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PART II  What Genes Are and What They Do
                                6

                       chapter



                                    DNA Structure,



                                 Replication, and


                                   Recombination




                                                                            The double-helical structure of DNA provides an explanation for
                                                                            the transmission of genetic information from generation to
                                                                              generation over billions of years.
                                                                            © Adrian Neal/Getty Images RF

                                                                            chapter outline

                                                                            •     6.1 Experimental Evidence for DNA as the
                                                                              Genetic Material
                                                                            •     6.2 The Watson and Crick Double Helix Model
                                                                              of DNA
                       FOR NEARLY 4 BILLION YEARS, the double-stranded DNA   •     6.3 Genetic Information in Nucleotide Sequence
                       molecule has served as the bearer of genetic information.   •     6.4 DNA Replication
                       It was present in the earliest single-celled organisms and   •     6.5 Homologous Recombination at the DNA Level
                       in every other organism that has existed since. Over that
                       long period of time, the hardware—the structure of the   •     6.6 Site-Specific Recombination
                       molecule itself—has not changed. In contrast, evolution
                       has honed and vastly expanded the  software—the pro-
                       grams of genetic information that the molecule stores, expresses, and transmits from
                       one generation to the next.
                          Under special conditions of little or no oxygen, DNA can withstand a wide range
                       of temperature, pressure, and humidity and remain relatively intact for hundreds, thou-
                       sands, even tens of thousands of years. Molecular sleuths have retrieved the evidence:
                       38,000-year-old DNA from a Neanderthal skeleton (Fig. 6.1). Amazingly, this ancient
                       DNA still carries readable sequences—shards of decipherable information that act as
                       time machines for the viewing of genes in this long-vanished species. Comparisons
                       with homologous DNA segments from living people make it possible to identify the
                       precise mutations that have fueled evolution.
                          For example, comparisons of Neanderthal and human DNA have helped anthro-
                       pologists settle a long-running debate about the genetic relationship of the two. The
                       evidence shows that Neanderthals and our own species, Homo sapiens, last shared a
                       common ancestor between 600,000 and 800,000 years ago. Neanderthal ancestors
                       migrated to Europe about 400,000 years ago while our own ancestors remained in
                         Africa. The two groups remained out of contact until 40,000 years ago, when Homo
                       sapiens first arrived in Europe. Within a few millennia, the Neanderthals were extinct.
                       However, their recently recovered DNA suggests that during the 10,000 years that
                       Neanderthals shared Europe with Homo sapiens, some interbreeding took place; 1–4%
                       of the genomes of modern non-Africans can be traced to Neanderthals.
                          Francis Crick, co-discoverer of DNA’s double-helical structure and a leading
                       twentieth-century theoretician of molecular biology, wrote that “almost all aspects
                       of life are engineered at the molecular level, and without understanding molecules,

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