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Figure 6.24 Two possible arrangements of the a and b loci and their centromere. Distances are in map units.
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Tamarin: Principles of Genetics, Seventh Edition
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The McGraw Hill Companies, 2001
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tetratypes plus the number of nonparental ditypes, all divided by the total number of asci, expressed as a percentage. For ordered spores (Neurospora), the distance from a locus to its centromere is one-half the percentage of second-division segregants. Mapping the distance between two loci is similar to the process in unordered spores.
Yellow spot Singed spot
S O M AT I C ( M I T O T I C ) CROSSING OVER
Crossing over is known to occur in somatic cells as well as during meiosis. It apparently occurs when two homologous chromatids come to lie next to each other and breakage and reunion follow, most likely as a consequence of DNA repair (see chapter 12). Unlike in meiosis, no synaptonemal complex forms. The occurrence of mitotic crossing over is relatively rare. In the fungus Aspergillus nidulans, mitotic crossing over occurs about once in every one hundred cell divisions. Mitotic recombination was discovered in 1936 by Curt Stern, who noticed the occurrence of twin spots in fruit ies that were dihybrid for the yellow allele for body color (y) and the singed allele (sn) for bristle morphology ( g. 6.25). A twin spot could be explained by mitotic crossing over between the sn locus and its centromere ( g. 6.26). A crossover in the sn y region would produce only a yellow spot, whereas a double crossover, one between y and sn and the other between sn and the centromere, would produce only a singed spot. (Verify this for yourself.) These three phenotypes were found in the relative frequencies expected. That is, given that the gene locations are drawn to scale in gure 6.26, we would expect double spots to be most common, followed by yellow spots, with singed spots rarest of all because they require a double crossover. This in fact occurred, and no other obvious explanation was consistent with these facts. Mitotic crossing over has been used in fungal genetics as a supplemental, or even a primary, method for determining linkage relations. Although gene orders are consistent between mitotic and meiotic mapping, relative distances are usually not, which is not totally unexpected. We know that neither meiotic nor mitotic crossing over is uniform along a chromosome. Apparently, the factors that cause deviation from uniformity differ in the two processes.
Yellow and singed twin spots on the thorax of a female Drosophila.
Curt Stern (1902 1981)
(Courtesy of the Science Council of Japan.)
ci c crosses coupled with the relatively small number of offspring) make these techniques of human chromosome mapping very dif cult. However, some progress has been made based on pedigrees, especially in assigning genes to the X chromosome. As the pedigree analysis in the previous chapter has shown, X chromosomal traits have unique patterns of inheritance, and loci on the X chromosome are easy to identify. Currently over four hundred loci are known to be on the X chromosome. It has been estimated, by several different methods, that between fty and one hundred thousand loci exist on human chromosomes. In later chapters, we will discuss several additional methods of human chromosomal mapping that use molecular genetic techniques.
HUMAN CHROMOSOMAL MAPS
In theory, we can map human chromosomes as we would those of any other organism. Realistically, the problems mentioned earlier (the inability to make spe-
X Linkage
After determining that a human gene is X linked, the next problem is to determine the position of the locus on the X chromosome and the map units between loci. Sometimes we can do this with the proper pedigrees, if crossing over can be ascertained. An example of this grand-
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
6. Linkage and Mapping in Eukaryotes
The McGraw Hill Companies, 2001
Human Chromosomal Maps
sn+ sn+ sn sn
y y y+ y+
Dihybrid at prophase (mitosis)
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