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186    Chapter 6    DNA Structure, Replication, and Recombination


              Figure 6.6  Experiments with viruses provide convincing evidence that genes are made of DNA. (a) and (b) Bacteriophage
              T2 structure and life cycle. The phage particle consists of DNA contained within a protein coat. The virus attaches to the bacterial host cell and
              injects its genes (the DNA) through the bacterial cell wall into the host cell cytoplasm. Inside the host cell, these genes direct the formation of
              new phage DNA and proteins, which assemble into progeny phages that are released into the environment when the cell bursts.
                  (a)                                        (b)
                      Protein coat                                                          1. Phage attaches
                                                                                                to bacterium
                         DNA                                                                    (host).
                                                             5. Cell bursts,
                                                                 releasing new
                                                                 phages.



                                                                                                     2. Phage injects
                                                                                                         its genes into
                                          Core                                                           host cell.
                          Host cell wall                            4. Phage
                                                                        particles
                                                                        assemble.         3. Phage DNA replicates;
                       T2 phage                                                               new phage proteins
                                                                                              are made.




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              Figure 6.7  The Hershey-Chase Waring blender experiment. T2 bacteriophage particles either with  P-labeled DNA (orange) or
                  35
              with  S-labeled proteins (purple) were used to infect bacterial cells. After a short incubation, Hershey and Chase shook the cultures in a
              Waring blender and spun the samples in a centrifuge to separate the empty viral ghosts from the heavier infected cells. Most of the
              35 S-labeled proteins remained with the ghosts, while most of the  P-labeled T2 DNA was found in the sedimented infected cells.
                                                              32
                                                                                                      Ghosts
                                                                32
                                                                 P DNA
                                                                                              Blend
                                                                                              briefly
                                                                                               Radioactivity     Cell
                 T2 phage        Infect E. coli and grow  Phages with       Introduce phages into  recovered in host and
                                   32
                                 in  P-containing medium.  32 P-labeled DNA.  bacteria culture.  passed on to phage progeny.
                                                                                                       Ghosts
                                                                35
                                                                 S protein
                                                                                              Blend
                                                                                              briefly
                                                                                                                 Cell
                                                                                                    Radioactivity
                 T2 phage        Infect E. coli and grow  Phages with        Introduce phages into  recovered
                                   35
                                 in  S-containing medium.  35 S-labeled protein.   bacteria culture.  in phage ghosts.


              maintained in two different culture media, one infused with   each one, effectively separating the viral ghosts from the
                                           32
              radioactively labeled phosphorus ( P), the other with radi-  bacteria harboring the viral genes. Centrifugation of the
                                   35
              oactively labeled sulfur ( S). Because proteins incorporate   cultures then separated the heavier infected cells, which
              sulfur but no phosphorus and DNA contains phosphorus   ended up in a pellet at the bottom of the tube, away from
                                          35
              but no sulfur, phages grown on  S would have radioac-  the lighter phage ghosts, which remained suspended in
                                                                                                               32
                                                       32
              tively labeled protein while particles grown on  P would   the supernatant solution. Most of the radioactive  P (in
              have radioactive DNA. The radioactive tags would serve as   DNA) went to the pellet, while most of the radioactive
              markers for the location of each material when the phages   35 S (in protein) remained in the supernatant. This result
              infected fresh cultures of bacterial cells.          confirmed  that  the  extracellular  ghosts  were  indeed
                  After exposing one fresh culture of bacteria to   mostly protein, while the injected viral material specify-
                                                      35
              32 P-labeled phages and another culture to  S-labeled   ing production of more phages was mostly DNA. Bacte-
              phages, Hershey and Chase used a Waring blender to disrupt   ria containing the radio-labeled phage DNA behaved just
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