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       plasmic reticulum (RER). These ribosomes syn-  Hence, the Golgi apparatus represents a
       thesize export proteins as well as transmem-  central modification, sorting and distribution
       brane proteins (! G) for the plasma mem-  center for proteins and lipids received from the
       brane, endoplasmic reticulum, Golgi appara-  endoplasmic reticulum.
       tus, lysosomes, etc. The start of protein synthe-  Regulation of gene expression takes place
       sis (at the amino end) by such ribosomes (still  on the level of transcription (! C1a), RNA
       unattached) induces a signal sequence to  modification (! C1b), mRNA export (! C1c),
       which a signal recognition particle (SRP) in the  RNA degradation (! C1d), translation (! C1e),
       cytosol attaches. As a result, (a) synthesis is
                                       modification and sorting (! F,f), and protein
    Fundamentals and Cell Physiology  tein synthesis, a translocator protein conveys  site of oxidation of carbohydrates and lipids to
                                       degradation (! F,g).
       temporarily halted and (b) the ribosome (me-
                                        The mitochondria (! A, B; p. 17 B) are the
       diated by the SRP and a SRP receptor) attaches
       to a ribosome receptor on the ER membrane.
                                       CO 2 and H 2O and associated O 2 expenditure.
       After that, synthesis continues. In export pro-
                                       The Krebs cycle (citric acid cycle), respiratory
                                       chain and related ATP synthesis also occur in
       the peptide chain to the cisternal space once
                                       mitochondria. Cells intensely active in meta-
       synthesis is completed. Synthesis of membrane
       proteins is interrupted several times (depend-
                                       bolic and transport activities are rich in mito-
       ing on the number of membrane-spanning
                                       chondria—e.g., hepatocytes, intestinal cells,
       domains (! G2) by translocator protein clo-
                                       and renal epithelial cells. Mitochondria are en-
       peptide sequence is pushed into the phos-
                                       smooth outer membrane and an inner mem-
                                       brane. The latter is deeply infolded, forming a
       pholipid membrane. The smooth endoplasmic
    1  sure, and the corresponding (hydrophobic)  closed in a double membrane consisting of a
                                       series of projections (cristae); it also has im-
       reticulum (SER) contains no ribosomes and is
       the production site of lipids (e.g., for lipo-  portant transport functions (! p. 17 B). Mito-
       proteins, ! p. 254 ff.) and other substances.  chondria probably evolved as a result of sym-
       The ER membrane containing the synthesized  biosis between aerobic bacteria and anaerobic
       membrane proteins or export proteins forms  cells (symbiosis hypothesis). The mitochondrial
       vesicles which are transported to the Golgi ap-  DNA (mtDNA) of bacterial origin and the
       paratus.                        double membrane of mitochondria are relicts
         The Golgi complex or Golgi apparatus (! F)  of their ancient history. Mitochondria also
       has sequentially linked functional compart-  contain ribosomes which synthesize all pro-
       ments for further processing of products from  teins encoded by mtDNA.
       the endoplasmic reticulum. It consists of a cis-  Lysosomes are vesicles (! F) that arise from
       Golgi network (entry side facing the ER),  the ER (via the Golgi apparatus) and are in-
       stacked flattened cisternae (Golgi stacks) and a  volved in the intracellular digestion of macro-
       trans-Golgi network (sorting and distribution).  molecules. These are taken up into the cell
       Functions of the Golgi complex:  either by endocytosis (e.g., uptake of albumin
       ! polysaccharide synthesis;     into the renal tubules; ! p. 158) or by phagocy-
       ! protein processing (posttranslational modi-  tosis (e.g., uptake of bacteria by macrophages;
       fication), e.g., glycosylation of membrane pro-  ! p. 94 ff.). They may also originate from the
       teins on certain amino acids (in part in the ER)  degradation of a cell’s own organelles (auto-
       that are later borne as glycocalyces on the ex-  phagia, e.g., of mitochondria) delivered inside
       ternal cell surface (see below) and γ-carboxy-  autophagosomes (! B, F). A portion of the en-
       lation of glutamate residues (! p. 102 );  docytosed membrane material recycles (e.g.,
       ! phosphorylation of sugars of glycoproteins  receptor recycling in receptor-mediated en-
       (e.g., to mannose-6-phosphate, as described  docytosis; ! p. 28). Early and late endosomes
       below);                         are intermediate stages in this vesicular trans-
       ! “packaging” of proteins meant for export  port. Late endosomes and lysosomes contain
       into secretory vesicles (secretory granules), the  acidic hydrolases (proteases, nucleases, li-
       contents of which are exocytosed into the ex-  pases, glycosidases, phosphatases, etc., that
   12  tracellular space; see p. 246, for example.  are active only under acidic conditions). The
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       Despopoulos, Color Atlas of Physiology © 2003 Thieme
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