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226       Organelles



             ER: structure and function                       will take and whether its transport will be
                                                              constitutive or regulated depends on the sig-
             The endoplasmic reticulum (ER) is an exten-      nal sequences or signal structures that pro-
             sive closed membrane system consisting of        teins carry with them like address labels (see
             tubular and saccular structures. In the area     p. 228). In addition to proteins, the Golgi ap-
             of the nucleus, the ER turns into the external   paratus also transports membrane lipids to
             nuclear membrane. Morphologically, a dis-        their targets.
             tinction is made between the rough ER (rER)
             and the smooth ER (sER). Large numbers of
             ribosomes are found on the membranes of the      B. Smooth endoplasmic reticulum
             rER, whichare lackingonthe sER. On the           Regions of the ER that have no bound ribo-
             other hand, the sER is rich in membrane-         somesare known asthe smooth endoplasmic
             bound enzymes, which catalyze partial reac-      reticulum (sER). In most cells, the proportion
             tions in the lipid metabolism as well as bio-    representedbythe sER is small. AmarkedsER
             transformations.                                 is seen in cells that have an active lipid me-
                                                              tabolism, such as hepatocytes and Leydig
                                                              cells. The sER is usually made up of branching,
             A. Rough endoplasmic reticulum and the
                                                              closed tubules.
             Golgi apparatus
                                                                 Membrane-located enzymes in the sER
             The rER (1)is a site of active protein           catalyze lipid synthesis. Phospholipid synthe-
             biosynthesis. This is where proteins destined    sis (see p. 170) is located in the sER, for exam-
             for membranes, lysosomes, and export from        ple, and several steps in cholesterol biosyn-
             the cell are synthesized. The remaining pro-     thesis (see p. 172) also take place there. In
             teins are produced in the cytoplasm on ribo-     endocrine cells that form steroid hormones, a
             somes that are not bound to membranes.           large proportion of the reaction steps in-
                Proteins synthesized at the rER (1)are        volved also take place in the sER (see p. 376).
             folded and modified after translation (protein      In the liver’s hepatocytes, the proportion
             maturation; see p. 230). They remain either in   represented by the sER is particularly high.
             the rER as membrane proteins, or pass with       It contains enzymes that catalyze so-called
             the help of transport vesicles (2)to the Golgi   biotransformations. These are reactions in
             apparatus (3). Transport vesicles are formed     which apolar foreign substances, as well as
             by buddingfrom existingmembranes,and             endogenous substances—e. g., steroid hormo-
             they disappear again by fusing with them         nes—are chemically altered in order to inacti-
             (see p. 228).                                    vate them and/or prepare them for conjuga-
                The Golgi apparatus (3)is a complexnet-       tion with polar substances (phase I reactions;
             work, also enclosed, consisting of flattened     see p. 316). Numerous cytochrome P450
             membrane saccules (“cisterns”), which are        enzymes are involved in these conversions
             stacked on top of each other in layers. Pro-     (see p. 318) and can therefore be regarded
             teins maturehereand aresorted and packed.        as the major molecules of the sER.
             A distinction is made between the cis, medial,      The sER also functions as an intracellular
             and trans Golgi regions, aswell asa trans        calcium store, which normally keeps the Ca 2+
             Golgi network (tGN). The post–translational      level in the cytoplasm low. This function is
             modification of proteins, which starts in the    particularly marked in the sarcoplasmic retic-
             ER, continues in these sections.                 ulum, a specialized form of the sER in muscle
                From the Golgi apparatus, the proteins are    cells (see p. 334). For release and uptake of
                                                                2+
             transported by vesicles to various targets in    Ca , the membranes of the sER contain sig-
             the cells—e. g., to lysosomes (4), the plasma    nal-controlled Ca 2+  channels and energy-de-
             membrane (6), and secretory vesicles (5)that     pendent Ca 2+  ATPases (seep.220). In thelu-
             release their contents into the extracellular    men of the sER, the high Ca 2+  concentration is
                                                                            2+
             space by fusion with the plasma membrane         buffered by Ca -binding proteins.
             (exocytosis; see p. 228). Protein transport can
             either proceed continuously (constitutive), or
             it can be regulated by chemical signals. The
             decision regarding which pathway a protein


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