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Tamarin: Principles of Genetics, Seventh Edition
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II. Mendelism and the Chromosomal Theory
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6. Linkage and Mapping in Eukaryotes
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The McGraw Hill Companies, 2001
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Linkage and Mapping in Eukaryotes
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Black, purple, curved b b pr pr c c
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Wild-type b + b + pr + pr + c + c +
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Testcross the trihybrid
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Wild-type (trihybrid) b + b pr + pr c +c
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Black, purple, curved b b pr pr c c
If unlinked 1/8 b/b pr /pr c /c 1/8 b/b pr /pr c +/c 1/8 b/b pr +/ pr c /c 1/8 b/b pr +/ pr c +/c 1/8 b +/b pr /pr c /c 1/8 b +/b pr /pr c +/ c 1/8 b +/b pr +/ pr c /c 1/8 b +/b pr +/ pr c +/ c Figure 6.6
If completely linked 1/2 b pr c /b pr c 1/2 b + pr + c + /b pr c
Possible results in the testcross progeny of the b pr c trihybrid.
pr and c ( g. 6.8). Groupings according to these recombinant events are shown at the right in table 6.1. In the nal column of table 6.1, recombination between b and c is scored. Only those recombinant classes that have a new arrangement of b and c alleles, as compared with the parentals, are counted. This last column shows us what a b c, two-point cross would have revealed had we been unaware of the pr locus in the middle.
Map Distances
The percent row in table 6.1 reveals that 5.9% (887/15,000) of the offspring in the Drosophila trihybrid
testcross resulted from recombination between b and pr, 19.5% between pr and c, and 23.7% between b and c. These numbers allow us to form a tentative map of the loci ( g. 6.9). There is, however, a discrepancy. The distance between b and c can be calculated in two ways. By adding the two distances, b pr and pr c, we get 5.9 19.5 25.4 map units; yet by directly counting the recombinants (the last column of table 6.1), we get a distance of only 23.7 map units. What causes this discrepancy of 1.7 map units Returning to the last column of table 6.1, we observe that the double crossovers (60 and 72) are not counted, yet each actually represents two crossovers in this re-
Table 6.1 Results of Testcrossing Female Drosophila Heterozygous for Black Body Color,
Purple Eye Color, and Curved Wings (b b pr pr c c bb prpr cc)
Number Recombinant Between b and pr pr and c b and c
Phenotype Wild-type Black, purple, curved Purple, curved Black Curved Black, purple Purple Black, curved Total Percent
Genotype b b pr pr c c bb prpr cc b b prpr cc bb pr pr c c b b pr pr cc bb prpr c c b b prpr c c bb pr pr cc
Number 5,701 5,617 388 367 1,412 1,383 60 72 15,000
Alleles from Trihybrid Female b pr c b pr c b pr c b pr c b pr c b pr c b pr c b pr c
388 367 1,412 1,383 60 72 887 5.9 60 72 2,927 19.5
388 367 1,412 1,383
3,550 23.7
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
6. Linkage and Mapping in Eukaryotes
The McGraw Hill Companies, 2001
Diploid Mapping
Meiotic tetrad
Gametes
b b b+ b+
pr pr pr + pr +
c c c+ c+
Gametes
b b b+ b+
pr pr pr pr +
Nonrecombinant
c+ c c+
Recombinant Nonrecombinant
Results of a crossover between the black and purple loci in Drosophila.
Meiotic tetrad
Gametes
b b b+ b+
pr pr pr + pr +
c c c+ c+
Gametes
b b b+ b+
pr pr + pr pr +
Nonrecombinant
c c+ c+
Double recombinant Nonrecombinant
Results of a double crossover in the b pr c region of the Drosophila chromosome.
gion.The reason they are not counted is simply that if we observed only the end loci of this chromosomal segment, we would not detect the double crossovers; the rst one of the two crossovers causes a recombination between the two end loci, whereas the second one returns these outer loci to their original con guration (see g. 6.8). If we took the 3,550 recombinants between b and c and added in twice the total of the double recombinants, 264, we would get a total of 3,814. This is 25.4 map units, which is the more precise gure we calculated before. The farther two loci are apart on a chromosome, the more double crossovers occur between them. Double crossovers tend to mask recombinants, as in our example, so that distantly linked loci usually appear closer than they really are. Thus, the most accurate map dis-
tances are those established on very closely linked loci. In other words, summed short distances are more accurate than directly measured larger distances. The results of the previous experiment show that we can obtain at least two map distances between any two loci: measured and actual. Measured map distance between two loci is the value obtained from a two-point cross. Actual map distance is an idealized, more accurate value obtained from summing short distances between many intervening loci. We obtain the short distances from crosses involving other loci between the original two.When we plot measured map distance against actual map distance, we obtain the curve in gure 6.10. This curve is called a mapping function. This graph is of both practical and theoretical value. Pragmatically, it allows
25.4 (sum of two shorter distances; best estimate of true map distance)
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