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Lipids are broadly defined as any fat-soluble (lipophilic), naturally-occurring molecules, such as fats, oils, waxes, cholesterol, steroids, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. Lipids main biological functions include energy storage, acting as structural components of cell membranes, and important signaling molecules. The smooth endoplasmic reticulum also processes these lipids, which store energy.
Although the term lipid is sometimes used as a synonym for fat, fats are in fact a subgroup of lipids called triglycerides and should not be confused with the term fatty acid. The term is also used more specifically to refer to fatty acids and their derivatives (including tri-, di-, and monoglycerides and phospholipids), as well as other fat-soluble steroid-containing metabolites such as cholesterol.
Lipids are a diverse and ubiquitous group of compounds which have many key biological functions, such as acting as structural components of cell membranes, serving as energy storage sources and participating in signaling pathways. Lipids may be broadly defined as hydrophobic or amphipathic small molecules that originate entirely or in part from two distinct types of biochemical subunits or "building blocks": ketoacyl and isoprene groups. Using this approach, lipids may be divided into eight categories : Fatty Acyls, Glycerolipids, Glycerophospholipids, Sphingolipids, Saccharolipids and Polyketides (derived from condensation of ketoacyl subunits); and Sterol Lipids and Prenol Lipids (derived from condensation of isoprene subunits). Although the term lipid is sometimes used as a synonym for fat, fats are generally triesters of glycerol and fatty acids and are covered by a category of lipids called Glycerolipids (which include mono-, di- and triglycerides).
Categories of Lipids
The glycerophospholipids are the main structural component of biological membranes, such as the cellular plasma membrane and the intracellular membranes of organelles. In animal cells the plasma membrane physically separates the intracellular components from the extracellular environment. All eukaryotic cells are compartmentalized into membrane-bound organelles which carry out different functions. These glycerophospholipids are amphipathic molecules that contain a glycerol core linked to two fatty acid-derived "tails" by ester or, more rarely, ether linkages and to one "head" group by a phosphate ester linkage. While glycerophospholipids are the major component of biological membranes, other non-glyceride lipid components such as sphingomyelin and sterols (mainly cholesterol in animal cell membranes) are also found in biological membranes. In plants and algae, the galactosyldiacylglycerols, and sulfoquinovosyldiacylglycerol, which lack a phosphate group, are important components of membranes of chloroplasts and related organelles and are the most abundant lipids in photosynthetic tissues, including those of higher plants, algae and certain bacteria.
A biological membrane is a form of lipid bilayer, as is a liposome. The formation of lipid bilayers is an energetically-preferred process when the glycerophospholipids described above are in an aqueous environment. In an aqueous system, the polar heads of lipids orientate towards the polar, aqueous environment, while the hydrophobic tails minimise their contact with water. The lipophilic tails of lipids (U) tend to cluster together, forming a lipid bilayer (1) or a micelle (2). Other aggregations are also observed and form part of the polymorphism of amphiphile (lipid) behaviour. The polar heads (P) face the aqueous environment, curving away from the water. Phase behaviour is a complicated area within biophysics and is the subject of current academic research. Micelles and bilayers form in the polar medium by a process known as the hydrophobic effect. When dissolving a lipophilic or amphiphilic substance in a polar environment, the polar molecules (i.e. water in an aqueous solution) become more ordered around the dissolved lipophilic substance, since the polar molecules cannot form hydrogen bonds to the lipophilic areas of the amphiphile. So in an aqueous environment the water molecules form an ordered "clathrate" cage around the dissolved lipophilic molecule.
Energy storage and metabolism
Triacylglycerols, stored in adipose tissue, are a major form of energy storage in animals. Animals use triglycerides for energy storage because of its high caloric content (9 KCal/g), whereas plants, which do not require energy for movement, can afford to store food for energy in a less compact but more easily accessible form, such as starch (carbohydrate). Triglycerides and phospholipids are broken down into free fatty acids by the action of lipases. Beta oxidation is the process by which fatty acids, in the form of acyl-CoA molecules, are broken down in the mitochondria and/or in peroxisomes to generate acetyl-CoA. The acetyl CoA is then ultimately converted into ATP, CO2, and H2O using the citric acid cycle and the electron transport chain. Conversely, fatty acid biosynthesis (Lipogenesis) takes place in the cytoplasm, using acetyl-CoA (derived from carbohydrates, amino acids or fatty acids) as the precursor. The fatty acids may be subsequently converted to triacylglycerols that are packaged in lipoproteins (VLDL's) and secreted from the liver.
