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Plasma Membrane All living cells, prokaryotic and eukaryotic, have a plasma membrane that encloses their contents

and serves as a semi-porous barrier to the outside environment. The membrane acts as a boundary, holding the cell constituents together and keeping other substances from entering. The plasma membrane is permeable to specific molecules, however, and allows nutrients and other essential elements to enter the cell and waste materials to leave the cell. Small molecules, such as oxygen, carbon dioxide, and water, are able to pass freely across the membrane, but the passage of larger molecules, such as amino acids and sugars, is carefully regulated.

According to the accepted current theory, known as the fluid mosaic model, the plasma membrane is composed of a double layer (bilayer) of lipids, oily substances found in all cells (see Figure 1). Most of the lipids in the bilayer can be more precisely described as phospholipids, that is, lipids that feature a phosphate group at one end of each molecule. Phospholipids are characteristically hydrophilic ("water-loving") at their phosphate ends and hydrophobic ("water-fearing") along their lipid tail regions. In each layer of a plasma membrane, the hydrophobic lipid tails are oriented inwards and the hydrophilic phosphate groups are aligned so they face outwards, either toward the aqueous cytosol of the cell or the outside environment. Phospholipids tend to spontaneously aggregate by this mechanism whenever they are exposed to water. Within the phospholipid bilayer of the plasma membrane, many diverse proteins are embedded, while other proteins simply adhere to the surfaces of the bilayer. Some of these proteins, primarily those that are at least partially exposed on the external side of the membrane, have carbohydrates attached to their outer surfaces and are, therefore, referred to as glycoproteins. The positioning of proteins along the plasma membrane is related in part to the organization of the filaments that comprise the cytoskeleton, which help anchor them in place. The arrangement of proteins also involves the hydrophobic and hydrophilic regions found on the surfaces of the proteins: hydrophobic regions associate with the hydrophobic interior of the plasma membrane and hydrophilic regions extend past the surface of the membrane into either the inside of the cell or the outer environment. Plasma membrane proteins function in several different ways. Many of the proteins play a role in the selective transport of certain substances across the phospholipid bilayer, either acting as channels or active transport molecules. Others function as receptors, which bind information-providing molecules, such as hormones, and transmit corresponding signals based on the obtained information to the interior of the cell. Membrane proteins may also exhibit enzymatic activity, catalyzing various reactions related to the plasma membrane. Since the 1970s, the plasma membrane has been frequently described as a fluid mosaic, which is reflective of the discovery that oftentimes the lipid molecules in the bilayer can move about in the plane of the membrane. However, depending upon a number of factors, including the exact composition of the bilayer and temperature, plasma membranes can undergo phase transitions which render their molecules less dynamic and produce a more gel-like or nearly solid state. Cells are able to regulate the fluidity of their plasma membranes to meet their particular needs by synthesizing more of certain types of molecules, such as those with specific kinds of bonds that keep them fluid at lower temperatures. The presence of cholesterol and glycolipids, which are found in most cell membranes, can also affect molecular dynamics and inhibit phase transitions. In prokaryotes and plants, the plasma membrane is an inner layer of protection since a rigid cell wall forms the outside boundary for their cells. The cell wall has pores that allow materials to enter and leave the cell, but they are not very selective about what passes through. The plasma membrane, which lines the cell wall, provides the final filter between the cell interior and the environment. Eukaryotic animal cells are generally thought to have descended from prokaryotes that lost their cell walls. With only the flexible plasma membrane left to enclose them, these primordial creatures would have been able to expand in size and complexity. Eukaryotic cells are generally ten times larger than prokaryotic cells and have membranes enclosing interior components, the organelles. Like the exterior

plasma membrane, these membranes also regulate the flow of materials, allowing the cell to segregate its chemical functions into discrete internal compartments.

