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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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(table 15.5). Structurally, the (Greek, zeta) subunit (a component of embryonic hemoglobin) is -like, whereas the rest are -like ( g. 15.34; see also g. 10.29). Fetal hemoglobin has a higher af nity for oxygen than does adult hemoglobin, thus allowing fetuses to draw oxygen from their mother s blood. From a comparative study of the DNA sequences, the evolution of the various hemoglobin genes has been inferred ( g. 15.35). The genes are located in a cluster on chromosome 16; the genes are located in a cluster on chromosome 11 ( g. 15.36). These two clusters provide a clear case history of gene duplication, presumably by unequal crossing
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over, followed by divergence. Having a second or third copy of a gene allows one of the duplicates to diverge (and perhaps to become nonfunctional in the process), whereas the original still performs the required function. Many diseases of genetic interest involve the hemoglobins. In fact, hemoglobinopathies, including sicklecell anemia and the thalassemias, are the most common genetic disorders in the world population. The bestknown mutation of a hemoglobin gene itself is the one that causes sickle-cell anemia, a mutation of the sixth amino acid of the chain. In the homozygous state, the disease is usually fatal. However, heterozygotes show an
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Figure 15.34 The structure of adult human and -globin genes. The numbers refer to amino acids (or translated codons).
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Figure 15.35 The presumed evolution of the various human globin genes from an ancestral primitive gene. The diagram represents a branching tree that begins on the left and progresses to the right. Each branch point is an evolutionary step in which the genes presumably were duplicated and then either diverged or simply endured as duplicates, as in present-day genes (on the right).
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Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
15. The Eukaryotic Chromosome
The McGraw Hill Companies, 2001
Summary
Table 15.5 Types of Human Hemoglobin
Type Embryonic Fetal (Hb F) Adult (Hb A) Adult (Hb A2) The - and -globin gene clusters in human beings. The 1 and 2 and the 1 refer to nontranscribing genes (pseudogenes). Mutation has rendered the pseudogenes inactive. Within each gene box, solid color refers to exons and open regions refer to introns. (Reproduced, with permission, from the
Annual Review of Genetics, Volume 14, 1980 by Annual Reviews, Inc.)
Generally When Present Up until eight weeks of gestation and beyond Eight weeks to birth Just before birth and beyond In immature cells
Composition
2 2 2 2
Note: Subscripts refer to the numbers of subunits present.
Tom Maniatis (1943 ).
(Courtesy of Dr. Tom Maniatis.)
increased resistance to malaria. One of the rami cations is that the sickle-cell allele is maintained at relatively high frequencies in malarial regions (see chapter 21). The thalassemias are a group of diseases that affect the regulation of the and hemoglobin genes. (Thalassemia comes from the Greek for sea blood, because the disease is best known in individuals living around the Mediterranean Sea.) In and thalassemias, the or subunit, respectively, is present in very low quantities or entirely absent. Many of the genetic defects are deletions, possibly due to unequal crossing over within the globin gene complexes. T. Maniatis showed that thalassemia is caused by a mutation in the -globin gene that disrupts RNA splicing. The body compensates by forming 4 or 4
hemoglobin in thalassemias, or 2 2 or 2 2 in thalassemias. These are relatively unsuitable or inef cient responses; the diseases range from very mild to very severe and frequently fatal. More information is needed regarding the control of hemoglobin production in the thalassemias.
S U M M A R Y
STUDY OBJECTIVE 1: To examine the arrangement of
DNA and proteins compromising the eukarayotic chromosome 440 452
To study developmental control in eukaryotes, we must understand the eukaryotic chromosome, which is uninemic: It consists of one DNA double helix per chromosome. Nucleoprotein is composed of DNA, histones, and nonhistone proteins. The nucleosome, a uniform packaging of the DNA, is made of histones. The majority of the nonhistone proteins create the scaffold structure of the chromosome and are not involved in gene regulation. Presumably, very small quantities of the nonhistone proteins take part in the regulation of transcription.
Core DNA, wrapped around nucleosomes, is separated by linker DNA between nucleosomes. There are regions of DNA, vulnerable to nucleases, that do not contain nucleosomes; these are referred to as nuclease-hypersensitive sites. Nucleosomes generally inhibit transcription. The 110 nucleosomed DNA forms a 300 ber by coiling into a solenoidlike structure. Coiling of this ber presumably forms the thick, 2,400 ber seen in metaphase chromosomes. STUDY OBJECTIVE 2: To look at the nature of centromeres
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