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214       Organelles



             Structure and components                         B. Membrane lipids
                                                              The illustration shows amodel of asmall
             A. Structure of the plasma membrane              section of a membrane. The phospholipids
                                                              are the most important group of membrane
             Allbiologicalmembranes are constructed ac-
             cording to a standard pattern. They consist of   lipids.  They   include   phosphatidylcholine
                                                              (lecithin), phosphatidylethanolamine, phos-
             a continuous bilayer of amphipathic lipids ap-   phatidylserine,   phosphatidylinositol,  and
             proximately 5 nm thick, into which proteins      sphingomyelin   (for  their  structures,  see
             are embedded. In addition, some membranes        p. 50). In addition, membranes in animal cells
             also carry carbohydrates (mono- and oligo-
             saccharides) on their exterior, which are        also contain cholesterol (with the exception
             bound to lipids and proteins. The proportions    of inner mitochondrial membranes). Glycoli-
                                                              pids (a ganglioside is shown here) are mainly
             of lipids, proteins, and carbohydrates differ    found on the outside of the plasma mem-
             markedly depending on the type of cell and       brane. Together with the glycoproteins, they
             membrane (see p. 216).
                Membrane lipids are strongly amphipathic      form the exterior coating of the cell (the gly-
             molecules with a polar hydrophilic “head         cocalyx).
             group” and an apolar hydrophobic “tail.” In
             membranes, they are primarily held together      C. Membrane proteins
             by the hydrophobic effect (see p. 28) and
             weak Van der Waals forces, and are therefore     Proteins can be anchored in or on membranes
             mobile relative to each other. This gives mem-   in various ways. Integral membrane proteins
                                                              cross right through the lipid bilayer. The sec-
             branes a more or less fluid quality.
                The fluidity of membranes primarily de-       tions of the peptide chains that lie within the
             pends on their lipid composition and on tem-     bilayer usually consist of 20 to 25 mainly
             perature. At a specific transition temperature,  hydrophobic amino acid residues that form a
                                                              right-handed α-helix.
             membranes pass from a semicrystalline state
             to a more fluid state. The double bonds in the      Type I and II membrane proteins only
             alkyl chains of unsaturated acyl residues in     contain one transmembrane helix of this
                                                              type, whiletypeIII proteins contain several.
             the membrane lipids disturb the semicrystal-
             line state. The higher the proportion of unsa-   Rarely, type I and II polypeptides can aggre-
                                                              gate to form a type IV transmembrane pro-
             turated lipids present, therefore, the lower the  tein. Several groups of integral membrane
             transition temperature. The cholesterol con-
             tent also influences membrane fluidity. While    proteins—e. g., theporins (seep.212)—pene-
                                                              trate the membrane with antiparallel β-sheet
             cholesterol increases the fluidity of semicrys-
             talline, closely-packed membranes, it stabil-    structures. Due to its shape, this tertiary
                                                              structure is known as a “β-barrel.”
             izes fluid membranes that contain a high pro-
             portion of unsaturated lipids.                      Type V and VI proteins carry lipid anchors.
                Like lipids, proteins are also mobile within  These are fatty acids (palmitic acid, myristic
             the membrane. If they are not fixed in place     acid), isoprenoids (e. g., farnesol), or glycoli-
             by special mechanisms, they float within the     pids such as glycosyl phosphatidylinositol
                                                              (GPI) that are covalently bound to the peptide
             lipid layer as if in a two-dimensional liquid;
             biological membranes are therefore also de-      chain.
                                                                 Peripheral membrane proteins are associ-
             scribedasbeing a “fluidmosaic.”                  ated with the head groups of phospholipids
                Lipids and proteins can shift easily within
             one layer of a membrane, but switching be-       or with another integral membrane protein
                                                              (not shown).
             tween the two layers (“flip/flop”)is not possi-
             ble for proteins and is only possible with dif-
             ficulty for lipids (with the exception of cho-
             lesterol). To move to the other side, phospho-
             lipids require special auxiliary proteins
             (translocators, “flipases”).






           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
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