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Figure 2.27 Pathway of niacin synthesis in Neurospora. Each arrow represents an enzyme-mediated step. Each question mark represents a presumed but (at the time Beadle and Tatum were working) unknown compound.
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Tamarin: Principles of Genetics, Seventh Edition
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II. Mendelism and the Chromosomal Theory
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The McGraw Hill Companies, 2001
Biochemical Genetics
Table 2.5 Growth Performance of Neurospora Mutants (plus sign indicates growth;
minus sign indicates no growth)
Additive Tryptophan Wild-type Mutant A Mutant B Kynurenine 3-Hydroxyanthranilic Acid Niacin
ucts must be in the pathway after the step interrupted in mutant A. Conversely, since neither of these mutant organisms could grow when given only tryptophan, Beadle and Tatum knew that tryptophan occurred in the pathway before the steps with the de cient enzymes. By this type of analysis, they discovered the steps in several biochemical pathways of Neurospora. Many biochemical pathways are similar in a huge range of organisms, and thus Beadle and Tatum s work was of general importance. (We will spend more time studying Neurospora in chapter 6.) Beadle and Tatum could further verify their work by observing which substances accumulated in the mutant organisms. If a biochemical pathway is blocked at a certain point, then the substrate at that point cannot convert into the next product, and it builds up in the cell. For example, in the niacin pathway ( g. 2.27), if a block occurs just after 3-hydroxyanthranilic acid, that substance will build up in the cell because it cannot convert into the next substance on the way to niacin. This analysis could be misleading, however, if the built-up substance is being siphoned off into other biochemical pathways in the cell. Also, the cell might attempt to break down or sequester toxic substances. This would mean there might not be an obvious buildup of the substance just before the blocked step. Beadle and Tatum concluded from their studies that one gene controls the production of one enzyme. The one-gene-one-enzyme hypothesis is an oversimpli cation that we will clarify later in the book. As a rule of thumb, however, the hypothesis is valid, and it has served to direct attention to the functional relationship between genes and enzymes in biochemical pathways. Although a change in a single enzyme usually disrupts a single biochemical pathway, it frequently has more than one effect on phenotype. Multiple effects are referred to as pleiotropy. A well-known example is sickle-cell anemia, caused by a mutation in the gene for the chain of the hemoglobin molecule. In a homozygote, this mutation causes a sickling of red blood cells ( g. 2.28). The sickling of these cells has two major rami cations.
First, the liver destroys the sickled cells, causing anemia. The phenotypic effects of this anemia include physical weakness, slow development, and hypertrophy of the bone marrow, resulting in the tower skull seen in some of those af icted with the disease. The second major effect of sickle-cell anemia is that the sickled cells interfere with capillary blood ow, clumping together and resulting in damage to every major organ. The individual can suffer pain, heart failure, rheumatism, and other ill effects. Hence, a single mutation shows itself in many aspects of the phenotype.
Sickle-shaped red blood cells from a person with sickle-cell anemia. Red blood cells are about 7 to 8 m in diameter. (Courtesy of Dr. Patricia N. Farnsworth.)
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
S U M M A R Y
STUDY OBJECTIVE 1: To understand that genes are discrete units that control the appearance of an organism 17 18 Genes control phenotypic traits such as size and color.They are inherited as discrete units. STUDY OBJECTIVE 2: To understand Mendel s rules of inheritance: segregation and independent assortment 18 22 Higher organisms contain two alleles of each gene, but only one allele enters each gamete. Zygote formation restores the double number of alleles in the cell. This is Mendel s rule of segregation. Alleles of different genes segregate independently of each other. Mendel was the rst to recognize the 3:1 phenotypic ratio as a pattern of inheritance; the 9:3:3:1 ratio demonstrates independent assortment in hybrids. Mendel was successful in his endeavor because he performed careful experiments using discrete characteristics, large numbers of offspring, and an organism (the pea plant) amenable to controlled fertilizations. STUDY OBJECTIVE 3: To understand that dominance is a There can be many alleles for one gene, although each individual organism has only two alleles for each gene. A phenotype is dominant if it is expressed when one or two copies of its allele are present (heterozygote or homozygote). Dominance depends, however, on the level of the phenotype one looks at. Genes usually control the production of enzymes, which control steps in metabolic pathways. Many human metabolic diseases are due to homozygosity of an allele that produces a nonfunctioning enzyme. Nonallelic genes can interact in producing a phenotype so that alleles of one gene mask the expression of alleles of another gene. This process, termed epistasis, alters the expected phenotypic ratios. STUDY OBJECTIVE 4: To de ne how genes generally con-
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