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6.2 The Watson and Crick Double Helix Model of DNA   189


                       other end of the chain—the 3′ end—it is the 3′ carbon of the   published their findings in the scientific journal Nature in
                       final nucleotide that is free. Along the chain between the two   April 1953.
                       ends, this 5′-to-3′ polarity is conserved from nucleotide to
                       nucleotide. By convention, a DNA chain is described in
                       terms of its bases, written with the 5′-to-3′ direction going   Evidence from X-ray diffraction
                       from left to right (unless otherwise noted). The chain   The diffraction patterns of oriented DNA fibers do not, on
                         depicted in Fig. 6.9c, for instance, would be 5′ TACG 3′.  their own, contain sufficient information to reveal structure.
                                                                           For instance, the number of diffraction spots, whose intensi-
                                                                           ties and positions constitute the X-ray data (review Fig. 6.8), is
                       DNA’s information content                           considerably lower than the number of unknown coordinates
                       Information can be encoded only in a sequence of symbols   of all the atoms in an oriented DNA molecule. Nevertheless,
                       whose order varies according to the message to be conveyed.   the photographs do reveal a wealth of structural information to
                       Without this sequence variation, there is no potential for car-  the trained eye. Excellent X-ray images produced by Rosalind
                       rying information. Because DNA’s backbone of alternating   Franklin and Maurice Wilkins showed that the molecule is
                       sugar and phosphate is chemically identical for every nucle-  spiral-shaped, or helical; the spacing between repeating units
                       otide in a DNA chain, the only difference between nucleo-  along the axis of the helix is 3.4 Å (3.4 × 10 −10  meters); the
                       tides is in the identity of the nitrogenous base. Thus, the   helix undergoes one complete turn every 34 Å; and the diam-
                       genetic information in DNA must consist of variations in the   eter of the molecule is 20 Å. This diameter is roughly twice the
                       sequence of the A, G, T, and C bases. The information con-  width of a single nucleotide, suggesting that a DNA molecule
                       structed from the four-letter language of DNA bases is anal-  might be composed of two side-by-side DNA chains.
                       ogous to the information built from the 26-letter alphabet of
                       English or French or Italian. Just as you can combine the
                       26 letters of the alphabet in different ways to generate the   Complementary base pairing
                       words of a book, so, too, different combinations of the four   If a DNA molecule contains two side-by-side chains of
                       bases in very long sequences of nucleotides can encode the   nucleotides, what forces hold these chains together? Erwin
                       information for constructing an organism.           Chargaff provided an important clue with his data on the
                                                                           nucleotide  composition  of  DNA  from  various  species.
                                                                             Despite large variations in the relative amounts of the
                       The DNA Helix Consists of Two                       bases, the ratio of A to T is not significantly different from
                       Antiparallel Chains                                 1:1, and the ratio of G to C is also the same in every organ-
                                                                           ism (Table 6.1). Watson grasped that the roughly 1:1 ratios
                       Watson and Crick’s discovery of the structure of the DNA   of A to T and of G to C reflect a significant aspect of the
                       molecule ranks with Darwin’s theory of evolution by natu-  molecule’s inherent structure.
                       ral selection and Mendel’s laws of inheritance in its contri-  To explain Chargaff’s ratios in terms of chemical affin-
                       bution to our understanding of biological phenomena. The   ities between A and T and between G and C, Watson made
                       Watson-Crick structure, first embodied in a model that   cardboard cutouts of the bases in the chemical forms they
                       superficially resembled the Tinker Toys of preschool chil-  assume in a normal cellular environment. He then tried to
                       dren, was based on an interpretation of all the chemical   match these up in various combinations, like pieces in a
                       and physical data available at the time. Watson and Crick   jigsaw puzzle. He knew that the particular arrangements of



                           TABLE 6.1      Chargaff’s Data on Nucleotide Base Composition in the DNA of Various Organisms
                                                               Percentage of Base in DNA                       Ratios

                        Organism                        A           T            G             C           A:T         G:C
                        Escherichia coli              24.7         23.6         26.0         25.7         1.05         1.01
                        Saccharomyces cerevisiae      31.3         32.9         18.7         17.1         0.95         1.09
                        Caenorhabditis elegans        31.2         29.1         19.3         20.5         1.07         0.96
                        Drosophila melanogaster       27.3         27.6         22.5         22.5         0.99         1.00
                        Mus musculus                  29.2         29.4         21.7         19.7         0.99         1.10
                        Homo sapiens                  29.3         30.0         20.7         20.0         0.98         1.04

                       Note that even though the level of any one nucleotide is different in different organisms, the amount of A always approximately equals the amount of T, and the level of
                       G is always similar to that of C. Moreover, the total amount of purines (A plus G) nearly always equals the total amount of pyrimidines (C plus T).
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