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Gregor Mendel was an Austrian monk (of Br nn, Austria, which is now Brno, Czech Republic). In his experiments, he tried to crossbreed plants that had discrete, nonoverlapping characteristics and then to observe the distribution of these characteristics over the next several generations. Mendel worked with the common garden pea plant, Pisum sativum. He chose the pea plant for at least three reasons: (1) The garden pea was easy to cultivate and had a relatively short life cycle. (2) The plant had discontinuous characteristics such as ower color and pea texture. (3) In part because of its anatomy, pollination of the plant was easy to control. Foreign pollen could be kept out, and cross-fertilization could be accomplished arti cially. Figure 2.2 shows a cross section of the pea ower that indicates the keel, in which the male and female parts develop. Normally, self-fertilization occurs when pollen falls onto the stigma before the bud opens. Mendel cross-fertilized the plants by opening the keel of
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Gregor Johann Mendel (1822 84).
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permission of the Moravski Museum, Mendelianum.)
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Tamarin: Principles of Genetics, Seventh Edition
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II. Mendelism and the Chromosomal Theory
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2. Mendel s Principles
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Filament Stigma Anther Style Ovary Keel (half cut away)
ent forms of a gene that exist within a population are termed alleles. The terms dominant and recessive are used to describe both the relationship between the alleles and the traits they control. Thus, we say that both the allele for tallness and the trait, tall, are dominant. Dominance applies to the appearance of the trait when both a dominant and a recessive allele are present. It does not imply that the dominant trait is better, is more abundant, or will increase over time in a population. When the F1 offspring of gure 2.4 were selffertilized to produce the F2 generation, both tall and dwarf offspring occurred; the dwarf characteristic reappeared. Among the F2 offspring, Mendel observed 787 tall and 277 dwarf plants for a ratio of 2.84:1. It is an indication of Mendel s insight that he recognized in these numbers an approximation to a 3:1 ratio, a ratio that suggested to him the mechanism of inheritance at work in pea plant height.
Anatomy of the garden pea plant ower. The female part, the pistil, is composed of the stigma, its supporting style, and the ovary. The male part, the stamen, is composed of the pollen-producing anther and its supporting lament.
S E G R E G AT I O N
Rule of Segregation
Mendel assumed that each plant contained two determinants (which we now call genes) for the characteristic of height. For example, a hybrid F1 pea plant possesses the dominant allele for tallness and the recessive allele for dwarfness for the gene that determines plant height. A pair of alleles for dwarfness is required to develop the recessive phenotype. Only one of these alleles is passed into a single gamete, and the union of two gametes to form a zygote restores the double complement of alleles. The fact that the recessive trait reappears in the F2 generation shows that the allele controlling it was hidden in the F1 individual and passed on unaffected. This explanation of the passage of discrete trait determinants, or genes, comprises Mendel s rst principle, the rule of segregation. The rule of segregation can be summarized as follows: A gamete receives only one allele from the pair of alleles an organism possesses; fertilization (the union of two gametes) reestablishes the double number. We can visualize this process by redrawing gure 2.4 using letters to denote the alleles. Mendel used capital letters to denote alleles that control dominant traits and lowercase letters for alleles that control recessive traits. Following this notation, T refers to the allele controlling tallness and t refers to the allele controlling shortness (dwarf stature). From gure 2.5, we can see that Mendel s rule of segregation explains the homogeneity of the F1 generation (all tall) and the 3:1 ratio of tall-to-dwarf offspring in the F2 generation. Let us de ne some terms. The genotype of an organism is the gene combination it possesses. In gure 2.5,
a ower before the anthers matured and placing pollen from another plant on the stigma. In the more than ten thousand plants Mendel examined, only a few were fertilized other than the way he had intended (either self- or cross-pollinated). Mendel used plants obtained from suppliers and grew them for two years to ascertain that they were homogeneous, or true-breeding, for the particular characteristic under study. He chose for study the seven characteristics shown in gure 2.3. Take as an example the characteristic of plant height. Although height is often continuously distributed, Mendel used plants that displayed only two alternatives: tall or dwarf. He made the crosses shown in gure 2.4. In the parental, or P1, generation, dwarf plants pollinated tall plants, and, in a reciprocal cross, tall plants pollinated dwarf plants, to determine whether the results were independent of the parents sex. As we will see later on, some traits follow inheritance patterns related to the sex of the parent carrying the traits. In those cases, reciprocal crosses give different results; with Mendel s tall and dwarf pea plants, the results were the same. Offspring of the cross of P1 individuals are referred to as the rst lial generation, or F1. Mendel also referred to them as hybrids because they were the offspring of unlike parents (tall and dwarf). We will speci cally refer to the offspring of tall and dwarf peas as monohybrids because they are hybrid for only one characteristic (height). Since all the F1 offspring plants were tall, Mendel referred to tallness as the dominant trait. The alternative, dwarfness, he referred to as recessive. Differ-
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