..and half will have recombinant chromosomes.

nonrecombinants. The other two chromatids, which did participate in crossing over, now contain new combinations of alleles; gametes that receive these chromatids are recombinants. For each meiosis in which a single crossover takes place, then, two nonrecombinant gametes and two recombinant gametes will be produced. This result is the same as that produced by independent assortment (see Figure 7.5b); so, when crossing over between two loci takes place in every meiosis, it is impossible to determine whether the genes were linked and crossing over took place or whether the genes are on different chromosomes.

For closely linked genes, crossing over does not take place in every meiosis. In meioses in which there is no crossing over, only nonrecombinant gametes are produced. In meioses in which there is a single crossover, half the gametes are recombinants and half are nonrecombinants (because a single crossover only affects two of the four chromatids); so the total percentage of recombinant gametes is always half the percentage of meioses in which crossing over takes place. Even if crossing over between two genes takes place in every meiosis, only 50% of the resulting gametes will be recombinants. Thus, the frequency of recombinant gametes is always half the frequency of crossing over, and the maximum proportion of recombinant gametes is 50%.

Concepts ®

Linkage between genes causes them to be inherited together and reduces recombination; crossing over breaks up the associations of such genes. In a testcross for two linked genes, each crossover produces two recombinant gametes and two nonrecombinants. The frequency of recombinant gametes is half the frequency of crossing over, and the maximum frequency of recombinant gametes is 50%.

Application Let us apply what we have learned about linkage and recombination to a cross between tomato plants that differ in the genes that code for leaf type and plant height. Assume now that these genes are linked and that some crossing over takes place between them. Suppose a geneticist carried out the testcross outlined earlier:

When crossing over takes place between the genes for leave type and height, two of the four gametes produced will be recombinants. When there is no crossing over, all four resulting gametes will be nonrecombinants. Thus, over all, the majority of gametes will be nonrecombinants. These gametes then unite with gametes produced by the homozygous recessive parent, which contain only the recessive alleles, resulting in mostly nonrecombinant progeny and a few recombinant progeny ( FIGURE 7.7). In this cross, we see that 55 of the testcross progeny have normal leaves and are tall and 53 have mottled leaves and are dwarf. These plants are the nonrecom-binant progeny, containing the original combinations of traits that were present in the parents. Of the 123 progeny, 15 have new combinations of traits that were not seen in the parents: 8 are normal leaved and dwarf, and 7 are mottle leaved and tall. These plants are the recombinant progeny.

The results of a cross such as the one illustrated in Figure 7.7 reveal several things. A testcross for two independently assorting genes is expected to produce a 1:1:1:1 phenotypic ratio in the progeny. The progeny of this cross clearly do not exhibit such a ratio; so we might suspect that

recombinant frequency the genes are not assorting independently. When linked genes undergo crossing over, the result is mostly nonrecombinant progeny and fewer recombinant progeny. This result is what we observe among the progeny of the testcross illustrated in Figure 7.7; so we conclude that two genes show evidence of linkage with some crossing over.

Calculation of Recombination Frequency

The percentage of recombinant progeny produced in a cross is called the recombination frequency, which is calculated as follows:

recombinant frequency number of recombinant progeny inno. ' ; - X 1UU%

total number of progeny

In the testcross shown in Figure 7.7, 15 progeny exhibit new combinations of traits; so the recombination frequency is:

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