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Diploid Mapping 110 Two-Point Cross 110 Three-Point Cross 114 Cytological Demonstration of Crossing Over 120 Haploid Mapping (Tetrad Analysis) 122 Phenotypes of Fungi 124 Unordered Spores (Yeast) 124 Ordered Spores (Neurospora) 125 Somatic (Mitotic) Crossing Over 132 Human Chromosomal Maps 132 X Linkage 132 Autosomal Linkage 134 Summary 140 Solved Problems 140 Exercises and Problems 142 Critical Thinking Questions 147 Box 6.1 The Nobel Prize 112 Box 6.2 The First Chromosomal Map 121 Box 6.3 Lod Scores 135
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Scanning electron micrograph (false color) of a fruit y, Drosophila melanogaster.
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II. Mendelism and the Chromosomal Theory
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6. Linkage and Mapping in Eukaryotes
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fter Sutton suggested the chromosomal theory of inheritance in 1903, evidence accumulated that genes were located on chromosomes. For example, Morgan showed by an analysis of inheritance patterns that the white-eye locus in Drosophila is located on the X chromosome. Given that any organism has many more genes than chromosomes, it follows that each chromosome has many loci. Since chromosomes in eukaryotes are linear, it also follows that genes are arranged in a linear fashion on chromosomes, like beads on a string. Sturtevant rst demonstrated this in 1913. In this chapter, we look at analytical techniques for mapping chromosomes techniques for determining the relationship between different genes on the same chromosome. These techniques are powerful tools that allow us to nd out about the physical relationships of genes on chromosomes without ever having to see a gene or a chromosome. We determine that genes are on the same chromosome when the genes fail to undergo independent assortment, and then we use recombination frequencies to determine the distance between genes. If loci were locked together permanently on a chromosome, allelic combinations would always be the same. However, at meiosis, crossing over allows the alleles of associated loci to show some measure of independence. A geneticist can use crossing over between loci to determine how close one locus actually is to another on a chromosome and thus to map an entire chromosome and eventually the entire genome (genetic complement) of an organism. Loci carried on the same chromosome are said to be linked to each other. There are as many linkage groups (l) as there are autosomes in the haploid set plus sex chromosomes. Drosophila has ve linkage groups (2n 8; l 3 autosomes X Y), whereas human beings have twenty-four linkage groups (2n 46; l 22 autosomes X Y). Prokaryotes and viruses, which usually have a single chromosome, are discussed in chapter 7. Historically, classical mapping techniques, as described in this chapter and the next, gave researchers their only tools to determine the relationships of particular genes and their chromosomes. When geneticists know the locations of speci c genes, they can study them in relation to each other and begin to develop a comprehensive catalogue of the genome of an organism. Knowing the location of a gene also helps in isolating the gene and studying its function and structure. And mapping the genes of different types of organisms (diploid, haploid, eukaryotic, prokaryotic) gives geneticists insight into genetic processes. More recently, recombinant DNA technology has allowed researchers to sequence whole genomes, including the human and fruit y genomes; this means they now know the exact locations of all the genes on all the chromosomes of these organisms (see
chapter 13). Geneticists are now creating massive databases containing this information, much of it available for free or by subscription on the World Wide Web. Until investigators mine all this information for all organisms of interest, they will still use analytical techniques in the laboratory and eld to locate genes on chromosomes.
DIPLOID MAPPING
Two-Point Cross
In Drosophila, the recessive band gene (bn) causes a dark transverse band on the thorax, and the detached gene (det) causes the crossveins of the wings to be either detached or absent ( g. 6.1). A banded y was crossed with a detached y to produce wild-type, dihybrid offspring in the F1 generation. F1 females were then testcrossed to banded, detached males ( g. 6.2). (There is no crossing over in male fruit ies; in experiments designed to detect linkage, heterozygous females in which crossing over occurs are usually crossed with homozygous recessive males.) If the loci were assorting independently, we would expect a 1:1:1:1 ratio of the four possible phenotypes. However, of the rst one thousand offspring examined, experimenters recorded a ratio of 2:483: 512:3. Several points emerge from the data in gure 6.2. First, no simple ratio is apparent. If we divide by two, we get a ratio of 1:241:256:1.5. Although the rst and last categories seem about equal, as do the middle two, no simple numerical relation seems to exist between the middle and end categories. Second, the two cate-
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