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46        Biomolecules



             Overview                                         are derivatives of the polyunsaturated fatty
                                                              acid arachidonic acid (see p. 390).

             A. Classification
                                                              B. Biological roles
             The lipids are a large and heterogeneous
             group ofsubstances ofbiologicaloriginthat        1. Fuel. Lipids are an important source of en-
             are easily dissolved in organic solvents such    ergy in the diet. In quantitative terms, they
             as methanol, acetone, chloroform, and ben-       represent the principal energy reserve in ani-
             zene. By contrast, they are either insoluble or  mals. Neutral fats in particular are stored in
             only poorly soluble in water. Their low water    specialized cells, known as adipocytes.Fatty
             solubility is due to a lack of polarizing atoms  acids are released from these again as needed,
             such as O, N, S, and P (see p. 6).               and these are then oxidized in the mitochon-
                Lipids can be classified into substances that  dria to form water and carbon dioxide, with
             are either hydrolyzable—i. e., able to undergo   oxygen being consumed. This process also
             hydrolytic cleavage—or nonhydrolyzable.Only      gives rise to reduced coenzymes, which are
             a few examples of the many lipids known can      used for ATP production in the respiratory
             be mentioned here. The individual classes of     chain (see p.140).
             lipids are discussed in more detail in the fol-     2. Nutrients. Amphipathic lipids are used
             lowing pages.                                    by cells to build membranes (see p. 214). Typ-
                Hydrolyzable lipids (components shown in      ical membrane lipids include phospholipids,
             brackets). The simple esters include the fats    glycolipids, and cholesterol. Fats are only
             (triacylglycerol; one glycerol + three acyl res-  weakly amphiphilic and are therefore not
             idues); the waxes (one fatty alcohol + one acyl  suitableas membranecomponents.
             residue); and the sterol esters (one sterol + one   3. Insulation. Lipids are excellent insula-
             acyl residue). The phospholipids are esters      tors. In the higher animals, neutral fats are
             with more complex structures. Their charac-      found in the subcutaneous tissue and around
             teristic component is a phosphate residue.       various organs, where they serve as mechan-
             The phospholipids include the phosphatidic       ical and thermal insulators. As the principal
             acids (one glycerol + two acyl residues + one    constituent of cell membranes, lipids also in-
             phosphate) and the phosphatides (one glyc-       sulate cells from their environment mechan-
             erol + two acyl residues + one phosphate +       ically and electrically. The impermeability of
             one amino alcohol). In the sphingolipids,glyc-   lipid membranes to ions allows the formation
             eroland oneacylresidue arereplaced by            of themembranepotential (see p.126).
             sphingosine. Particularly important in this         4. Special tasks. Some lipids have adopted
             group are the sugar-containing glycolipids       special roles in the body. Steroids, eicosa-
             (one sphingosine + one fatty acid + sugar).      noids, and some metabolites of phospholipids
             The cerebrosides (one sphingosine + one fatty    have signaling functions. They serve as hor-
             acid + one sugar) and gangliosides (one sphin-   mones, mediators, and second messengers
             gosine + one fatty acid + several different      (see p. 370). Other lipids form anchors to at-
             sugars, including neuraminic acid) are repre-    tach proteins to membranes (see p. 214). The
             sentatives of this group.                        lipids also produce cofactors for enzymatic re-
                The components of the hydrolyzable lipids     actions—e. g., vitamin K (see p. 52) and ubiq-
             are linked to one another by ester bonds.They    uinone (see p.104). The carotenoid retinal, a
             are easily broken down either enzymatically      light-sensitive lipid, is of central importance
             or chemically.                                   in the process of vision (see p. 358).
                Non-hydrolyzable lipids. The hydrocarbons        Several lipids are not formed indepen-
             include the alkanes and carotenoids.The lipid    dently in the human body. These substances,
             alcohols are also not hydrolyzable. They in-     as essential fatty acids and fat-soluble vita-
             clude long-chained alkanols and cyclic sterols   mins, are indispensable components of nutri-
             such as cholesterol, and steroids such as es-    tion (see pp. 364ff.)
             tradiol and testosterone. The most important
             acids among the lipids are fatty acids. The
             eicosanoids also belong to this group; these




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