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23.7 (measured map distance in a two-point cross; double recombinants between b and c are masked) Figure 6.9 Tentative map of the black, purple, and curved chromosome in Drosophila. Numbers are map units (centimorgans).
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us to convert a measured map distance into a more accurate one. Theoretically, it shows that measured map distance never exceeds 50 map units in any one cross. Multiple crossovers reduce the apparent distance between two loci to a maximum of 50 map units, the value that independent assortment produces (50% parentals, 50% recombinants).
Gene Order
Although we performed the previous analysis merely assuming that pr was in the middle, the data in table 6.1 con rm our original assumption that the gene order is b pr c. Of the four pairs of reciprocal phenotypic classes in table 6.1, one pair has the highest frequency (5,701 and 5,617) and one pair has the lowest (60 and 72). The pair with highest frequency is the nonrecombinant group. The one with the lowest frequency is the double recombinant group, the one in which only the middle locus has been changed from the parental arrangement. A comparison of either of the double recombinant classes with either of the nonrecombinant classes shows the gene that is in the middle and, therefore, the gene order. In other words, the data allow us to determine gene order. Since b pr c was one of the nonrecombinant gametes, and b pr c was one of the double recombinant gametes, pr stands out as the changed locus, or the one in the middle. In a similar manner, comparing b pr c with b pr c would also point to pr as the inside locus (or inside marker). So would comparing b pr c with b pr c or b pr c with b pr c . In each case, the middle locus, pr, displays the different pattern, whereas the allelic arrangements of the outside markers, b and c, behave in concert. If this seems confusing, simply compare the double crossovers and nonrecombinants to nd one of each in which two alleles are identical. For example, the double recombinant b pr c and the nonrecombinant b pr c share the b and c alleles. The pr locus is mutant in one case and wild-type in the other. Hence, pr is the locus in the middle. From the data in table 6.1, we can con rm the association of alleles in the trihybrid parent. That is, since the data came from testcrossing a trihybrid, the allelic con guration in that trihybrid is re ected in the nonrecombinant classes of offspring. In this case, one is the result of a b pr c gamete, the other, of a b pr c gamete.Thus, the trihybrid had the genotype b pr c/b pr c : all alleles were in the cis con guration.
ble recombinants equal the expected number In the example, there were 132/15,000 double crossovers, or 0.88%. The expected number is based on the independent occurrence of crossing over in the two regions measured. That is, 5.9% of the time there is a crossover in the b pr region, which we can express as a probability of occurrence of 0.059. Similarly, 19.5% of the time there is a crossover in the pr c region, or a probability of occurrence of 0.195. A double crossover should occur as a product of the two probabilities: 0.059 0.195 0.0115. This means that 1.15% of the gametes (1.15% of 15,000 172.5) should be double recombinants. In our example, the observed number of double recombinant offspring is lower than expected (132 observed, 172.5 expected). This implies a positive interference, in which the occurrence of the rst crossover reduces the chance of the second. We can express this as a coef cient of coincidence, de ned as observed number of double recombinants expected number of double recombinants In the example, the coef cient of coincidence is 132/172.5 0.77. In other words, only 77% of the expected double crossovers occurred. Sometimes we express this reduced quantity of double crossovers as the degree of interference, de ned as interference 1 coef cient of coincidence
In our example, the interference is 23%. It is also possible to have negative interference, in which we observe more double recombinants than expected. In this situation, the occurrence of one crossover seems to enhance the probability that crossovers will occur in adjacent regions.
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