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Arenaviridae (lymphocytic choriomeningitis virus) Togaviridae (rubella virus) Flaviviridae (yellow fever virus) Retroviridae (human immunodeficiency virus) Picomaviridae (human poliovirus 1, human hepatitis A virus) Caliciviridae (vesicular exanthema of swine virus)
100 nm
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Figure 16.28 Representatives of families of animal viruses. The abbreviations ss and ds refer to single-stranded and double-stranded, respectively. (From R. I. B. Francki, et al., Classi cation and Nomenclature of Viruses, fth report, 1991. Springer-Verlag, Vienna. Reprinted by permission.)
the host organism. To the surprise of virtually everyone, these oncogenes were found in untransformed cells. Since transforming viruses can function quite well as viruses without their oncogenes, and since cellular oncogenes have introns and viral oncogenes do not, geneticists generally accept the theory that these oncogenes originated in the host and were picked up, presumably as messenger RNAs, by the retroviruses. We believe that retroviruses pick up cellular genes by transcription readthrough, transcribing beyond the end of the integrated virus and producing a messenger RNA that is then incorporated into a viral particle after intron removal. Retroviruses can thus pick up genes adjacent to their point of integration. To distinguish oncogenes within viruses and hosts, we pre x the name of a viral oncogene, such as src, with a v (v-src) and a cellular oncogene with a c (c-src). Cellu-
lar oncogenes within a nontransformed cell are called proto-oncogenes. How are proto-oncogenes induced to become oncogenes, and what do proto-oncogenes normally do in the cell
Oncogene Induction
Proto-oncogenes can be induced in at least three different ways. First, a mutation can cause a proto-oncogene to transform its host cell. For example, a ras proto-oncogene (see table 16.4) was converted to an oncogene when one codon, GGC (glycine), was converted to GTC (valine). Second, a proto-oncogene can be activated if it is moved to a region with a strong promoter or enhancer. Burkitt s lymphoma, for example, is associated with a translocation involving the proto-oncogene c-myc, which is normally located on chromosome 8. When translocated to
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Sixteen
Gene Expression: Control in Eukaryotes
Table 16.4 Some Oncogenes, Their Origins, and Their Protein Products
Oncogene abl src erbB fms mos sis Ha-ras Ki-ras fos myb erbA rel jun Virus Abelson murine leukemia virus Rous sarcoma virus Avian erythroblastosis virus McDonough feline sarcoma virus Avian myeloblastosis virus Simian sarcoma virus Harvey murine sarcoma virus Kirsten murine sarcoma virus FBJ osteosarcoma virus Avian myeloblastosis virus Avian erythroblastosis virus Reticuloendotheliosis virus Avian sarcoma virus 17 Species of Origin Mouse Chicken Chicken Cat Chicken Woolly monkey Rat Rat Mouse Chicken Chicken Turkey Chicken Gene Function Tyrosine kinase Tyrosine kinase Tyrosine kinase Growth factor Protein kinase Growth factor GTP-binding protein GTP-binding protein Binds DNA Binds DNA Binds DNA Binds DNA Binds DNA
Source: Reprinted with permission from J. Marx, What Do Oncogenes Do , Science, 223:673 76, 1984. Copyright 1984 American Association for the Advancement of Science.
A retroviral RNA genome. R is a repeated sequence; U3 and U5 are unique sequences; PBS is the primer-binding site, LTR is the long terminal repeat; gag, pol, and env are viral genes. During the process of reverse transcription, the LTRs are created at the ends of the DNA. Direct host repeats are created when the viral DNA integrates into the host chromosome.
chromosome 14, c-myc is placed contiguous with the immunoglobulin IgM constant gene. This gene is very active in lymphocytes (as we will see later). Hence c-myc is now transcribed at a much higher rate than normal, resulting in cellular transformation. The c-myc gene normally occurs near a fragile site, a region of a chromosome that has a tendency to break. Many protooncogenes occur near fragile sites on chromosomes.
The simple capture of a gene by a retrovirus might be enough for transformation, since the gene is brought under the influence of viral transcriptional control. However, not all genes captured this way are oncogenes. Third, a proto-oncogene can be activated if it is ampli ed. Several cases are known in which ampli ed genes (e.g., c-ras and c-abl) or genes on trisomic chromosomes are related to transformation.
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