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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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17. Non Mendelian Inheritance
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The McGraw Hill Companies, 2001
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Cytoplasmic Inheritance
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Among the mitochondrial DNAs that have been sequenced from different organisms, we see great variation in content and organization. Yeast mitochondrial DNA, for example, is not as economical as human mitochondrial DNA. Yeast mitochondrial DNA, about five times larger than human mitochondrial DNA, has noncoding regions as well as introns. Because mitochondria are similar in structure and biochemistry to prokaryotic cells, given the general lack of introns in
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prokaryotic genes, it was surprising to nd introns in yeast mitochondrial DNA. These genes most probably arose later as nuclear genes that were then captured by the mitochondria, possibly by recombination with nuclear DNA. Of the many mitochondria sequenced to date (about 175 at the beginning of 2001), the sizes range from less than 6 to more than 200 kilobases and from 3 to 97 genes. With this wide range of genes present, the only generality we can make about mitochondrial DNA is that the large and small segments of the mitochondrial ribosomal RNA, as well as most of the mitochondria s transfer RNAs, are usually coded by the mitochondria s own genome, as are several proteins in respiratory complexes III and IV (cytochrome c oxidase and cytochrome c oxidoreductase). Once the interaction within the mitochondrial-nuclear genetic system is clearly understood, we might expect to see several different inheritance patterns following either cytoplasmic or nuclear lines for the genetic defects that lead to interruption of cellular respiration. Among the beststudied phenotypes with such inheritance patterns are the petite mutations of yeast.
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The amino acid sequence of mouse dihydrofolate reductase. Numbers refer to sequential amino acids. The rst eighty- ve amino acids serve as the signal sequence for transport into mitochondria. Five -helical regions exist in the protein (A E). Positively and negatively charged amino acids are marked with ( ) and ( ) signs. (Reprinted by permission from E. C. Hurt and G. Schatz, A cytosolic
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protein contains a cryptic mitochondrial targeting signal, Nature, Volume 325, p. 499, 1987. Copyright 1987 Macmillan Magazines, Ltd.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Seventeen
Non-Mendelian Inheritance
Petites
Under aerobic conditions, yeast grows with a distinctive colony morphology. Under anaerobic conditions, the colonies are smaller, and the structures of the mitochondria are reduced. Occasionally, when growing aerobically, small, anaerobiclike colonies appear; but in these colonies, the mitochondria appear perfectly normal. These colonies are caused by petite mutations. When petites are crossed with the wild-type, three modes of inheritance emerge ( g. 17.6). The segregational petite, caused by mutation of a chromosomal gene, exhibits Mendelian inheritance. The neutral petite is lost immediately upon crossing to the wild-type. The suppressive petite shows variability in expression from one strain to the next but is able to convert the wild-type mitochondria to the petite form. All petites represent failures of mitochondrial function, whether the function is controlled by the mitochondria themselves or by the cell s nucleus; they usually lack one or another cytochrome. Although the mechanisms that produce neutral and suppressive petites are not known with certainty, their DNA has supplied some interesting information. In some petites, no change in the buoyant density of the DNA is found. (Buoyant density, a term that describes the position at which the DNA equilibrates during densitygradient centrifugation, is a measure of the composition of the molecule; see chapter 15.) In other petites, changes
in buoyant density range from very small to the complete absence of DNA. Petites, therefore, can be the result of an approximation to a point mutation (with no measurable change in the buoyant density of the DNA), marked changes in the DNA, or the total absence of DNA. In most petites, protein synthesis within the mitochondrion is lacking. Any and all of these changes produce the petite (anaerobiclike) phenotype. Neutral petites seem to have mitochondria that entirely lack DNA. When neutral petites are crossed with the wild-type to form diploid cells, the normal mitochondria dominate. During meiosis, virtually every spore receives large numbers of normal mitochondria; the progeny are, therefore, all normal. Suppressive petites could exert their in uence over normal mitochondria in one of two ways. The suppressive mitochondria might simply out-compete the normal mitochondria and take over; they might simply reproduce faster within a cell. Alternatively, crossing over between the DNA of the suppressive petite and the wildtype might affect the normal DNA if the suppressive petite s DNA were severely damaged. Presumably, recombination in mitochondrial DNAs occurs when two or more mitochondria fuse, bringing the two different sets of DNA in contact within the same organelle. Recombination would presumably take place by normal crossover mechanisms. If large portions of the DNA from the suppressive mitochondria were missing or altered, recombination with the normal mitochondria s DNA might exchange some of this damaged DNA. Several experiments have crossed a suppressive petite and a wild-type, each with mitochondrial DNA of known buoyant density. The DNAs of the offspring colonies, which were petites, were of various buoyant densities. For example, when a normal strain with mitochondrial DNA with a buoyant density of 1.684 g/cm3 was crossed with a suppressive petite with a buoyant density of 1.677 g/cm3, the offspring colonies mitochondrial DNA had buoyant densities of 1.671, 1.674, and 1.683 g/cm3. Such information supports the notion that the suppressive character takes over a colony by way of recombination.
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