In recent years, evidence has emerged showing that lipid signaling is a vital part of the cell signaling. Lipid signaling may occur via activation of GPCR's or nuclear receptors, and members of several different lipid categories have been identified as signaling molecules and cellular messengers. These include sphingosine-1-phosphate, a sphingolipid derived from ceramide that is a potent messenger molecule involved in regulating calcium mobilization, cell growth, apoptosis; diacylglycerol(DAG) and the phosphatidylinositol phosphates (PIPs), involved in calcium-mediated activation of protein kinase C; the prostaglandins, arachidonic acid -derived fatty acids involved in inflammation and immunity; the steroid hormones such as estrogen, testosterone and cortisol, which modulate a host of functions such as reproduction, metabolism and blood pressure; and the oxysterols such as 25-hydroxy-cholesterol that are Liver X receptor (LXR) agonists.
The "fat-soluble" vitamins (A, D, E and K) which are isoprene-based lipids are essential nutrients stored in the liver and fatty tissues. These have a diverse range of functions discussed elsewhere. Acyl-carnitines are involved in the transport and metabolism of fatty acids in and out of mitochondria, where they undergo beat oxidation. Polyprenols and their phosphorylated derivatives also play important transport roles, in this case the transport of oligosaccharides across membranes. Polyprenol phosphate sugars and polyprenol diphosphate sugars function in extra-cytoplasmic glycosylation reactions, in extra-cellular polysaccharide biosynthesis (for instance peptidoglycan polymerization in bacteria, and in eukaryotic protein N-glycosylation. Cardiolipins are a subclass of glycerophospholipids containing four acyl chains and three glycerol groups that are particularly abundant in the inner mitochondrial membrane. They are believed to activate enzymes invoved with oxidative phosphorylation.
Nutrition and health
Lipids play diverse and important roles in nutrition and health. Many lipids are absolutely essential for life. However, there is also considerable awareness that abnormal levels of certain lipids, particularly cholesterol (in hypercholesterolemia) and trans fatty acids, are risk factors for heart disease amongst others.
Humans have a requirement for certain essential fatty acids, such as linoleic acid (an omega-6 fatty acid) and alpha-linolenic acid (an omega-3 fatty acid) in the diet because they cannot be synthesized from simple precursors in the diet. Both of these fatty acids are 18-carbon polyunsaturated fatty acids differing in the number and position of the double bonds. Most vegetable oils are rich in linoleic acid (safflower, sunflower, and corn oils). Alpha-linolenic acid is found in the green leaves of plants, and in selected seeds, nuts and legumes (flax, canola, walnuts and soy). Fish oils are particularly rich in the longer-chain omega-6 fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA). Most of the lipid found in food is in the form of triacylglycerols, cholesterol and phospholipids.
Most of the saturated fatty acids (as triacylglycerols) in the diet are incorporated into adipose tissue stores, because the absence of double bonds allows a higher energy yield per carbon than is obtained from oxidation of unsaturated fatty acids. The longer chain fatty acids are incorporated into cell membranes as phospholipids regardless of degree of saturation. Since dietary fatty acids are exchanged with membrane fatty acids, dietary fat composition is reflected in membrane lipid composition. Thus dietary fatty acids can influence cell function through effects on membrane properties. Dietary fat provides an average energy intake which is approximately twice that of carbohydrate or protein. A minimum amount of dietary fat is necessary to facilitate absorption of fat-soluble vitamins (A, D, E and K) and carotenoids. A minimal amount of body fat is also necessary to provide insulation that prevents heat loss and protects vital organs from shock due to ordinary activities.
High fat intake contributes to increased risk of obesity, diabetes and atherosclerosis. Atherosclerosis is the primary cause of coronary and cardiovascular diseases and is primarly due to the buildup of plaque on the inside walls of arteries. Plaque is made up of cholesterol-rich low density lipoproteins (LDL), macrophages, smooth muscle cells, platelets, and other substances. In North America and most other western countries, atherosclerosis is the leading cause of illness and death, almost doubling the number of deaths from cancers. Despite significant medical advances, coronary artery disease and atherosclerotic stroke are responsible for more deaths than all other causes combined.A substantial amount of scientific evidence supports the impact of dietary fatty acids on cardiovascular health. Saturated fats have a profound hypercholesterolemic (increase blood cholesterol levels) effect and tend to increase plasma LDL. They are found predominantly in animal products (butter, cheese and meat) but coconut oil and palm oil are common vegetable sources. Intake of monounsaturated fats in oils such as olive oil is thought to be preferable to consumption of polyunsaturated fats in oils such as corn oil because the monounsaturated fats apparently do not lower high-density-lipoprotein (HDL) cholesterol levels. Keeping cholesterol in the normal range not only helps prevent heart attacks and strokes but may also prevent the progression of atherosclerosis. "Statins" are a class of drugs that lowers the level of cholesterol in the blood by inhibiting the enzyme HMG-CoA reductase. This is a key enzyme involved in the biosynthesis of cholesterol in the liver.
|This article is licensed under the GNU Free Documentation License. It uses material from the Wikipedia article "Lipid". A list of authors is available in Wikipedia.|