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208       Organelles



             Nucleus                                          proteins can enter the nucleus without dif -
                                                              culty. By contrast, larger proteins (over
                                                              40 kDa) can only pass through the nuclear
             A. Nucleus
                                                              pores if they carry a nuclear localization se-
             The nucleus is the largest organelle in the eu-  quence consisting of four successive basic
             karyotic cell. With a diameter of about 10 µm,   amino acids inside their peptide chains (see
             it is easily recognizable with the light micro-  p. 228). mRNAs and rRNAs formed in the nu-
             scope. This is the location for storage, replica-  cleus cross the pores into the cytoplasm as
             tion, and expression of genetic information.     complexes with proteins (see below).
                The nucleus is separated from the cyto-
             plasm by the nuclear envelope, which consists    C. Relationships between the nucleus and
             of the outer and inner nuclear membranes.
                                                              cytoplasm
             Each of the two nuclear membranes has two
             layers, and the membranes are separated          Almost all of the RNA in the cell is synthesized
             from each other by the perinuclear space.        in the nucleus. In this process, known as
             The outer nuclear membrane is continuous         transcription, the information stored in DNA
             with the rough endoplasmic reticulum and         is transcribed into RNA (see p. 242). As men-
             is covered with ribosomes. The inner side of     tioned above, ribosomal RNA (rRNA) is mainly
             the membrane is covered with a protein layer     produced in the nucleolus, while messenger
             (the nuclear lamina), in which the nuclear       and transfer RNA (mRNA and tRNA) are
             structures are anchored.                         formed in the region of the euchromatin. En-
                The nucleus contains almost all of the cell’s  zymatic duplication of DNA—replication—also
             DNA (around 1% of which is mitochondrial         only takes place in the nucleus (see p. 240).
             DNA). Together with histones and structural         Thenucleotidecomponents requiredfor
             proteins, the nuclear DNA forms the chroma-      transcription and replication have to be im-
             tin (see p. 238). It is only during cell division  ported into the nucleus from the cytoplasm.
             that chromatin condenses into chromosomes,       Incorporation of these components into RNA
             which are also visible with the light micro-     leads to primary products, which are then
             scope. During this phase, the nuclear mem-       altered by cleavage, excision of introns, and
             brane temporarily disintegrates.                 the addition of extra nucleotides (RNA matu-
                During the phase between cell divisions,      ration; see p. 242). It is only once these pro-
             the interphase, it is possible to distinguish    cess have been completed that the RNA mol-
             between the more densely packed hetero-          ecules formed in the nucleus can be exported
             chromatin and loose euchromatin using an         into the cytoplasm for protein synthesis
             electron microscope. Active transcription of     (translation;see p. 250).
             DNA into mRNA takes place in the region of          The nucleus is not capable of synthesizing
             the euchromatin. A particularly electron-        proteins. All of the nuclear proteins therefore
             dense    region   is  noticeable   in   many     have to be imported—the histones with which
             nuclei—the nucleolus (several nucleoli are       DNA is associated in chromatin, and also the
             sometimes present). The DNA in the nucleolus     so–called non–histone proteins (DNA poly-
             contains numerous copies of the genes for        merases and RNA polymerases, auxiliary and
             rRNAs (see p. 242). They are constantly under-   structural proteins, transcription factors, and
             going transcription, leading to a high local     ribosomal proteins). Ribosomal RNA (rRNA)
             concentration of RNA.                            already associates with proteins in the nucle-
                                                              olus to form ribosome precursors.
                                                                 A special metabolic task carried out by the
             B. Nuclear pores                                                               +
                                                              nucleus is biosynthesis of NAD . The immedi-
             The exchange of substances between the nu-       ate precursor of this coenzyme, nicotinamide
                                                                                    +
             cleus and the cytoplasm is mediated by pore      mononucleotide (NMN ), arises in the cyto-
             complexes    with   complicated    structures,   plasm and is then transported into the nucle-
             which traverse the nuclear membrane. The         olus, where it is enzymatically converted into
                                                                                   +
                                                                                                 +
             nuclear pores consist of numerous proteins       the dinucleotide NAD . Finally, NAD then re-
             that form several connected rings of varying     turns to the cytoplasm.
             diameter. Low-molecular structures and small


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