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Connecting Concepts 9

two heterozygotes; the 1:1 ratio, produced by a cross between a heterozygote and a homozygote; and the uniform progeny produced by a cross between two homozygotes. These simple phenotypic and genotypic ratios and the parental genotypes that produce them provide the key to understanding crosses for a single locus and, as you will see in the next section, for multiple loci.

Multiple-Loci Crosses

We will now extend Mendel's principle of segregation to more complex crosses for alleles at multiple loci. Understanding the nature of these crosses will require an additional principle, the principle of independent assortment.

Dihybrid Crosses

In addition to his work on monohybrid crosses, Mendel also crossed varieties of peas that differed in two characteristics (dihybrid crosses). For example, he had one homozy-gous variety of pea that produced round seeds and yellow endosperm; another homozygous variety produced wrinkled seeds and green endosperm. When he crossed the two, all the F1 progeny had round seeds and yellow endosperm. He then self-fertilized the F1 and obtained the following progeny in the F2: 315 round, yellow seeds; 101 wrinkled, yellow seeds; 108 round, green seeds; and 32 wrinkled, green seeds. Mendel recognized that these traits appeared approximately in a 9:3:3:1 ratio; that is, 9/16 of the progeny were round and yellow, 3/16 were wrinkled and yellow, 3/16 were round and green, and 1/16 were wrinkled and green.

The Principle of Independent Assortment

Mendel carried out a number of dihybrid crosses for pairs of characteristics and always obtained a 9:3:3:1 ratio in the F2. This ratio makes perfect sense in regard to segregation and dominance if we add a third principle, which Mendel recognized in his dihybrid crosses: the principle of independent assortment (Mendel's second law). This principle states that alleles at different loci separate independently of one another.

A common mistake is to think that the principle of segregation and the principle of independent assortment refer to two different processes. The principle of independent assortment is really an extension of the principle of segregation. The principle of segregation states that the two alleles of a locus separate when gametes are formed; the principle of independent assortment states that, when these two al-leles separate, their separation is independent of the separation of alleles at other loci.

Let's see how the principle of independent assortment explains the results that Mendel obtained in his dihybrid cross. Each plant possesses two alleles coding for each characteristic, so the parental plants must have had genotypes RRYY and rryy (^Figure 3.11a). The principle of segrega-

P generation

Round, yellow seeds

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