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Replication
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The genetic material must be capable of precisely directing its own replication so that every daughter cell receives an exact copy. Some mutability, or the ability to change, is also required, because we know that the genetic material has changed, or evolved, over the history of life on earth. In their 1953 paper, Watson and Crick had already worked out the replication process based on the structure of DNA. The delity of the replication process is so great that the error rate is only about one in a billion.
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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9. Chemistry of the Gene1
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Chemistry of the Gene
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Structure of compound CH3 CH OH CHCOOH NH2
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Name of enzyme
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Name of compound Threonine
Enzyme
Threonine dehydratase NH3 CH3 CH2 Pyruvate Acetolactate synthase CO2 C2H5 CH3 C O C OH Acetolactate mutase NAD(P)H Reductase NAD(P)+ CH3 H CH3 CH2 C OH C OH Dihydroxyacid dehydratase H2O CH3 CH3 CH2 C H Glutamate -ketoglutarate CH3 CH3 CH2 CH H C NH2 Figure 9.2 COOH Isoleucine Valine transaminase C O COOH -keto- -methylvaleric acid Glucose-6-phosphate COOH , -dihydroxy -methylvaleric acid Enzyme-substrate complex ADP Enzyme COOH -aceto- -hydroxybutyric acid C O COOH -ketobutyric acid Glucose Enzyme
The active site of an enzyme recognizes a speci c substance. In this case, ATP plus glucose is converted into ADP and glucose-6-phosphate by the enzyme hexokinase. The active site is diagrammed in red. The terminal phosphate group of ATP is tan.
Metabolic pathway of conversion of the amino acid threonine into isoleucine.
Location
It has been known since the turn of the century that genes, the discrete functional units of genetic material, are located in chromosomes within the nuclei of eukaryotic cells: the way chromosomes behave during the cel-
lular division stages of mitosis and meiosis mimics the behavior of genes. Thus, the genetic material in eukaryotes must be a part of the chromosomes. For a long time, proteins were considered the most probable genetic material because they have the neces-
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
9. Chemistry of the Gene1
The McGraw Hill Companies, 2001
In Search of the Genetic Material
Oswald T. Avery (1877 1955).
(Courtesy of the National Academy of Sciences.)
sary molecular complexity. The twenty naturally occurring amino acids can be combined in an almost unlimited variety, creating thousands and thousands of different proteins. The rst proof that the genetic material is deoxyribonucleic acid (DNA) came in 1944 from Oswald Avery and his colleagues. The Watson and Crick model in 1953 ended a period when many thought DNA was the genetic material, but its structure was unknown.
Petri plate with smooth and rough colonies of Streptococcus pneumoniae. R (rough) strain colonies appear on the left and S (smooth) colonies on the right on the same agar. Magni cation 3.5 . (O. T. Avery, C. M. Macleod, and
M. McCarty, Studies on the chemical nature of the substance inducing transformation of pneumococcal types. Reproduced from the Journal of Experimental Medicine 79 (1944):137 58, g. 1 by copyright permission of the Rockefeller University Press. Reproduced by permission. Photograph made by Mr. Joseph B. Haulenbeek.)
Evidence for DNA as the Genetic Material
Transformation
In 1928, F. Grif th reported that heat-killed bacteria of one type could transform living bacteria of a different type. Grif th demonstrated this transformation using two strains of the bacterium Streptococcus pneumoniae. One strain (S) produced smooth colonies on media in a petri plate because the cells had polysaccharide capsules. It caused a fatal bacteremia (bacterial infection) in mice. Another strain (R), which lacked polysaccharide capsules, produced rough colonies on petri plates ( g. 9.4); it did not have a pathological effect on mice. Bacteria of the rough strain are engulfed by the mice s white blood cells; bacteria of the virulent smooth strain survive because their polysaccharide coating protects them. Grif th found that neither heat-killed S-type nor live R-type cells, by themselves, caused bacteremia in mice. However, if he injected a mixture of live R-type and heatkilled S-type cells into mice, the mice developed a bacteremia identical to that caused by living S-type cells ( g. 9.5). Thus, something in the heat-killed S cells transformed the R-type bacteria into S-type cells. In 1944, Oswald Avery and two of his associates, C. MacLeod and M. McCarty, reported the nature of the transforming substance. Avery and his colleagues did their work in vitro (literally, in glass), using colony morphology on culture media rather than bacteremia in mice as evidence of transformation. They ruled out proteins, carbohydrates, and lipids by their extraction procedure, by the chemical analysis of the transforming material, and by demonstrating that the only enzymes that destroyed the transforming ability were enzymes that de-
stroyed DNA. This study provided the rst experimental evidence that DNA was the genetic material: DNA transformed R-type bacteria into S-type bacteria.
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