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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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15. The Eukaryotic Chromosome
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The Eukaryotic Chromosome
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n chapter 14, we looked at the control of gene expression in prokaryotes and bacteriophages. Compared to eukaryotes, bacteriophages and prokaryotes are relatively simple. Of fundamental importance is that, in these lower forms, the operon model of induction and repression of transcription is a unifying theme for control of gene expression. Despite nuances such as catabolite repression and attenuator control, the operon model provides a relatively clear picture of how genes are turned on and off in phages and prokaryotes.This model does not exist for eukaryotes. In attempting to elucidate models for control of gene expression in eukaryotes, we must take one very important factor into account: the complexity of the structure of the eukaryotic chromosome. In this chapter, we cover the current understanding of how these very large structures are organized.
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other limited forms of differentiation, they are primarily one-celled organisms. And, although some eukaryotes are single-celled (e.g., yeast), the essence of eukaryotes is differentiation. In human beings, a zygote gives rise to every other cell type in the body in a relatively predictable manner. To fully appreciate the complexity of eukaryotes, we begin by looking at the eukaryotic chromosome. In the next chapter, we look at the patterns of development in eukaryotes and some mechanisms of control of gene expression.
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THE EUKARYOTIC CHROMOSOME
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Evidence that the eukaryotic chromosome is uninemic that is, contains one double helix of DNA comes from several sources.The best data are provided by radioactivelabeling studies, rst done by J. Taylor and his colleagues in 1957. If a eukaryote is allowed to undergo one DNA replication in the presence of tritiated (3H-) thymidine, each of the daughter chromatids would be expected to contain a double helix with one unlabeled DNA template strand and one labeled strand of newly synthesized bases ( g. 15.1). This con guration is expected on the basis of semiconservative replication, with each chromatid containing one double helix. A second round of DNA replication, in the absence of 3H-thymidine, should produce chromosomes in which one chromatid would have unlabeled DNA and one would have labeled DNA. Figure 15.2 shows the chromosomes after this second replication in nonlabeled media. As expected, one chromatid of every pair is labeled and one is not. In another kind of experiment, R. Kavenoff, L. Klotz, and B. Zimm demonstrated that Drosophila nuclei contained pieces of DNA of the size predicted from their DNA content, based on the premise that each chromosome contains one DNA molecule. They isolated the DNA and measured the size of the largest DNA molecules using the viscoelastic property of DNA, the rate at which
Ruth Kavenoff (1944 ).
(Courtesy of Dr. Ruth Kavenoff.)
THE EUKARYOTIC CELL
Eukaryotes and prokaryotes are the two superkingdoms of organisms. The following comparisons, using E. coli as a general model for prokaryotes, show how much more complex eukaryotes are: 1. An E. coli chromosome contains approximately 4.2 106 base pairs of DNA. The haploid human genome contains nearly one thousand times as much DNA. 2. Eukaryotic DNA is in the form of nucleoprotein, a DNA-histone protein complex. Although a few histonelike proteins have been found in E. coli, its chromosomal DNA is not complexed with protein to anywhere near the same extent. 3. An E. coli cell has very little internal structure. Eukaryotes have a number of internal organelles and an extensive lipid membrane system, including the nuclear envelope itself. 4. An E. coli cell is small (0.5 to 5.0 m in length for bacteria). Eukaryotic cells are generally larger than prokaryotes (10 to 50 m in length for animal tissue cells; box 15.1). 5. The messenger RNA of E. coli is translated while it is being transcribed. Eukaryotic messenger RNA is modi ed within the nucleus before it is transported out for translation in the cytoplasm. 6. Almost no messenger RNA isolated from eukaryotic cells, including the messenger RNA of animal viruses, has been found to be polycistronic (containing many genes). Most prokaryotic messenger RNAs are polycistronic. 7. Most E. coli genes are parts of inducible or repressible operons; there are almost no operons in eukaryotes. 8. E. coli exists as a simple, single cell. Although some prokaryotes do aggregate, sporulate, and show a few
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