The Fatty Acid Collection Fatty Acids Important constituents of lipids, fatty acid molecules characteristically consist of an even number of carbon atoms linked in a chain bonded with hydrogen atoms that features a carboxyl group at one end. When all of the carbon-to-carbon bonds that hold the chain together are single, the fatty acid is said to be saturated. However, if one or more of the bonds is double, the molecule is unsaturated. As implied by their name, monounsaturated fatty acids only exhibit one double bond, while polyunsaturated acids possess two or more. A few fatty acids that contain triple bonds are also known to exist, although they are far less common than other fatty acids.

Lauric Acid Although lipids and fatty acids are relatively large molecules, they are not biopolymers made up of consecutive units like their cousins DNA, RNA, proteins and polysaccharides. Fatty acids have two chemically different structures combined to allow the unusual properties necessary for these biochemicals to perform their biochemical functions. The generalized structure for fatty acids contains a hydrophilic (water-loving) acidic "head" and a hydrophobic (water-fearing) hydrocarbon "tail". This structural combination is similar to that found in common soaps and fatty acids display a number of soap-like properties. Due to concerns about cholesterol and heart disease, scientists have been very interested in determining which types of fats and oils are best for one's health. All such lipids contain mixtures of fatty acids, but are designated saturated, monounsaturated, or polyunsaturated based upon their predominant component. Fats chiefly consisting of saturated fatty acids are quite stable and are typically found in solid form at room temperature. Studies have shown that these fats may raise blood cholesterol and can contribute to an increased risk of heart disease. Monosaturated fats, which are usually liquid oils at room temperature but may begin to solidify if exposed to colder environments, have often been found to be less unhealthy that saturated varieties. Polyunsaturated fats, however, are an even greater improvement over saturated fats, and substituting these oils, which remain in liquid form even when placed in a refrigerator, for solid saturated varieties may help lower total blood cholesterol.

Oleic Acid Making the issue of "good" versus "bad" fats an even more complex issue, however, is the common process of hydrogenating vegetable and fish oils. In nature, unsaturated fatty acids are typically found in the cis form, in which their hydrogen atoms are located on the same side of their double carbon bonds. Yet, when lipids are hydrogenated, hydrogen atoms can be found on opposite sides of the double bonds they contain, forming what are known as trans fatty acids. In order to extend the shelf life of various food items or to create a solid product, such as margarine, from polyunsaturated oils, which may become rancid rather quickly in their unaltered form, companies often hydrogenate the fatty acids they contain. Though these products were originally believed to be better for the health of consumers than if they contained saturated fats, little evidence remains to support this notion. Most recent studies have suggested that trans fatty acids raise low-density lipoprotein (LDL) and total blood cholesterol levels, while reducing beneficial high-density lipoprotein (HDL) levels.

Docosahexaenoic Acid Fatty acids perform a variety of biochemical functions in the human body, ranging from aiding in the maintenance of the immune system to facilitating the development of healthy cell membranes and enabling the production of prostaglandins, thromboxanes, and other eicosanoids, which are involved in the regulation of vasoconstriction, blood viscosity, blood pressure, and similar activities. However, humans, as well as many other animals, are unable to synthesize all of the fatty acids they need, and some must be obtained from the diet. These essential polyunsaturated fatty acids generally consist of linoleic and alpha-linolenic acids, but arachidonic acid is sometimes also included in the group although it may be synthesized from linolenic acids. Some good dietary sources of linoleic acid, which is part of the Omega-6 family, are green leafy vegetables, nuts, grains, seeds, and the oils made from them. A constituent of the Omega-3 group, alpha-linolenic acid is found in considerable quantities in similar items and is especially

prevalent in flaxseed and fish oils. The human body is able to metabolize fatty acids through the progressive division of pairs of carbon atoms, which are then converted into acetyl coenzyme A. This coenzyme is oxidized as part of the citric acid cycle that is also involved in the breakdown of the sugar glucose. Through this process, fatty acids yield relatively large amounts of adenosine triphosphate ( ATP), making the molecules an excellent source of energy, a fact that is supported by the tendency of the human body to store excess fuel as fat. Nevertheless, some parts of the body, such as the brain, are unable to utilize fatty acids as an energy source and must instead depend upon glucose metabolism to support their activity.

