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84        Biomolecules



             DNA                                              sugar and phosphate residues in the back-
                                                              bone. Along the whole length of the DNA
                                                              molecule, there are two depressions—re-
             A. DNA: structure
                                                              ferred to as the “minor groove” and the “ma-
             Like RNAs (see p. 82), deoxyribonucleic acids    jor groove”—that lie between the strands.
             (DNAs) are polymeric molecules consisting of
             nucleotide building blocks. Instead of ribose,
                                                              B. Coding of genetic information
             however, DNA contains 2 -deoxyribose, and
             the uracil base in RNA is replaced by thymine.   In all living cells, DNA serves to store genetic
             Thespatial structureof the two molecules         information.  Specific  segments    of  DNA
             also differs (see p. 86).                        (“genes”) are transcribed as needed into
                The first evidence of the special structure   RNAs, which either carry out structural or
             of DNA was the observation that the amounts      catalytic tasks themselves or provide the basis
             of adenine and thymine are almost equal in       for synthesizing proteins (see p. 82). In the
             every type of DNA. The same applies to gua-      latter case, the DNA codes for the primary
             nine and cytosine. The model of DNA struc-       structure of proteins. The “language” used in
             ture formulated in 1953 explains these con-      this process has four letters (A, G, C, and T). All
             stant base ratios: intact DNA consists of two    of the words (“codons”) contain three letters
             polydeoxynucleotide molecules (“strands”).       (“triplets”), and each triplet stands for one of
             Each base in one strand is linked to a comple-   the 20 proteinogenic amino acids.
             mentary base in the other strand by H-bonds.        Thetwo strands ofDNA arenot function-
             Adenine is complementary to thymine, and         ally equivalent. The template strand (the (–)
             guanine is complementary to cytosine. One        strand or “codogenic strand,” shown in light
             purine base and one pyrimidine base are          gray in Fig.1) is the one that is read during the
             thus involved in each base pair.                 synthesis of RNA (transcription; see p. 242).
                The complementarity of A with T and of G      Its sequence is complementary to the RNA
             with C can be understood by considering the      formed. The sense strand (the (+) strand or
             H bonds that arepossiblebetween thediffer-       “coding strand,” shown in color in Figs. 1 and
             ent bases. Potential donors (see p. 6) are       2)has the same sequence as the RNA, except
             amino groups (Ade, Cyt, Gua) and ring NH         that T is exchanged for U. By convention, it is
             groups. Possible acceptors are carbonyl oxy-     agreed that gene sequences are expressed by
             gen atoms (Thy, Cyt, Gua) and ring nitrogen      reading the sequence of the sense strand in
             atoms. Two linear and therefore highly stable    the 5  3  direction. Using the genetic code
             bonds can thus be formed in A–T pairs, and       (see p. 248), in this case the protein sequence
             three in G–C pairs.                              (3) is obtained directly in the reading direc-
                Base pairings of this type are only possible,  tion usual for proteins—i. e., from the N termi-
             however, when the polarity of the two strands    nus to the C terminus.
             differs—i. e., when they run in opposite direc-
             tions (see p. 80). In addition, the two strands
             have to be intertwined to form a double helix.
             Due to steric hindrance by the 2 -OH groups
             of theriboseresidues, RNA is unable to form a
             double helix. The structure of RNA is therefore
             less regular than that of DNA (see p. 82).
                The conformation of DNA that predomi-
             nates within the cell (known as B-DNA)is
             shown schematically in Fig. A2 and as a van
             der Waals model in Fig. B1.In the schematic
             diagram (A2), the deoxyribose–phosphate
             “backbone” is shown as a ribbon. The bases
             (indicated by lines) are located on the inside
             of the double helix. This area of DNA is there-
             fore apolar. By contrast, the molecule’s surface
             is polar and negatively charged, due to the


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