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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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Higher-Order Structure of Chromatin
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Since the nucleosome has a width of only 110 , and metaphase chromosomes appear to be constructed of a ber having a diameter of about 2,400 ( g. 15.9), several additional levels of chromatin compaction lead to the metaphase chromosome.Various experiments, which change the ionic strength the chromatin is subjected to, indicate that the 110 DNA spontaneously forms a 300 , solenoidlike ber with increased ionic strength. It seems that this ber results from the coiling of the nucleosomal DNA ( g. 15.10). This 300 ber is not, however, the nal form of the DNA. We can account for the contraction of the 300 ber to the 2,400 ber found in metaphase chromosomes by the formation of a second solenoidlike structure from the winding of the 300 ber ( g. 15.11). If the histones are removed from a chromosome, the DNA billows out, leaving a proteinaceous structure termed a scaffold ( g. 15.12). This scaffold structure is formed from nonhistone proteins; two of them predominate, namely SC1 and SC2. SC1 has been identi ed
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as topoisomerase II. It would not be unreasonable to expect several hundred different proteins, many in minute quantities, to be associated with the chromosome and involved in replication, repair, and transcription.
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Figure 15.10 Solenoid model for the formation of the 300 chromatin ber. Nucleosomal DNA wraps in a helical fashion, forming a hollow core. Although histone H1 is not shown, it is known to be on the inside of the solenoid.
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2,400
300
Chinese hamster chromosome. Note the bers making up the chromosome; they are approximately 2,400 in diameter. Magni cation 11,800 . (Source: Courtesy of Dr. Hans Ris.)
Figure 15.11 The 2,400 ber of the eukaryotic chromosome is a hollow, solenoidlike structure. It is formed by the coiling of the 300 ber, which itself is a solenoid.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
The Eukaryotic Chromosome
Figure 15.13 A chromosome puff on the left arm of chromosome 3 of the midge Chironomus pallidivittatus.
Academic Press.)
(Jan-
Erik Edstr m, et al., Developmental Biology 91:131 37, 1982, Figure 1B,
Band
Band
Scaffold protein. When the histones are removed from a eukaryotic chromosome, a brous scaffold remains. The DNA loops out from this scaffold. The bar is 2 m long.
Puff
Puff
(J. Paulson and U. Laemmli, The structure of histone-depleted metaphase chromosomes, Cell, 12:817 28, 1977. Micrograph courtesy of James R. Paulson.)
Band Interband 300 A fibers
Band Interband
Polyteny, Puffs, and Balbiani Rings
Drosophila s salivary glands, as well as some other tissues of Drosophila and other diptera, contain giant banded chromosomes (see g. 6.12) that result from the replication of the chromosomes and the synapsis of homologues without cell division (endomitosis). These chromosomes consist of more than one thousand copies of the same chromatid and appear as alternating dark bands and lighter interband regions. The dark bands are referred to as chromomeres. Also seen are diffuse areas called chromosome puffs ( g. 15.13). Chromosome puffs are also referred to as Balbiani rings. These rings were originally de ned as puffs in the midge, Chironomus, whose polytene chromosomes were discovered by E. G. Balbiani in 1881. Currently, the term applies to all puffs, or at least the larger puffs, in all species with polytene chromosomes. The structure of the polytene chromosome can be explained by the diagram in gure 15.14. Dark bands (chromomeres) are due to tight coiling of the 300 ber; light interband regions are due to looser coiling. The gure
Figure 15.14 Polytene chromosome with bands and a puff. Three of the approximately one thousand synapsed chromatids are shown diagrammatically on the right.
also shows how chromosome puffs would come about as bers unfold in regions of active transcription. Staining with reagents speci c for RNA, such as toluidine blue, or autoradiography with tritiated (3H) uridine, have been used to demonstrate that active transcription is going on in the puffs but not in neighboring regions of the polytene chromosomes. The messenger RNA isolated from cells with puffs has also been shown to hybridize only to the puffed regions of the chromosomes. Thus, these regions of the DNA are complementary to the messenger RNA ( g. 15.15) and represent areas of active transcription. Modern recombinant DNA techniques have also shown that many puffs probably represent the transcription of only one gene, although there are exceptions. Puffs generally fall into four categories. Stage-speci c puffs appear during a certain stage of development, such
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