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Retroviruses
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Animal viruses come in many different varieties, with DNA or RNA as their genetic material ( g. 16.28). Several classes of viruses, both DNA and RNA, can transform cells, a process that may or may not be caused by an oncogene the virus carries. Some DNA viruses do carry oncogenes, such as the adenovirus that carries the gene for the E1A protein, which may act by binding to the retinoblastoma repressor protein. Oncogenes, however, were originally discovered in retroviruses, a group of very simple RNA viruses that contain the enzyme reverse transcriptase. After the virus enters the host cell, this enzyme converts the viral RNA into DNA. In 1910, Peyton Rous, who much later won the Nobel Prize for his work, discovered that a sarcoma in chickens could be induced by a cell-free extract from a tumor in another chicken. The transmitted agent was later found to be a retrovirus, named Rous sarcoma virus, the rst retrovirus to be discovered.
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Peyton Rous (1879 1970).
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(Courtesy of Rockefeller University Archives.)
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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16. Gene Expression: Control in Eukaryotes
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The McGraw Hill Companies, 2001
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Gene Expression: Control in Eukaryotes
p53 No DNA damage
Free MDM2 Degradation of p53 in proteasome
MDM2
Activate DNA DNA damage p53 Phosphorylation of p53 (active transcription factor) CHK2 P
P DNA
Stop cell cycle Figure 16.27
Induce apoptosis
Induce MDM2
Normally, MDM2 ubiquinates the p53 protein, which leads to its degradation in the proteasome. If there is DNA damage that produces broken DNA ends (double-strand breaks), MDM2 binds to these ends and activates the checkpoint kinase CHK2. CHK2 phosphorylates p53, making it stable and an active transcription factor. Next, the active p53 binds to the promoters of synexpression group genes for stopping the cell cycle and apoptosis. In addition, p53 induces the MDM2 gene, which can then cause p53 degradation if the broken DNA has been repaired. (MDM2 is named for the cell culture in which it was cloned murine double minute chromosome clone number 2.)
The retrovirus, which often carries only three genes, integrates into the host genome in a series of steps ( g. 16.29).When the virus enters the host, it is in the form of a plus ( ) RNA strand (capable of acting as a messenger RNA; the minus strand is the complement to the strand). At either end is a repeated sequence (R) located outside two unique sequences (U3 and U5). Through reverse transcription, using the reverse transcriptase the virus brings in, the viral RNA is converted to a doublestranded DNA. During that process, the ends of the DNA take on the con guration of long terminal repeats (LTRs), repetitions of U3-R-U5. The linear DNA then circularizes and integrates into the host genome just as a transposon does, generating short direct repeats at either end. As we mentioned, retroviruses can cause cellular transformation through direct integration or from the oncogenes they carry. Transformation from integration comes about because the integrated provirus either inactivates a tumor-suppressor gene or activates an oncogene
in a process called insertion mutagenesis. The U3 region of the retrovirus contains both an enhancer and a promoter. Since a long terminal repeat lies at either end of the provirus, cellular genes can be turned on when the virus integrates.
Oncogenes
Genetic analysis and recombinant DNA studies showed that Rous sarcoma virus transforms cells through the action of a single gene. This gene, called src for sarcoma, was the rst viral oncogene discovered. Since then at least fty have been discovered, and each has been given a three-letter designation (table 16.4). Unlike tumor suppressors, which lead to cancer when in the homozygous mutant condition, oncogenes act in a dominant fashion: Only one copy of the activated gene need be present for transformation to occur. With the viral oncogene in hand, researchers could create a probe for the gene and look within the DNA of
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Cancer
Enveloped
dsDNA dsDNA
Nonenveloped
Adenoviridae (human adenovirus 2) Herpesviridae (herpes simplex virus) Poxviridae, Choropoxvirinae (vaccinia virus) Papovaviridae (polyoma virus)
Iridoviridae (frog virus)
Hepadnaviridae (hepatitis B virus)
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