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Let us work out one more three-point cross, in which neither the middle gene nor the cis-trans relationship of the alleles in the trihybrid F1 parent is given. On the third chromosome of Drosophila, hairy (h) causes extra bristles on the body, thread (th) causes a thread-shaped arista (antenna tip), and rosy (ry) causes the eyes to be reddish brown. All three traits are recessive. Trihybrid females were testcrossed; the phenotypes from one thousand offspring are listed in table 6.2. At this point, it is possible to use the data to determine the parental genotypes (the P1 generation, assuming that they were homozygotes), the gene order, the map distances, and the coef cient of coincidence. The table presents the data in no particular order, as an experimenter might have recorded them. Phenotypes are tabulated and, from these, the genotypes can be reconstructed. Notice that the data can be put into the form found in table 6.1;
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Coef cient of Coincidence
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The next question in our analysis of this three-point cross is, are crossovers occurring independently of each other That is, does the observed number of dou-
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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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Table 6.2 Offspring from a Trihybrid (h h ry ry
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th th) Testcross (x hh ryry thth) in Drosophila
Genotype (order unknown) h ry th/h ry th h ry th/h ry th h ry th/h ry th h ry th/h ry th h ry th /h ry th h ry th /h ry th h ry th /h ry th h ry th /h ry th
Table 6.3 Data from Table 6.2 Arranged to Show
Recombinant Regions
Trihybrid s Gamete
Number 359 351 98 92 47 43 4 6 1,000
h th
th ry
h ry
Phenotype Thread Rosy, thread Hairy, rosy, thread Hairy, thread Rosy Hairy, rosy Wild-type Hairy
Number 359 47 4 98 92 351 6 43
h th ry h th ry h th ry h th ry h th ry h th ry h th ry h th ry Total
98 92 47 43 4 6 200 4 6 100
98 92 47 43
we see a large reciprocal set (359 and 351), a small reciprocal set (4 and 6), and large and small intermediate sets (98 and 92, 47 and 43). From the data presented, is it obvious that the three loci are linked The pattern, as just mentioned, is identical to that of the previous example, in which the three loci were linked. (What pattern would appear if two of the loci were linked and one assorted independently See problem 6 at the end of the chapter.) Next, what is the allelic arrangement in the trihybrid parent The offspring with the parental, or nonrecombinant, arrangements are the reciprocal pair in highest frequency. Table 6.2 shows that thread and hairy, rosy offspring are the nonrecombinants.Thus, the nonrecombinant gametes of the trihybrid F1 parent were h ry th and h ry th, which is the allelic arrangement of the trihybrid with the actual order still unknown h ry th /h ry th. (What were the genotypes of the parents of this trihybrid, assuming they were homozygotes ) Continuing, which gene is in the middle From table 6.2, we know that h ry th and h ry th are the double recombinant gametes of the trihybrid parent because they occur in such low numbers. Comparison of these chromosomes with either of the nonrecombinant chromosomes (h ry th or h ry th ) shows that the thread (th) locus is in the middle. We now know that the original trihybrid had the following chromosomal composition: h th ry/h th ry . The h and ry alleles are in the cis con guration, with th in the trans con guration. We can now compare the chromosome from the trihybrid in each of the eight offspring categories with the parental arrangement and determine the regions that had crossovers. Table 6.3 does this. We can see that the h th distance is 20 map units, the th ry distance is 10 map units, and the apparent h ry distance is 28 map units
( g. 6.11). As in the earlier example, the h ry discrepancy is from not counting the double crossovers twice each: 280 2(10) 300, which is 30 map units and the more accurate gure. Last, we wish to know what the coef cient of coincidence is. The expected occurrence of double recombinants is 0.200 0.100 0.020, or 2%. Two percent of 1,000 20. Thus coef cient of coincidence observed number of double recombinants expected number of double recombinants 10/20 0.50 Only 50% of the expected double crossovers occurred. Geneticists have mapped the chromosomes of many eukaryotic organisms from three-point crosses of this type those of Drosophila are probably the most extensively studied. Drosophila and other species of ies have giant polytene salivary gland chromosomes, which arise as a result of endomitosis. In this process,
30 (all recombinants; best estimate of true map distance)
28.0 (measured map distance) Figure 6.11 Map of the h th ry region of the Drosophila chromosome, with numerical discrepancy in distances. Numbers are map units (centimorgans).
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