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The examples of mouse coat color, corn kernel color, and snapdragon ower petal color demonstrate that genes control the formation of enzymes, proteins that control the steps in biochemical pathways. For the most part, dominant alleles control functioning enzymes that catalyze biochemical steps. Recessive alleles often produce nonfunctioning enzymes that cannot catalyze speci c steps. Often a heterozygote is normal because one allele produces a functional enzyme; usually only half the enzyme quantity of the dominant homozygote is enough.The study of the relationship between genes and enzymes is generally called biochemical genetics because it involves the genetic control of biochemical pathways. A. E. Garrod, a British physician, pointed out this general concept of human gene action in Inborn Errors of Metabolism, published in 1909. Only nine years after Mendel was rediscovered, Garrod described several human conditions, such as albinism and alkaptonuria, that occur in individuals who are homozygous for recessive alleles (see g. 2.24).
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9:3:4 Figure 2.26
Flower color inheritance in snapdragons. This is an example of epistasis: an nn genotype masks the expression of alleles (EE, Ee, or ee) at the eosinea gene.
Table 2.4 Some Examples of Epistatic Interactions Among Alleles of Two Genes
Characteristic Corn and sweet pea color Mouse coat color Shepherd s purse seed capsule shape Summer squash shape Fowl color Phenotype of F1 Dihybrid (AaBb) Purple Agouti Triangular Disk White Phenotypic F2 Ratio Purple:white 9:7 Agouti:black:albino 9:3:4 Triangular:oval 15:1 Disk:sphere:elongate 9:6:1 White:colored 13:3
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
For example, people normally degrade homogentisic acid (alkapton) into maleylacetoacetic acid. Persons with the disease alkaptonuria are homozygous for a nonfunctional form of the enzyme essential to the process: homogentisic acid oxidase, found in the liver. Absence of this enzyme blocks the degradation reaction so that homogentisic acid builds up. This acid darkens upon oxidation. Thus, affected persons can be identi ed by the black color of their urine after its exposure to air. Eventually, alkaptonuria causes problems in the joints and a darkening of cartilage that is visible in the ears and the eye sclera.
One-Gene-One-Enzyme Hypothesis
Pioneering work in the concept that genes control the production of enzymes, which in turn control the steps in biochemical pathways, was done by George Beadle and Edward Tatum, who eventually shared the Nobel Prize for their work. They not only put forth the one-gene-oneenzyme hypothesis, but also used mutants to work out the details of biochemical pathways. In 1941, Beadle and Tatum were the rst scientists to isolate mutants with nutritional requirements that de ned steps in a biochemical pathway. In the early 1940s, they united the elds of biochemistry and genetics by using strains of a bread mold with speci c nutritional requirements to discover the steps in biochemical pathways in that organism. Through this century, the study of mutations has been the driving force in genetics. The process of mutation produces alleles that differ from the wild-type and shows us that a particular aspect of the phenotype is under genetic control. Beadle and Tatum used mutants to work out the steps in the biosynthesis of niacin (vitamin B3) in pink bread mold, Neurospora crassa. Normally, Neurospora synthesizes niacin via the pathway shown in gure 2.27. Beadle and Tatum isolated mutants that could not grow unless niacin was provided in
George W. Beadle (1903 89). Courtesy of the
Archives, California Institute of Technology.
Edward L. Tatum (1909 75). Courtesy of the
Proceedings for the National Academy of Sciences.
the culture medium; these mutants had enzyme de ciencies in the synthesizing pathway that ends with niacin. Thus, although wild-type Neurospora could grow on a medium without additives, the mutants could not. Beadle and Tatum had a general idea, based on the structure of niacin, as to what substances would be in the niacin biosynthesis pathway. They could thus make educated guesses as to what substances they might add to the culture medium to enable the mutants to grow. Mutant B (table 2.5), for example, could grow if given niacin or, alternatively, 3-hydroxyanthranilic acid. It could not grow if given only kynurenine. Thus, Beadle and Tatum knew that the B mutation affected the pathway between kynurenine and 3-hydroxyanthranilic acid. Similarly, mutant A could grow if given 3-hydroxyanthranilic acid or kynurenine instead of niacin. Therefore, these two prod-
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