Linoleic Acid The Molecular Expressions collection of fatty acids contains only a few members of this important class of biochemicals. Fatty acids are very difficult to crystallize and this makes it difficult to obtain high-quality photomicrographs. Also, they have been hard for us to obtain, and we are looking for new fatty acid and lipid samples. Should you be able to provide us with any samples in this arena please contact us by phone or e-mail.

Cell Membranes Cell membranes protect and organize cells. All cells have an outer plasma membrane that regulates not only what enters the cell, but also how much of any given substance comes in. Unlike prokaryotes, eukaryotic cells also possess internal membranes that encase their organelles and control the exchange of essential cell components. Both types of membranes have a specialized structure that facilitates their gatekeeping function. What Are Cellular Membranes Made Of? With few exceptions, cellular membranes including plasma membranes and internal membranes are made of glycerophospholipids, molecules composed of glycerol, a phosphate group, and two fatty acid chains. Glycerol is a three-carbon molecule that functions as the backbone of these membrane lipids. Within an individual glycerophospholipid, fatty acids are attached to the first and second carbons, and the phosphate group is attached to the third carbon of the glycerol backbone. Variable head groups are attached to the phosphate. Space-filling models of these molecules reveal their cylindrical shape, a geometry that allows glycerophospholipids to align side-by-side to form broad sheets (Figure 1).
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Figure 1: The lipid bilayer and the structure and composition of a glycerophospholipid molecule (A) The plasma membrane of a cell is a bilayer of glycerophospholipid molecules. (B) A single glycerophospholipid molecule is composed of two major regions: a hydrophilic head (green) and hydrophobic tails (purple). (C) The subregions of a glycerophospholipid molecule; phosphatidylcholine is shown as an example. The hydrophilic head is composed of a choline structure (blue) and a phosphate (orange). This head is connected to a glycerol (green) with two hydrophobic tails (purple) called fatty acids. (D) This view shows the specific atoms within the various subregions of the phosphatidylcholine molecule. Note that a double bond between two of the carbon atoms in one of the hydrocarbon (fatty acid) tails causes a slight kink on this molecule, so it appears bent. 2010 Nature Education All rights reserved. Glycerophospholipids are by far the most abundant lipids in cell membranes. Like all lipids, they are insoluble in water, but their unique geometry causes them to aggregate into bilayers without any energy input. This is because they are two-faced molecules, with hydrophilic (water-loving) phosphate heads and hydrophobic (water-fearing) hydrocarbon tails of fatty acids. In water, these molecules spontaneously align with their heads facing outward and their tails lining up in the bilayer's interior. Thus, the hydrophilic heads of the glycerophospholipids in a cell's plasma membrane face both the water-based cytoplasm and the exterior of the cell. Altogether, lipids account for about half the mass of cell membranes. Cholesterol molecules, although less abundant than glycerophospholipids, account for about 20 percent of the lipids in animal cell plasma membranes. However, cholesterol is not present in bacterial membranes or mitochondrial membranes. Also, cholesterol helps regulate the stiffness of membranes, while other less prominent lipids play roles in cell signaling and cell recognition.
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Figure 2: The glycerophospholipid bilayer with embedded transmembrane proteins

