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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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Viral oncogenes can cause transformation by the same mechanisms. Either a mutation of the oncogene itself or the placement of the gene next to an active viral promoter can cause high levels of transcription of the oncogene and, hence, transformation of the cell. What are the gene products of these proto-oncogenes
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Oncogene Function
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We know that proto-oncogenes are important to the cell because they have been conserved evolutionarily. For example, c-src is found in fruit ies as well as in vertebrates; c-ras is found in yeasts and in human beings. They are all genes that can promote cell growth; they are speci c transcription factors or other components of growth-stimulating signal pathways. The known protein products of oncogenes can be classified into at least four categories: tyrosine kinases, growth factors, GTP-binding proteins, and DNA-binding proteins (see table 16.4). As proto-oncogenes, they normally function at low levels; in transformed cells, they function at high levels. Can these proteins explain cancerous growth Tyrosine kinases are enzymes that add a phosphate group to tyrosine residues in proteins. Other kinases phosphorylate serine and threonine. (These three amino acids have OH groups available for phosphorylation.) Proteins that are phosphorylated at their tyrosine residues are involved in signal pathways, cytoskeleton shape (transformed cells are shaped differently than normal cells), and glycolysis (cancer cells tend toward the anaerobic glycolytic pathway). Overactivation of the cellular oncogene can result in inappropriate kinase activity, thereby changing many of the cellular activities and leading to cancer. The c-sis oncogene encodes platelet-derived growth factor, which stimulates cells to grow. Its potential in transformation is obvious. GTP-binding proteins, the product of v-ras, for example, play a role in transmitting endocrine signals across membranes. Increased quantities of GTP-binding proteins can send continuous or ampli ed signals to certain cells and thus enhance growth.The v-myc gene product is a protein that binds to
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DNA; speci c transcription factors can signal inappropriate transcription, also inducing transformation. From the initial lesion in a gene to full-blown cancer normally takes many steps (clonal evolution; g. 16.30). For example, in the colorectal cancer familial adenomatous polyposis (FAP), at least seven genetic changes are needed. Through these steps, a normal mass of cells passes through hyperplasia, an increased growth without any obvious change in cells; to dysplasia, in which overgrowth continues with changes in cell and nuclear structures (polyp formation); to the cancerous state, with invasion of surrounding tissues and metastasis. B.Vogelstein and colleagues have discovered many of the genetic changes involved in the formation of this cancer. First, the APC gene (adenomatous polyposis coli) mutates, leading through hyperplasia to dysplasia, a condition referred to as aberrant cryptic foci. Although the exact role of the APC protein is not known, it does bind -catenin, which is involved in cell adhesion and activates the cyclin D1 promoter, exerting a direct effect on cellular proliferation. Thus, mutation of APC results in an accumulation of -catenin, which then has effects on cell cycle progression and cell adhesion. The next genetic change results in an early adenoma (a benign growth). Mutation of the ras oncogene leads to intermediate adenoma, due to the autonomous growth signals sent by the Ras GTP-binding protein. This is followed by late adenoma caused by the mutation of a gene in region q21 of chromosome 18, a gene called deleted in colorectal cancer (DCC). This gene codes for a transmembrane protein involved in the adhesion of cells to each other. At this point, mutations resulting in the loss of p53 protein result in full-blown cancer. Throughout this series of events, it is a cell from the previous state that mutates into the next state, consistent with our concept of clonal evolution. Although it may seem odd that so many mutations appear consecutively in the same cells, remember that mutations in some genes, such as p53, result in an overall higher mutation rate within cells. In one study, when cancer cells were compared with noncancerous progenitor cells, the cancer cells showed eleven thousand genetic changes.
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