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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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as molting. Tissue-speci c puffs are active in one tissue but not another. (In dipteran larvae, tissues other than the salivary glands, such as the midgut and Malpighian tubules, have polytene chromosomes.) Constitutive puffs are active almost all the time in a speci c tissue. And environmentally induced puffs appear after some environmental change, such as heat shock ( g. 15.16). In Drosophila, about 80% of the puffs are stage speci c; in Chironomus, only about 20% are. For example, at the time of molt in insects, the hormone ecdysone is secreted by the prothoracic gland. At the same time, many puff patterns change ( g. 15.17). Similar changes in puff patterns can be induced by the injection of ecdysone. Hence, molting, a stage-speci c developmental sequence, is related to a sequential transcription sequence in the chromosomes.
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Lampbrush Chromosomes
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Lampbrush chromosomes, which occur in amphibian oocytes, are so named because their looped-out con guration has the appearance of a brush for cleaning lamps, now a relatively uncommon household item ( g. 15.18). The loops of the lampbrush chromosomes are covered by an RNA matrix and are the sites of active transcription. Presumably, the loops are unwindings of the single chromosome, similar to the unwindings in the polytene chromosome shown in gure 15.14. Thus, under certain circumstances, such as in polytene chromosomal puffs
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Figure 15.16 Puff 4-81B of the salivary gland in Drosophila hydei is induced by heat shock (37 C for one-half hour). At the top, normal activity. At the bottom, temperature shock in vitro, resulting in the puff. (Source: H. D. Berendes, et al.,
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Experimental puffs in salivary gland chromosomes of Drosophila hydei, Chromosoma [Berl.] 16:35 46, Fig. 4a b, 1965. Springer-Verlag.)
Time
Hybridization at a Chironomus tentans salivary gland chromosome puff. The chromosomal DNA is hybridized with labeled RNA (black dots) transcribed from the locus. The activity of the locus is forming the puff. (Reprinted by permission
Figure 15.17 Puff patterns on a segment of a Chironomus tentans salivary gland chromosome during molt. As time proceeds, puffs appear and disappear and change in size.
from B. Lambert, Repeated DNA sequences in a Balbiani ring, Journal of Molecular Biology, 72:65 75, 1972. Copyright by Academic Press, Inc. (London) Ltd.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
The Eukaryotic Chromosome
Lampbrush chromosome of the newt, Notophthalmus viridescens. Centromere is at the the left (arrow); the two long homologues are held together by three chiasmata. Magni cation 238 . (Source: Joseph G. Gall, gure 2 in
D. M. Prescott, ed., Methods in Cell Physiology, vol. 2 [New York: Academic Press, 1966], 39. Reproduced by permission.)
and in lampbrush chromosomes, active transcription can be seen in the light microscope. Since only certain bands puff at any one moment in polytene chromosomes, and since the loops of lampbrush chromosomes are of various sizes (with some regions not looped at all), we have evidence of speci c transcription. However, we have no indication, so far, of the nature of the control of that transcription.
Chromosomal Banding
Several chromosomal staining techniques reveal consistent banding patterns. By means of these patterns, all of the human chromosomes can be differentiated (see g. 5.1). Of possibly greater importance is the fact that these staining techniques have provided some insight into the structure of the chromosome. The techniques for staining the C, G, and R chromosomal bands will serve as an illustration. G-bands are obtained with Giemsa stain, a complex of stains speci c for the phosphate groups of DNA. Treatment of xed chromatin with trypsin or hot salts brings out the G-bands. Giemsa stain enhances banding that is already visible in mitotic chromosomes. The banding pattern is caused by the arrangement of chromomeres. Under careful observation, the major G-bands prove to consist of many smaller chromomeres. This banding appearance has led D. Comings to suggest the mechanism of chromosomal folding shown in gure 15.19. C-bands are Giemsa-stained bands after the chromosomes are treated with NaOH. The C is for centromere, because these bands represent constitutive heterochro-
Figure 15.19 Model of eukaryotic (mammalian) chromosomal banding. G-bands are chromomere clusters, which result from the contraction of smaller chromomeres. These, in turn, result from looping of the 300 ber. (Reproduced with permission, from
the Annual Review of Genetics, Volume 12, 1978 by Annual Reviews, Inc.)
matin surrounding the centromeres ( g. 15.20).The DNA is also usually satellite rich. Satellite DNA differs in buoyant density from the major portion of cellular DNA. When eukaryotic DNA is isolated and centrifuged in CsCl, forming a density gradient, the majority of the DNA forms one band in the gradient at a single buoyant density. The buoyancy is determined by the G-C content of the DNA. However, smaller secondary bands are also usually present, indicating regions of DNA having sequences different from the majority of the cell s DNA ( g. 15.21). DNA isolated this way is referred to as satellite DNA because of the secondary, or satellite, bands formed in the density gradient. As we will see, this DNA is found primarily around centromeres and consists of numerous repetitions of a short sequence.
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