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We can now determine the distance from each locus to its centromere and the linkage arrangement of the loci if they are both linked to the same centromere. We can establish by inspection that the two loci are linked to each
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Table 6.5 Genetic Patterns Following Meiosis in an a a Heterozygous Neurospora (Ten Asci Examined)
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Ascus Number Spore Number 1 2 3 4 5 6 7 8 1 a a a a a a a a FDS 2 a a a a a a a a FDS 3 a a a a a a a a FDS
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(1/2)% SDS (1/2) 40% 20 map units
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4 a a a a a a a a SDS
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9 a a a a a a a a SDS
10 a a a a a a a a FDS
Note: Map distance (a locus to centromere)
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
6. Linkage and Mapping in Eukaryotes
The McGraw Hill Companies, 2001
Six
Linkage and Mapping in Eukaryotes
First-division segregation with no crossover between a locus and centromere
Table 6.6 Eight of the Thirty-Six Possible Spore
Patterns in Neurospora Scored for Two Loci, a and b (All Random Variants of the Same Genetic Event)
Ascus Number Spore Number 1 ab ab ab ab a b a b a b a b 2 ab ab ab ab a b a b a b a b 3 ab ab ab ab a b a b a b a b 4 ab ab ab ab a b a b a b a b 5 a b a b a b a b ab ab ab ab 6 a b a b a b a b ab ab ab ab 7 a b a b a b a b ab ab ab ab 8 a b a b a b a b ab ab ab ab
a+ a+
a+ a+ a+ a+
a+ a a+
1 2 3 4
a a a a
a a a a a+ a+ a+ a+
5 6 7
a+ a+ a
Second-division segregation with crossover between a locus and centromere
Table 6.7 The Seven Unique Classes of Asci
a+ a a+ a
Resulting from Meiosis in a Dihybrid Neurospora, ab/a b
Ascus Number
a+ a+ a+
a a+ a a a+
a a+ a a
2:2:2:2
Spore Number 1 2
2:2:2:2
1 ab ab ab ab a b a b a b a b 729
2 ab ab ab ab a b a b a b a b 2
3 ab ab ab ab a b a b a b a b 101 101
4 ab ab a b a b a b a b ab ab 9 9
5 ab ab a b a b a b a b ab ab 150 150 150 PD
6 ab ab a b a b a b a b ab ab 1 1 1 NPD
7 ab ab a b a b a b a b ab ab 8 8 8 TT
a a a+ a+ a a a+ a+
3 4 5 6 7 8 SDS for a locus SDS for b locus Unordered:
a+ a+ a a a a a+ a+
2:4:2
a a a+ a+ a+ a+ a a
2:4:2
The six possible Neurospora ascospore patterns in respect to one locus.
other and therefore to the same centromere by examining classes 1 (parental ditype) and 2 (nonparental ditype) in table 6.7. If the two loci are unlinked, these two categories would represent two equally likely alternative events when no crossover takes place. Since category 1 represents almost 75% of all the asci, we can be sure the two loci are linked. To determine the distance of each locus to the centromere, we calculate one-half the percentage of second-division segregation patterns for each locus. For the a locus, classes 4, 5, 6, and 7 are second-division segregation patterns. For the b locus, classes 3, 5, 6, and 7 are second-division segregation patterns. Therefore,
the distances to the centromere, in map units, for each locus are for locus a: (1 2) 9 150 1 1,000 8 100
8.4 centimorgans for locus b: (1 2) 101 150 1 1,000 8 100
13.0 centimorgans
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
6. Linkage and Mapping in Eukaryotes
The McGraw Hill Companies, 2001
Haploid Mapping (Tetrad Analysis)
a a+ b
b+ b+
a b+
a+ a+ b b+
(5) or
a+ b+
a+ b
a b+
It should now be possible to describe exactly what type of crossover event produced each of the seven classes in table 6.7. Unfortunately, these two distances do not provide a unique determination of gene order. In gure 6.24, we see that two alternatives are possible: one has a map distance between the loci of 21.4 map units; the other has 4.6 map units between loci. How do we determine which of these is correct The simplest way is to calculate the a b distance using the unordered spore information. That is, the map distance is
map units (1 2) the number of TT asci the number of NPD asci total number of asci
a+ a+ b b+
a b+
(1 2)118 1,000
b+ b
a+ a+ b
a+ a+ b b+
a b+
b+ b+
a+ a+ b+ b
a+ a+
(8) b
a b+
b+ b
The eight random arrangements possible when a single crossover occurs between the a and b loci in Neurospora (see table 6.6). Circular arrows represent the rotation of a centromere from its position in the original con guration.
Since 6.2 map units is much closer to the a b distance expected if both loci are on the same side of the centromere, we accept alternative 2 in gure 6.24. A second way to choose between the alternatives in gure 6.24 is to nd out what happens to the b locus when a crossover occurs between the a locus and its centromere. If the order in alternative 1 is correct, crossovers between the a locus and its centromere should have no effect on the b locus; if 2 is correct, most of the crossovers that move the a locus in relation to its centromere should also move the b locus. Asci classes 4, 5, 6, and 7 include all the SDS patterns for the a locus. Of 168 asci, 150 (class 5) have similar SDS patterns for the b locus. Thus, 89% of the time, a crossover between the a locus and its centromere is also a crossover between the b locus and its centromere compelling evidence in favor of alternative 2. (What form would the data take if alternative 1 were correct ) In summary, mapping by tetrad analysis proceeds as follows. For both ordered and unordered spores, linkage is indicated by an excess of parental ditypes over nonparental ditypes. For unordered spores (yeast), the distance between two loci is one-half the number of
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