2010 Nature Education All rights reserved. In addition to lipids, membranes are loaded with proteins. In fact, proteins account for roughly half the mass of most cellular membranes. Many of these proteins are embedded into the membrane and stick out on both sides; these are called transmembrane proteins. The portions of these proteins that are nested amid the hydrocarbon tails have hydrophobic surface characteristics, and the parts that stick out are hydrophilic (Figure 2). At physiological temperatures, cell membranes are fluid; at cooler temperatures, they become gel-like. Scientists who model membrane structure and dynamics describe the membrane as a fluid mosaic in which transmembrane proteins can move laterally in the lipid bilayer. Therefore, the collection of lipids and proteins that make up a cellular membrane relies on natural biophysical properties to form and function. In living cells, however, many proteins are not free to move. They are often anchored in place within the membrane by tethers to proteins outside the cell, cytoskeletal elements inside the cell, or both. What Do Membranes Do? Cell membranes serve as barriers and gatekeepers. They are semi-permeable, which means that some molecules can diffuse across the lipid bilayer but others cannot. Small hydrophobic molecules and gases like oxygen and carbon dioxide cross membranes rapidly. Small polar molecules, such as water and ethanol, can also pass through membranes, but they do so more slowly. On the other hand, cell membranes restrict diffusion of highly charged molecules, such as ions, and large molecules, such as sugars and amino acids. The passage of these molecules relies on specific transport proteins embedded in the membrane.
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Figure 3: Selective transport Specialized proteins in the cell membrane regulate the concentration of specific molecules inside the cell. 2010 Nature Education All rights reserved. Membrane transport proteins are specific and selective for the molecules they move, and they often use energy to catalyze passage. Also, these proteins transport some nutrients against the concentration gradient, which requires additional energy. The ability to maintain concentration gradients and sometimes move materials against them is vital to cell health and maintenance. Thanks to membrane barriers and transport proteins, the cell can accumulate nutrients in higher concentrations than exist in the environment and, conversely, dispose of waste products (Figure 3). Other transmembrane proteins have communication-related jobs. These proteins bind signals, such as hormones or immune mediators, to their extracellular portions. Binding causes a conformational change in the protein that transmits a signal to intracellular messenger molecules. Like transport proteins, receptor proteins are specific and selective for the molecules they bind (Figure 4).
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Figure 4: Examples of the action of transmembrane proteins Transporters carry a molecule (such as glucose) from one side of the plasma membrane to the other. Receptors can bind an extracellular molecule (triangle), and this activates an intracellular process. Enzymes in the membrane can do the same thing they do in the cytoplasm of a cell: transform a molecule into another form. Anchor proteins can physically link intracellular structures with extracellular structures. 2010 Nature Education All rights reserved. Peripheral membrane proteins are associated with the membrane but are not inserted into the bilayer. Rather, they are usually bound to other proteins in the membrane. Some peripheral proteins form a filamentous network just under the membrane that provides attachment sites for transmembrane proteins. Other peripheral proteins are secreted by the cell and form an extracellular matrix that functions in cell recognition. How Diverse Are Cell Membranes? In contrast to prokaryotes, eukaryotic cells have not only a plasma membrane that encases the entire cell, but also intracellular membranes that surround various organelles. In such cells, the plasma membrane is part of an extensive endomembrane system that includes the endoplasmic reticulum (ER), the nuclear membrane, the Golgi apparatus, and lysosomes. Membrane components are exchanged throughout the endomembrane system in an organized fashion. For instance, the membranes of the ER and the Golgi apparatus have different compositions, and the proteins that are found in these membranes contain sorting signals, which are like molecular zip codes that specify their final destination. Mitochondria and chloroplasts are also surrounded by membranes, but they have unusual membrane structures specifically, each of these organelles has two surrounding membranes instead of just one. The outer membrane of mitochondria and chloroplasts has pores that allow small molecules to pass easily. The inner membrane is loaded with the proteins that make up the electron transport chain and help generate energy for the cell. The double membrane enclosures of mitochondria and chloroplasts are similar to certain modern-day prokaryotes and are thought to reflect these organelles' evolutionary origins. Conclusion Membranes are made of lipids and proteins, and they serve a variety of barrier functions for cells and intracellular organelles. Membranes keep the outside "out" and the inside "in," allowing only certain molecules to cross and relaying messages via a chain of molecular events