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Determining a Genetic Distance ~ Llncferstandlng Inheritance DETERMINING A GENETIC DISTANCE ~ The classical method for determining the The participants in the first breeding are a meioses in the female that involve a single genetic distance between the loci of two female fruit fly that is heterozygous for both crossover between the two loci (or any odd allele pairs known to reside on the same traits (and therefore has red eyes and nor- number of crossovers) yield in addition two homologous chromosome pair of an organ- mal wings) and a male fruit fly that is ho- other types of eggs: those possessing a ism involves observing the phenotypes of mozygous for both recessive trait variants chromosome with the allele combination Ab the offspring of one of two particular breed- (and therefore has purple eyes and vestigial and those possessing a chromosome with ings. During the course of Thomas Hunt wings). Furthermore, the female is known the allele combination aB. In other words, a Morgan’s work on fruit flies, he happened to to be a product of the breeding AABB x single crossover between the two loci es- carry out both breedings and was rewarded aabb. Therefore the distribution of the alle- tablishes linkage between one dominant not only with the first clear evidence of les A, a, B, and bon the homologous auto- and one recessive allele, On the other crossing over but also with the first unam- some pair of the female is known: Both hand, meioses in the doubly homozygous biguous genetic-distance data. Morgan’s dominant alleles (A and B) reside on one male, whether or not they invove cross- experiments and data are used here to member of the homologous autosome pair, overs between the two loci, yield sperms illustrate the procedure. and both recessive alleles (a and b) reside possessing only the allele combination ab. on the other member. Such an allele distri- Thus the offspring of breeding 1 possess The allele pairs in question reside on one of bution is denoted by writing the genotype of four genotypes, each corresponding to one the homologous autosome pairs of Dro- the female as AB/ab. The distribution of the of the four possible phenotypes: sophila frrekmogaster. One allele pair af- alleles a, a, b, and b on the homologous fects eye color: a dominant allele A that autosome pair of the male is also known AB/ab female x ab/ab male — ‘> specifies red eye color and a recessive (because the male is homozygous for both AB/ab + ab/ab + Ab/ab + aB/ab. allele a that specifies purple eye color. The traits) and is denoted in a similar fashion as other allele pair affects wing length: a domi- ab/ab. Thus the first breeding can be sym- Morgan examined more than 2800 progeny nant allele B that specifies wild-type wings bolized by of breeding 1 and found that 47.2 percent and {3 recessive allele b that specifies ves- had red eyes and normal wings (AB/ah), tigial (very short) wings. AB/ab female x ab/ab male, (1) 42.1 percent had purple eyes and vestigial wings (ab/ah), 5.3 percent had red eyes and Meioses in the heterozygous female that vestigial wings (Ab/ah), and 5.4 percent had involve no crossovers between the two loci purple eyes and normal wings (aB/ab). All yield two types of eggs: those possessing the offspring exhibiting the last two pheno- the chromosome with the allele combina- types (the combinations of one recessive tion AB and those possessing the chromo- trait variant and one dominant trait variant) some with the allele combination ab, In result only from crossovers during meioses .. other words, the two dominant alleles and in the female parent. Thus the data indicate }: the two recessive alleles remain linked that the probability of new allele linkages (resident on the same chromo- being formed by crossing over is 0,107 = . ,/;.- -=’:J< some), just as they are in the 0,053 + 0.054, That value for the so-called t ‘\ il .. &p female herself. But those recombination fraction corresponds to a k ~:>._=.:<..\.\--- #~.. { >+ ‘ genetic distance of about 12 centimorgans. :> ‘“’,\ ‘;’;2:/==% (The relationship between recombinaticm .,i::,.~; .’.,f - fraction and genetic distance is presented “;:2?21 {“”’”””w’ in “Classical Linkage Mapping” in ‘fMappirlg /“” ““’’”””’”’”$~ the Genome,”) ‘.:i :> ‘G;~{/ / “’* ‘4’ /,,T’) ~~ The participants in the other breeding that ““”’””’”’””’l “* .&y,3- ,-- ‘ / provides unambiguous recombination-frac- ,1’ — .Aj * .-..r+ ‘ ,, / “’%’ tion data are, like the participants in breed- ,- ing 1, a doubly heterozygous female and a .// / .$,\ doubly homozygous-recessive male. How- /~ .’ 