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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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Replication in 3H-thymidine
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Second metaphase in hamster cells in culture after one replication in the presence of 3H-thymidine followed by one in nonradioactive medium, verifying the uninemic nature of the eukaryotic chromosome. Cases in which the label apparently switches from one chromatid to the other are caused by sister chromatid exchanges (at arrows). (Source:
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G. Marin and D. M. Prescott, The frequency of sister chromatid exchanges
Replication in unlabeled medium
following exposure to varying doses of 3H-thymidine or X-ray, Journal of Cell Biology, 21, (1964): 159 67, by copyright permission of the Rockefeller University Press.)
Radioactive labeling of a uninemic eukaryotic chromosome following semiconservative replication. Replication occurs rst in the presence of 3H-thymidine and then in its absence. Red represents labeling. After the second round of replication, one chromatid of each chromosome is labeled, whereas the other is not, con rming that there is only one DNA molecule per chromatid and that the chromosome is thus uninemic.
The longest molecule that the investigators found was 1.2 cm long, equivalent to between 24 and 32 109 daltons ( g. 15.3), close to the predicted size. Thus, the evidence is in complete concordance with the simple uninemic model of eukaryotic chromosomal structure (box 15.2).
Nucleoprotein Composition
Nucleosome Structure
Since each eukaryotic chromosome consists of a single, relatively long piece of duplex DNA, the average diploid cell contains many of these long pieces of DNA. For chromosomes to be properly distributed to each daughter cell during mitosis and meiosis, they must be condensed into structures that are more easily managed. Wrapping the DNA around spools of protein constitutes the rst step in a series of coiling and folding processes that eventually result in the fully compacted chromosome we see at metaphase. Interphase nuclei can be disrupted by placing them in a hypotonic liquid such as water. When this happens, chromatin material is released. When this material is observed under the electron microscope, small particles
equivalent change in the viscoelastic estimates to between 52 and 64 109 daltons. The conclusion from these studies is that the largest Drosophila chromosome, and by extension every eukaryotic chromosome, contains a single DNA molecule running from end to end, encompassing both arms. The viscoelastic values were corroborated by carefully isolating and measuring the lengths of long DNA molecules, an especially dif cult task given DNA s propensity to break.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
The Eukaryotic Chromosome
Autoradiograph of a 1.2 cm radioactive DNA molecule carefully isolated from Drosophila melanogaster chromosomes. Drops of DNA solution were placed on microscope slides, then tilted to allow the DNA to spread slowly down the slide. A photographic emulsion was applied and later developed after a ve-month exposure period.
1 mm
(From Ruth Kavenoff, Lynn C. Klotz, and Bruno H. Zimm, Symposia on Quantitative Biology (Cold Spring Harbor), 38(1973):4.)
called nucleosomes can be seen ( g. 15.4). These are the spools that the DNA is wrapped around. They are made of histone proteins and associated DNA (table 15.1). The histones, a group of arginine- and lysine-rich basic proteins, have been well characterized.They are especially well suited to bind to the negatively charged DNA (table 15.2). When chromatin is treated with micrococcal nuclease, individual nucleosomes can be isolated, indicating that the DNA between nucleosomes is accessible to digestion. The results of these studies indicate that a length of 168 base pairs (bp) of DNA, the core DNA, is intimately associated with the nucleosome, and another 50 to 75 base pairs, depending on species, connects the nucleosomes (linker DNA; g. 15.5). When the quantities of the various histones were measured, there were two each of histones H2A, H2B, H3, and H4 per nucleosome and only one molecule of histone H1. Reconstitution and degradation studies have indicated that histone H1 is not a necessary component in the formation of nucleosomes. We believe that histone H1 is associated with the linker DNA as it enters and emerges from the nucleosome ( g. 15.6), although its exact position is not known with certainty. Histone H1 may be more off center and internally located than illustrated. The term chromatosome has been suggested for the core nucleosome plus the H1 protein, a unit that includes approximately 168 base pairs of DNA. Nucleosomes, then, are a rst-order packaging of DNA; they reduce its length and undoubtedly make the coiling and contraction required during mitosis and meiosis more ef cient ( g. 15.7). When DNA is replicated, twice as many nucleosomes are needed since one double helix becomes two. Recent studies indicate that a parental nucleosome is partly disassembled during DNA replication and reassembled on one or the other daughter strand, apparently randomly. The other DNA strand has a new nucleosome constructed
Electron micrograph of chromatin bers. Photo shows nucleosome structures (spheres) and connecting strands of DNA called linkers. The bar is 100 nm long. (Source: D. E.
Olins and A. L. Olins, Nucleosomes: The structural quantum in chromosomes, American Scientist, 66: 704 11, November 1978. Reproduced by permission.)
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