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Although the study of development in animals has progressed markedly by using Drosophila as a model, other organisms have been used as well. Historically, amphibians were the focus of developmental research because they have large eggs that can be easily observed and manipulated. The same reasoning made the chick embryo a classical model of development. The nematode Caenorhabditis elegans has emerged as another model organism for developmental studies because of its simplicity ( g. 16.24). Each individual consists of only about one thousand cells; its life cycle lasts only 3.5 days; and with only 8 107 base pairs of DNA, it has the smallest genome of any multicellular organism. In 1963, S. Brenner proposed learning the lineage of every cell in the adult. With the efforts of numerous colleagues, that work was completed in about twenty years. From the fertilized egg to the adult, the division and fate of every cell of this nematode worm is known. The worm has been especially useful in studying homeotic mutants and
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Sydney Brenner (1927 ).
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(Courtesy of Dr. Sydney Brenner.)
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Figure 16.24 The roundworm Caenorhabditis elegans. (a) Self-fertilizing hermaphrodite. (b) Male. The worms are about 0.3 mm long. (J. E. Sulston and H. R. Horvitz, PostEmbryonic Cell Lineages of the Nematode Caenorhabditis elegans,
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Developmental Biology, 56:1101 56, 1977, Academic Press.)
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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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Sixteen
Gene Expression: Control in Eukaryotes
apoptosis, programmed cell death. As we will see later, apoptosis is important in development as well as in the elimination of infected or cancerous cells. Also being used as animal models are mice and zebra sh. In plants, the snapdragon, Antirrhinum majus, is another model organism. Development is a growth process that is, among other things, an orderly process of cell division. As we discussed in chapter 3, the cell cycle is controlled by checkpoints; the cycle is allowed to continue if the cell is ready for the next stage. Ready means, among other things, that the cell has successfully completed DNA replication and repair of DNA damage. If the cell is not ready, the cell cycle stops until the cell is ready. If the cell is damaged beyond repair, including being cancerous, programmed cell death (apoptosis) is initiated. It is clear that numerous checks and balances are involved in assuring that only healthy, ready cells continue in the cell cycle. Interference to these checks and balances can lead to uncontrolled cell growth cancer.
normal cell to a cancerous one. As we will see, most cancers come about from a series of genetic changes, progressing from an aberrant cell to an aggressively cancerous one. This view is called the clonal evolution theory of cancer. Historically, cancers were understood to be caused by either mutation or by viruses. We now know that viruses can bring cancer-causing genes into cells, where their mutated form or inappropriate location can lead to cancer. Thus, both the mutational and viral views of cancer are ultimately concerned with mutation. In essence, cancers result from the inappropriate activity of certain genes, whether those genes were changed by mutation or were imported or activated by viruses.
Mutational Nature of Cancer
Mutations, both point and chromosomal, have been implicated in carcinogenesis (table 16.3). For example, the disease xeroderma pigmentosum in human beings is caused by mutations in any of seven loci (XpA-XpG) that inactivate the mutation repair system that corrects UVlight damage (see chapter 12); exposure to the sun then results in skin lesions that often become malignant. A related disease, ataxia-telangiectasia, is caused by a defect in the double-strand break repair mechanism, often the result of X-ray induced damage. (Ataxia refers to dif culty in balance; telangiectasia refers to dilated blood vessels in the eye membranes.) By binding to the ends of DNA, as would happen when a double-strand break occurs, ATM (the protein product of the ataxiatelangiectasia locus, atm) begins a signaling pathway that tells the cell there are broken ends of DNA. Persons with this defect are at risk for acute and chronic leukemia and lymphomas; women with it are also at risk for ovarian cancers. Most cancers are associated with chromosomal defects; improved chromosomal banding techniques have demonstrated that a speci c chromosomal defect is often associated with a speci c cancer (table 16.3, g. 16.25, box 16.2). The implication is that when a gene is in a new location (because of translocation or the deletion of intervening material), that gene may fall under the control of more powerful promoters or promoters outside the range of that gene s normal control. As we shall see, genes that are known to be able to transform cells (oncogenes) are often the ones that are relocated into regions of new control. These oncogenes then become more active, and transformation follows.
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