34 Understanding Inheritance ever, the second female is known to be a during meioses in the ,\---- product 01 the breeding Ab/Ab x aB/aB male parent rather than in the (rather than the breeding AB/AB x ab/ab). female parent. However, for some Therefore the distribution of alleles on her unknown reason crossing over simply homologous autosome pair is Ab/aB (rather does not occur in male fruit flies. But fruit than AB/a~5). (The difference in allele distri- flies are exceptional in that respect, and The reader can accept on faith or verify butions of the two doubly heterozygous genetic distances for other species can be personally that breedings 1 and 2 are the females is often referred to as a difference determined by carrying out either breeding only breedings that provide unambiguous in linkage phase.) The second breeding is 1, say, or its reciprocal. recombination-fraction and hence genetic- thus symbolized by distance data. Note, in addition, that obtain- Breedings 1 and 2 are those that provide ing even ambiguous data requires that one Ab/aB female x ab/ab male. (2) unambiguous recombination-fraction data. parent be doubly heterozygous. As an example of the ambiguities that can arise, consider the fruit-fly breeding Determining a genetic distance is thus rela- Breeding 2 yields offspring that exhibit the tively easy when the breeding of the organ- same genotypes and phenotypes as breed- AB/ab female x AB/ab male. (3) ism in question can be manipulated at will. ing 1: But determining the genetic distance be- Assume first that crossing over between the tween the loci of two human allele pairs is Ab/aE? female x ab/ab male two loci does not occur during meioses in much more difficult, since the breeding of Ab/’b + aB/ab + AB/ab + ab/ab. the female parent. Then the offspring of humans cannot be manipulated, the geno- breeding 3 exhibit two phenotypes: red eyes types and allele distributions of human par- Morgan examined more than 2300 progeny and normal wings (Af3/Af3 and A/3/ah) and ents are not always known, and human of breeding 2 and found that 41.3 percent purple eyes and vestigial wings (ab/ab). breedings generally produce so few off- had red eyes and vestigial wings (Ab/ah), Now assume that crossing over does occur spring that the statistical uncertainty in the 45.7 percent had purple eyes and normal during meioses in the female parent, Then measured recombination fraction is large. wings (aE1/ah), 6.7 percent had red eyes among the offspring of breeding 3 are some and normal wings (AB/ah), and 6.3 percent that exhibit the two other possible pheno- had purple eyes and vestigial wings (ah/ types: red eyes and vestigial wings (Ab/ab) ah), Agaitl, all the offspring exhibiting the and purple eyes and normal wings (aB/ab). last two phenotypes result only from cross- All offspring that exhibit those two pheno- overs during meioses in the female parent. types result only from crossing over, How- Thus the clata indicate that the recombina- ever, crossing over also leads to offspring tion fraction for the two allele pairs is 0.130, that exhibit one of the phenotypes produced which corresponds to a genetic distance of in the absence of crossing over, namely, red about 15 centimorgans, eyes and normal wings (Ab\ABand aB/Af3). In other words, whereas the offspring pro- Note that the two data sets yield different duced by breeding 1 or 2 can values forihe same genetic distance. How- unambiguously be sorted by pheno- >>+.<—‘1 , ever, the clifference between the values is type into two categories-those that ‘“ }.,.<$1<- ,..: ! ~,: Yj within the statistical uncertainties associ- are the result of crossovers and those ,-.’ ated with measurements of probabilistic that are not—the offspring resulting from’ “”~ events. Note also that the same genetic breeding 3 cannot be so sorted because distance could in principle be determined meioses that do and do not involve cross- by carrying out the reciprocal of breeding 1 overs result in the doubly dominant pheno- or breedimg 2 (that is, a breeding between type, a doubly heterozygous male and a doubly homozygous-recessive female). Then, the crossovers detected are those that occur 35 Understanding Inheritance cell displays a characteristic pattern of dark and light bands when stained with an appropriate dye (see “Chromosomes: The Sites of Hereditary Information”). BecaLlse the banding pattern characteristic of a pair of homologous metaphase chromosomes varies along the length of the chromosomes, it can also be used to identify different regions of the chromosomes. The advent of chromosome banding led to recognition of the occasional occurrence of aberrant chromosomes. (The incidence of aberrant chromosomes, like the incidence of gene mutations, can be increased by exposure to x rays or other mutagenic agents.) Several types of chromosome aberrations, or rearrangements, were noted, including translocations (the exchange of chromosome regions between nonhomologous chromosomes) and inversions (the reversal of the orientation of a chromosome region). Obviously a chromosome rearrangement can lead to changes in the complement ,of genes present on a chromosome or to changes in their relative locations.
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