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Mendel also analyzed the inheritance pattern of traits observed two at a time. He looked, for instance, at plants that differed in the form and color of their peas: he crossed true-breeding (homozygous) plants that had seeds that were round and yellow with plants that produced seeds that were wrinkled and green. Mendel s results appear in gure 2.14. The F1 plants all had round, yellow seeds, which demonstrated that round was dominant to wrinkled and yellow was dominant to green. When these F1 plants were self-fertilized, they produced
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Reginald C. Punnett (1875 1967).
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From Genetics, 58 (1968): frontispiece. Courtesy of the Genetics Society of America.
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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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Round, yellow
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Wrinkled, green
RRYY
Gametes
rryy ry
RrYy
Round, yellow (RRYY )
Wrinkled, green (rryy )
Gametes
1 : 1 : 1 : 1
X Self
Pollen
Ry RRYy
rY RrYY
ry RrYy
RRYY
Round, yellow (RrYy ) Ovules
RRYy
RRyy
RrYy
Rryy
RrYY
RrYy
rrYY
rrYy
Round, yellow (315) (RRYY; RRYy; RrYY; RrYy )
Round, green (108) (RRyy; Rryy )
RrYy
Rryy
rrYy
rryy
Assigning genotypes to the cross in gure 2.14.
Wrinkled, yellow (101) (rrYY; rrYy ) Figure 2.14
Wrinkled, green (32) (rryy )
Independent assortment in garden peas.
Rule of Independent Assortment
This ratio comes about because the two characteristics behave independently. The F1 plants produce four types of gametes (check g. 2.15): RY, Ry, rY, and ry. These gametes occur in equal frequencies. Regardless of which seed shape allele a gamete ends up with, it has a 50:50
chance of getting either of the alleles for color the two genes are segregating, or assorting, independently. This is the essence of Mendel s second rule, the rule of independent assortment, which states that alleles for one gene can segregate independently of alleles for other genes. Are the alleles for the two characteristics of color and form segregating properly according to Mendel s rst principle If we look only at seed shape (see g. 2.14), we nd that a homozygote with round seeds was crossed with a homozygote with wrinkled seeds in the P1 generation (RR rr). This cross yields only heterozygous plants with round seeds (Rr) in the F1 generation. When these
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
2. Mendel s Principles
The McGraw Hill Companies, 2001
Two Mendel s Principles
BOX 2.1
n February and March of 1865, Mendel delivered two lectures to the Natural History Society of Br nn. These were published as a single forty-eight-page article handwritten in German. The article appeared in the 1865 Proceedings of the Society, which came out in 1866. It was entitled Versuche ber P anzen-Hybriden, which means Experiments in Plant Hybridization. Following are some paragraphs from the English translation to give us some sense of the original. In his introductory remarks, Mendel writes:
That, so far, no generally applicable law governing the formation and development of hybrids has been successfully formulated can hardly be wondered at by anyone who is acquainted with the extent of the task, and can appreciate the dif culties with which experiments of this class have to contend. A nal decision can only be arrived at when we shall have before us the results of detailed experiments made on plants belonging to the most diverse orders. Those who survey the work done in this department will arrive at the conviction that among all the numerous experiments made, not one has been carried out to such an extent and in such a way as to make it possible to determine the number of different forms under which the offspring of hybrids appear, or to arrange these forms with certainty according to their separate genera-
Historical Perspectives
Excerpts from Mendel s Original Paper
tions, or de nitely to ascertain their statistical relations. . . . The paper now presented records the results of such a detailed experiment. This experiment was practically con ned to a small plant group, and is now, after eight years pursuit, concluded in all essentials. Whether the plan upon which the separate experiments were conducted and carried out was the best suited to attain the desired end is left to the friendly decision of the reader.
are transmitted entire, or almost unchanged in the hybridization, and therefore in themselves constitute the characters of the hybrid, are termed the dominant, and those which become latent in the process, recessive. The expression recessive has been chosen because the characters thereby designated withdraw or entirely disappear in the hybrids, but nevertheless reappear unchanged in their progeny, as will be demonstrated later on.
He then writes about the F2 generation:
In this generation there reappear, together with the dominant characters, also the recessive ones with their peculiarities fully developed, and this occurs in the de nitely expressed average proportion of three to one, so that among each four plants of this generation three display the dominant character and one the recessive. This relates without exception to all the characters which were investigated in the experiments. The angular wrinkled form of the seed, the green colour of the albumen, the white colour of the seed-coats and the owers, the constrictions of the pods, the yellow colour of the unripe pod, of the stalk, of the calyx, and of the leaf venation, the umbel-like form of the in orescence, and the dwarfed stem, all reappear in the numerical proportion given, without any essential alteration. Transitional forms were not observed in any experiment. . . .
After discussing the origin of his seeds and the nature of the experiments, Mendel discusses the F1, or hybrid, generation:
This is precisely the case with the Pea hybrids. In the case of each of the seven crosses the hybridcharacter resembles that of one of the parental forms so closely that the other either escapes observation completely or cannot be detected with certainty.This circumstance is of great importance in the determination and classi cation of the forms under which the offspring of the hybrids appear. Henceforth in this paper those characters which
F1 plants are self-fertilized, the result is 315 108 round seeds (RR or Rr) and 101 32 wrinkled seeds (rr) in the F2 generation. This is a 423:133 or a 3.18:1.00 phenotypic ratio very close to the expected 3:1 ratio. So the gene for seed shape is segregating normally. In a similar manner, if we look only at the gene for color, we see that the F2 ratio of yellow to green seeds is 416:140, or
2.97:1.00 again, very close to a 3:1 ratio. Thus, when two genes are segregating normally according to the rule of segregation, their independent behavior demonstrates the rule of independent assortment (box 2.1). From the Punnett square in gure 2.15, you can see that because of dominance, all phenotypic classes except the homozygous recessive one wrinkled, green
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