "fluid" in the sense that they are free to diffuse and exhibit rapid lateral diffusion along the layer they are present in. However, movement of phospholipid molecules between layers is not energetically favourable and does not occur to an appreciable extent. Lipid rafts and caveolae are examples of cholesterol-enriched microdomains in the cell membrane.

In animal cells cholesterol is normally found dispersed in varying degrees throughout cell membranes, in the irregular spaces between the hydrophobic tails of the membrane lipids, where it confers a stiffening and strengthening effect on the membrane.^[1]

Carbohydrates

About 5% of the plasma membrane weight is carbohydrate, predominantly glycoprotein, but with some lipoprotein (cerebrosides and gangliosides). Mostly, no glycosylation occurs on other unit membranes, one way in which the plasma membrane differs from others.

The glycocalyx is an important feature in all cells, especially epithelia with microvilli. Recent data suggest the glycocalyx participates in cell adhesion, lymphocyte homing, and many others.

The penultimate sugar is galactose and the terminal sugar is sialic acid as the sugar backbone is modified in the golgi apparatus. Sialic acid carries a negative charge, providing an external barrier to charged particles.

Proteins

Type	Description	Examples
Integral proteins or transmembrane proteins	Span the membrane and have a hydrophilic cytosolic domain which interacts with internal molecules, a hydrophobic membrane-spanning domain which anchors it within the cell membrane, and a hydrophilic extracellular domain which interacts with external molecules. The hydrophobic domain consists of one, multiple, or a combination of α-helices and β sheet protein motifs.	Ion channels, proton pumps, G protein-coupled receptor
Lipid anchored proteins	Covalently bound to single or multiple lipid molecules; hydrophobically insert into the cell membrane and anchor the protein. The protein itself is not in contact with the membrane.	G proteins
Peripheral proteins	Attached to integral membrane proteins, or associated with peripheral regions of the lipid bilayer. These proteins tend to have only temporary interactions with biological membranes, and once reacted the molecule dissociates to carry on its work in the cytoplasm.	Some enzymes, some hormones

The cell membrane plays host to a large amount of protein which is responsible for its various activities. The amount of protein differs between species and according to function, however the typical amount in a cell membrane is 50%.^[4] These proteins are undoubtedly important to a cell: approximately a third of the genes in yeast code specifically for them, and this number is even higher in multicellular organisms.^[3]

The cell membrane, being exposed to the outside environment, is an important site of cell-cell communication. As such, a large variety of protein receptors and identification proteins, such as antigens, are present on the surface of the membrane.

References

↑ ^1.0 ^1.1 ^1.2 Alberts B, Johnson A, Lewis J; et al. Molecular Biology of the Cell (4th ed. ed.). ISBN 0-8153-3218-1.
↑ The fluid mosaic model of the structure of cell membranes by S. J. Singer and G. L. Nicolson in Science (1972) Volume 175, pages 720-731.
↑ ^3.0 ^3.1 Lodish H, Berk A, Zipursky LS; et al. (2004). Molecular Cell Biology (4th ed. ed.). ISBN 0-7167-3136-31986 Check |isbn= value: length (help).
↑ ^4.0 ^4.1 Jesse Gray, Shana Groeschler, Tony Le, Zara Gonzalez (2002). "Membrane Structure" (SWF). Davidson College. Retrieved 2007-01-11.

External links

Template:Organelles Template:Structures of the cell membrane Template:WH Template:WS Template:Jb1

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[MBOC-1] 1.0 ^1.1 ^1.2 Alberts B, Johnson A, Lewis J; et al. Molecular Biology of the Cell (4th ed. ed.). ISBN 0-8153-3218-1.

[2] The fluid mosaic model of the structure of cell membranes by S. J. Singer and G. L. Nicolson in Science (1972) Volume 175, pages 720-731.

[Lodish-3] 3.0 ^3.1 Lodish H, Berk A, Zipursky LS; et al. (2004). Molecular Cell Biology (4th ed. ed.). ISBN 0-7167-3136-31986 Check |isbn= value: length (help).

[flashbio-4] 4.0 ^4.1 Jesse Gray, Shana Groeschler, Tony Le, Zara Gonzalez (2002). "Membrane Structure" (SWF). Davidson College. Retrieved 2007-01-11.

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