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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 McGraw Hill Companies, 2001
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The Eukaryotic Chromosome
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R-bands are visible with a technique that stains the regions between G-bands. The chromosomes are xed, stained with Giemsa, and then viewed with a phase contrast microscope. Since the dark-light pattern is the opposite of the G-band pattern, these bands are called reverse bands.
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From the information gleaned from these staining techniques, D. Comings distinguished between three basic chromatin types: euchromatin, constitutive heterochromatin, and intercalary heterochromatin (table 15.3). Presumably, the only chromatin involved in transcription is euchromatin. Constitutive heterochromatin surrounds the centromere and is rich in
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David E. Comings (1935 ).
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(Courtesy Dr. David E. Comings.)
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(a) C banding of chromosomes from a cell in the bone marrow of the house mouse, Mus musculus. The arrow indicates that the Y chromatids have already separated into two chromosomes. (b) Yellow uorescence indicates a satellite DNA probe in human chromosomes (centromeres). ([a] B. Vig,
Figure 15.21 Satellite DNA in Drosophila virilis. The quantity of DNA is graphed against the buoyant density (g/cc), resulting in four peaks. The large peak (at left) is the major DNA component of the cell; the other three bands are satellite DNA. The left-most of the satellite peaks (1.692) is DNA with a repeating sequence of ACAAACT; the middle satellite peak (1.688) is a sequence of ATAAACT; and the right-most satellite peak (1.671) has a sequence of ACAAATT. (From Joseph G. Gall,
et al., Cold Spring Harbor Laboratory Symposia on Quantitative Biology, 38:417 21. Copyright 1974 Cold Spring Harbor Laboratory, Cold Spring Harbor, NY. Reprinted by permission.)
Sequence of centromere separation: Role of centromeric heterochromatin, Genetics, 102:795 806, 1982. [b] Photograph Courtesy of Oncor, Inc. Gaithersburg, Maryland.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
The Eukaryotic Chromosome
Table 15.3 The Three Major Types of Chromatin in Eukaryotic Chromosomes
Euchromatin Relation to bands Location Condition during interphase Genetic activity Relation to chromomeres In R-bands Chromosome arms Usually dispersed Usually active Interchromomeric Centromeric Constitutive Heterochromatin In C-bands Usually centromeric Condensed Inactive Centromeric chromomere Intercalary Heterochromatin In G-bands Chromosome arms Condensed Probably inactive Intercalary chromomeres
satellite DNA. Intercalary heterochromatin is found throughout the chromosome. Thus, it becomes apparent that the eukaryotic chromosome is a relatively complex structure.
ML IL OL
Centromeres and Telomeres
Centromeres
Two regions of the eukaryotic chromosome have speci c functions the centromere and the telomeres. The centromere is involved in chromosomal movement during mitosis and meiosis, whereas the telomeres terminate the chromosomes. As we pointed out in chapter 3, the terms centromere and kinetochore, while occasionally used interchangeably, are distinct. The kinetochore is the interface between the visible constriction in the chromosome (the centromere) and the microtubules of the spindle. The kinetochore of higher organisms (e.g., mammals) contains proteins and some RNA. Microscopically, it is a trilaminar structure, attached to chromatin at the inner layer and to microtubules at the outer layer ( g. 15.22). Most of our knowledge of the genetics of centromeres has come from work in yeast (Saccharomyces cerevisiae). Cells did not maintain most arti cially created yeast plasmids because they were lost during mitosis. However, plasmids were isolated that did replicate normally during cell division. Presumably, they contained centromeres, allowing them to replicate and move in synchrony with the host s chromosomes. Further genetic engineering made it possible to isolate smaller and smaller regions that could serve as centromeres. After sequencing the centromeres of fteen of the sixteen yeast chromosomes, it was possible to conclude that the centromere from yeast is about 250 base pairs long with three consensus regions ( g. 15.23); we are de ning a centromere as a sequence of DNA called the CEN locus or CEN region. Recent data indicate that this region may contain a single, modi ed nucleosome associated with
The kinetochore of a metaphase chromosome of the rat kangaroo. IL, ML, and OL refer to inner, middle, and outer layers, respectively, of the kinetochore. Note the microtubules attached to the kinetochore and the large mass of dark-staining chromatin making up most of the gure. Magni cation 30,800 . (From B. R. Brinkley and J. Cartwright, Jr.,
J. Cell Biology, 50:416 31, 1971.)
region II. The 250 base-pair length of the CEN regions of yeast chromosomes is about 200 , the same as the diameter of a microtubule, indicating that only one microtubule attaches to each centromere during mitosis or meiosis in a yeast cell. This region is called a point centromere ( g. 15.24). Higher eukaryotes have larger centromeric regions that attach more microtubules. These regions are referred to as regional centromeres (see gs. 15.22 and also 3.12). Regional centromeres range from nineteen to one hundred kilobases (kb; 19,000 100,000 bases) with unique and satellite (repeated sequence) DNA that is heterochromatic and may include expressed genes. We know much less about regional centromeres than we do about point centromeres.
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