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Valuable information about the nature of the genetic material has also come from viruses. Of particular value are studies of bacterial viruses the bacteriophages, or phages. Since phages consist only of nucleic acid surrounded by protein, they lend themselves nicely to the determination of whether the protein or the nucleic acid is the genetic material. A. D. Hershey and M. Chase published, in 1952, the results of research that supported the notion that DNA is the genetic material and, in the process, helped to
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A. D. Hershey (1908 1997). (Courtesy of
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Dr. A. D. Hershey.)
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of Cold Springs Harbor Laboratory Archives.)
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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9. Chemistry of the Gene1
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The McGraw Hill Companies, 2001
Nine
Chemistry of the Gene
S-type cells
(Dead)
R-type cells
(No change)
Heat-killed S-type cells
(No change)
R-type plus heat-killed S-type cells
(Dead) Figure 9.5
Grif th s experiment with Streptococcus. S-type cells will kill mice; so will heat-killed S-type cells injected with live R-type cells. S-type cells are recovered from dead mice in both cases.
explain the nature of the viral infection process. Since all nucleic acids contain phosphorus, whereas proteins do not, and since most proteins contain sulfur (in the amino acids cysteine and methionine), whereas nucleic acids do not, Hershey and Chase designed an experiment using radioactive isotopes of sulfur and phosphorus to keep separate track of the viral proteins and nucleic acids during the infection process. They used the T2 bacteriophage and the bacterium Escherichia coli. The phages were labeled by having them infect bacteria growing in culture medium containing the radioactive isotopes 35S or 32P. Hershey and Chase then proceeded to identify the material injected into the cell by phages attached to the bacterial wall. When 32P-labeled phages were mixed with unlabeled E. coli cells, Hershey and Chase found that the 32P label entered the bacterial cells and that the next generation of phages that burst from the infected cells carried a significant amount of the 32P label. When 35S-labeled phages
were mixed with unlabeled E. coli, the researchers found that the 35S label stayed outside the bacteria for the most part. Hershey and Chase thus demonstrated that the outer protein coat of a phage does not enter the bacterium it infects, whereas the phage s inner material, consisting of DNA, does enter the bacterial cell ( g. 9.6). Since the DNA is responsible for the production of the new phages during the infection process, the DNA, not the protein, must be the genetic material.
RNA as Genetic Material
In some viruses, RNA (ribonucleic acid) is the genetic material. The tobacco mosaic virus that infects tobacco plants consists only of RNA and protein.The single, long RNA molecule is packaged within a rodlike structure formed by over two thousand copies of a single protein. No DNA is present in tobacco mosaic virus particles ( g. 9.7a). In 1955, H. Fraenkel-Conrat and R. Williams
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
9. Chemistry of the Gene1
The McGraw Hill Companies, 2001
Chemistry of Nucleic Acids
Bacterium in 35S medium
Bacterium in 32P medium
Phage (unlabeled)
Progeny phage with protein and unlabeled DNA
35S-labeled
Labeled phage progeny released
Progeny phage with unlabeled protein and 32P-labeled DNA
Labeled protein stays outside, unlabeled DNA enters cell
Infects unlabeled bacteria
Unlabeled protein stays outside, labeled DNA enters cell
Progeny phage virtually unlabeled (less than 1% of original 35S recovered) Figure 9.6
Progeny phage strongly labeled (more than 30% of original 32P recovered)
The Hershey and Chase experiments using 35S-labeled and 32P-labeled T2 bacteriophages. The nucleic acid label (32P) enters the E. coli bacteria during infection; the protein label (35S) does not.
showed that a virus can be separated, in vitro, into its component parts and reconstituted as a viable virus. This nding led Fraenkel-Conrat and B. Singer to reconstitute tobacco mosaic virus with parts from different strains ( g. 9.7b). For example, they combined the RNA from the common tobacco mosaic virus with the protein from the masked (M) strain of tobacco mosaic virus. They then made the reciprocal combination of common-type protein and M-type RNA. In both cases, the tobacco mosaic virus produced during the process of infection was the type associated with the RNA, not with the protein. Thus, it was the nucleic acid (RNA in this case) that was the genetic material. Subsequently, scientists rubbed pure tobacco mosaic virus RNA into plant leaves. Normal infection and a new generation of typical, protein-coated tobacco mosaic virus resulted, con rming RNA as the genetic material for this virus. We thus conclude that DNA is the genetic material. In the few viruses that do not have DNA, RNA serves as the
genetic material. The only exception to these statements is one type of disease that is transmitted by a protein without accompanying DNA or RNA (box 9.2).
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