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Detached Figure 6.1
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Detached
bn bn det det (homozygous for the bn allele and the det + allele)
Wild-type bn +bn det +det
bn+bn + det det (homozygous for the bn + allele and the det allele)
Banded, detached
Testcross
bn det bn det + bn bn det +det
Banded
bn bn det det
bn +det bn +bn det det
Detached
bn +det + bn +bn det +det
Wild-type
( bn det
Phenotype
bn bn det det
Banded, detached 2
Number Figure 6.2
Testcrossing a dihybrid Drosophila.
gories in very high frequency have the same phenotypes as the original parents in the cross (P1 of g. 6.2). That is, banded ies and detached ies were the original parents as well as the great majority of the testcross offspring. We call these phenotypic categories parentals, or nonrecombinants. On the other hand, the testcross offspring in low frequency combine the phenotypes of the two original parents (P1).These two categories are referred to as nonparentals, or recombinants. The simplest explanation for these results is that the banded and detached loci are located near each other on the same chromosome (they are a linkage group), and therefore they move together as associated alleles during meiosis. We can analyze the original cross by drawing the loci as points on a chromosome ( g. 6.3). This shows that 99.5% of the testcross offspring (the nonrecombinants) come about through the simple linkage of the two loci. The remaining 0.5% (the recombinants) must have arisen through a crossover of homologues, from a chiasma at meiosis, between the two loci ( g. 6.4). Note that since it is not possible to tell from these crosses which chromosome the loci are actually on or where the centromere is in relation to the loci, the centromeres are not included in the gures. The crossover event is viewed as a breakage and reunion of two chromatids lying adjacent to each other during prophase I of meiosis. Later in this chapter, we nd cytological proof for this; in chapter 12, we explore the molecular mechanisms of this breakage and reunion process. From the testcross in gure 6.3, we see that 99.5% of the gametes produced by the dihybrid are nonrecombinant, whereas only 0.5% are recombinant. This very small frequency of recombinant offspring indicates that
the two loci lie very close to each other on their particular chromosome. In fact, we can use the recombination percentages of gametes, and therefore of testcross offspring, as estimates of distance between loci on a chromosome: 1% recombinant offspring is referred to as one map unit (or one centimorgan, in honor of geneticist T. H. Morgan, the rst geneticist to win the Nobel Prize; box 6.1). Although a map unit is not a physical distance, it is a relative distance that makes it possible to know the order of and relative separation between loci on a chromosome. In this case, the two loci are 0.5 map units apart. (From sequencing various chromosomal segments see chapter 13 we have learned that the relationship between centimorgans and DNA base pairs is highly variable, depending on species, sex, and region of the chromosome. For example, in human beings, 1 centimorgan can vary between 100,000 and 10,000,000 base pairs. In the ssion yeast, Schizosaccharomyces pombe, 1 centimorgan is only about 6,000 base pairs.) The arrangement of the bn and det alleles in the dihybrid of gure 6.3 is termed the trans con guration, meaning across, because the two mutants are across from each other, as are the two wild-type alleles. The alternative arrangement, in which one chromosome carries both mutants and the other chromosome carries both wild-type alleles ( g. 6.5), is referred to as the cis con guration. (Two other terms, repulsion and coupling, have the same meanings as trans and cis, respectively.) A cross involving two loci is usually referred to as a two-point cross; it gives us a powerful tool for dissecting the makeup of a chromosome. The next step in our
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