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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 Eukaryotic Chromosome
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genome. They rely on the reverse transcriptase provided by the genes of LINES or retroviruses. One group of SINEs not derived from transfer RNA is derived from the RNA of the signal recognition particle (see chapter 11); members of this group occur in human beings in about ve-hundred thousand copies of a three-hundred-basepair sequence. Because these sequences are cleaved by the restriction endonuclease AluI, they are called the Alu family. The human genome is also permeated by remnants of at least a dozen distinct families of ancient retroviruses scattered throughout our chromosomes. At this point, we can see some potential explanations for the C-value paradox. Much eukaryotic DNA is junk, apparently doing no harm. In some cases, 97% of the host genome is composed of junk DNA. Recent work seems to indicate that gross differences in DNA content between higher organisms may be due to the differing abilities of different species to rid themselves of this parasitic DNA. If it builds up without being removed, the DNA content of the species can soar. Thus, the wide differences in DNA content among higher eukaryotes mentioned at the beginning of this section have little to do with the complexity of the organism, but rather with the ability of the organism to remove junk DNA as it forms.
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hybridization studies using probes, similar to the way that telomeric DNA was shown to be at the tips of the chromosomes (see g. 15.26). Repeated genes include the genes for transfer RNAs and histones. The average transfer RNA is repeated about a dozen times in Drosophila. Human beings have over thirteen hundred copies of transfer RNA genes in the haploid genome. In many species, the ve histone genes form a repeated cluster, although each gene is transcribed independently ( g. 15.33), while prokaryotic operons are transcribed as a unit. The arrangement of histone genes may be more complex in higher forms. There are indications that in mammals, histone genes may lie in small groups or even as individual genes. Several types of genes occur in similar but not identical forms that is, an original gene was duplicated but, unlike histone or ribosomal RNA genes, the copies diverged in function. These gene families include globin genes, immunoglobulin genes (see chapter 16), chorion protein (insect eggshell) genes, and Drosophila heat shock genes.
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The Globin Gene Family
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Globins are oxygen-transporting and storage molecules found in animals, some plants, and microorganisms. In higher vertebrates, there are two types of globins: myoglobin, which stores oxygen in muscles, and hemoglobin, found in red blood cells. Myoglobins function as single molecules, whereas hemoglobins occur as tetramers, two each of two protein chains. Evolution in the globin gene family can be traced by comparative studies of globins in different species as well as molecular studies of globins within a species (see chapter 21). Studying hemoglobins has provided a great deal of information on gene expression and evolution. We turn our attention to the globin gene family in human beings. During human development, four major hemoglobins appear: embryonic hemoglobin, Hb F, Hb A, and Hb A2
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Expressed Genes in Many Copies
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Several types of genes create a product that is needed in such large quantity that one copy of the gene could not ful ll the cell s needs. We are familiar with the nucleolus, the site of the ribosomal RNA genes (see g. 10.20). Human beings have about two hundred copies of the major ribosomal RNA gene and about two thousand copies of the 5S ribosomal RNA gene. Fruit ies have about two hundred and one hundred copies, respectively, of the two genes. In some cases, the normal number of multiple copies of a gene is still not enough. The cell must then resort to gene ampli cation, a process whereby the cell increases the number of copies of the gene. For example, during oogenesis, ribosomal RNA genes (rDNA) are often ampli ed. In Xenopus, rDNA is ampli ed about one thousand times, which allows an oocyte to accumulate about 1012 ribosomes. The ampli ed DNA is in the form of small, circular, extrachromosomal molecules of DNA. Several models have been proposed as to how cells actually amplify their DNA. One model relies on unequal crossing over (as in Bar eye in Drosophila), whereas another model is based on unscheduled extra DNA replication in a region, followed by recombinational events that generate linear and circular forms of the excess DNA. It is not presently clear which model is correct. In addition to ribosomal RNA genes, other genes are repeated, ensuring adequate gene products. The number and location of repeated genes are usually discovered by
The arrangement of histone genes (red) within the ve-gene cluster in sea urchins and fruit ies. Arrows indicate the direction of transcription. Spacer DNA (black) separates the genes.
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