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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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Inheritance pattern of larval and adult pigmentation in the our moth, Ephestia k hniella. A single locus controls the presence (a ) or absence (a) of kynurenin. In the cross on the left, the mother is aa (nonpigmented). Her aa offspring, in both the larval and adult stages, are also nonpigmented. In the reciprocal cross (right), the mother has the a a genotype and is pigmented. Her aa offspring are nonpigmented as adults but are pigmented as larvae because of residual kynurenin from the egg, which eventually dilutes out.
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Mitochondria
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The mitochondrion is an organelle in eukaryotic cells in which the electron transport chain takes place.The actual number of mitochondria per cell can be determined by serial sectioning of whole cells and examination under the electron microscope. This is a tedious and dif cult procedure. Estimates range between ten and ten thousand per cell, depending on the organism and cell type. As far as we are concerned, the most interesting aspect of the mitochondrion is that it has its own DNA. In most animal cells, the mitochondrial DNA (mtDNA) is a circle of about sixteen thousand base pairs ( g. 17.3). However, some organisms (yeast, higher plants) have mitochondrial DNAs ve to twenty- ve or more times larger than in animals. And some organisms have linear mitochondrial chromosomes. Two general patterns are found in mitochondrial inheritance in animals. First, the mitochondria are generally inherited in a maternal fashion; that is, the male gamete usually does not contribute mitochondria to the zygote. However, a small amount of leakiness occurs in this process. For example, it has recently been shown that about one mitochondrion per thousand is of paternal origin in mice. In some species, such as mussels, it appears that mitochondrial inheritance is biparental. That is, the population of mitochondria in an offspring derives almost equally from the male and female parent. In some gymnosperm plants, such as coastal redwoods, mitochondria are inherited paternally only paternal mitochondria are passed into the zygote. However, these are all exceptions to the general rule of maternal inheritance of mitochondria. The second general pattern of mitochondrial inheritance is homoplasmy, the existence of a uniform population of mitochondria within an organism. In general, all
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the mitochondria within an individual are genetically identical. Certainly, biparental inheritance and leakiness of paternal mitochondria violate that principle, resulting in heteroplasmy, a heterogeneity of mitochondria within a cell or organism.
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Electron micrograph of the circular DNA from within a mouse cell mitochondrion. Magni cation 48,000 .
(M. M. K. Nass, The circularity of mitochondrial DNA, Proceedings of the National Academy of Sciences, USA, 56 (1966):1215 22. Reproduced by permission of the author.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Seventeen
Non-Mendelian Inheritance
Mitochondrial Genomes
Numerous mitochondrial DNAs have been sequenced, including the human mitochondrial DNA, which is 16,569 base pairs long. It is a model of economy, with very few noncoding regions and no introns ( g. 17.4). Each strand of the duplex is transcribed into a single RNA product that is then cut into smaller pieces, primarily by freeing the twenty-two transfer RNAs interspersed throughout the genome. Also formed are a 16S and a 12S ribosomal RNA. Although proteins and small molecules such as ATP and tRNAs can move in and out of the mitochondrion, large RNAs cannot. Thus, the mitochondrion must be relatively self-suf cient in terms of the RNAs needed for protein synthesis.We previously discussed mitochondrial protein synthesis when we looked at unique attributes of the mitochondrial genetic code in chapter 11. Oxidative phosphorylation, the process that occurs within the mitochondrion, requires at least sixty-nine polypeptides. The human mitochondrion has the genes for thirteen of these: cytochrome b, two subunits of ATPase, three subunits of cytochrome-c oxidase, and seven subunits of NADH dehydrogenase. The remaining polypeptides needed for oxidative phosphorylation are transported into the mitochondrion; they are synthesized in the cytoplasm under the control of nuclear genes. Proteins targeted for entry into the mitochondrion have special signal sequences (see chapter 11).
The signal sequences range up to eighty- ve amino acids long. Signal sequences examined so far do not have consensus amino acids but do have certain attributes ( g. 17.5), including a somewhat regular alternation of basic (positively charged) and hydrophobic (negatively charged) residues. In addition, they form helices with opposite hydrophobic and hydrophilic faces that must somehow be important in the protein s ability to enter the mitochondrion. When a signal sequence (such as that in g. 17.5) is attached to nonmitochondrial proteins by DNA manipulations, those proteins are transported into the mitochondrion. The mitochondrial ribosomal RNA is more similar to prokaryotic ribosomal RNA than to eukaryotic ribosomal RNA. The mitochondrial ribosome, although constructed of imported cellular proteins, is sensitive to prokaryotic antibiotics; for example, streptomycin and chloramphenicol inhibit their function. This af nity (close resemblance) between mitochondria and prokaryotes is strong support for the symbiotic origin of mitochondria. That is, we now accept the model advocated by L. Margulis that organelles such as mitochondria and chloroplasts were originally free-living bacteria and cyanobacteria, respectively. These prokaryotes invaded or were eaten by early cells and, over evolutionary time, became the organelles we see today. Since they arose as prokaryotes, these organelles retain certain evolutionary similarities to other prokaryotes.
Gene map of the human mitochondrial chromosome. All but nine loci are on the heavy (H) strand. The light-strand (L) loci are labeled inside the circle; the H-strand loci are labeled on the outside. Also shown are the origins of Hand L-strand replication and the directions of transcription. The twenty-two tRNA genes are colored red. NADH refers to NADH dehydrogenase (subunits 1 4, 4L, 5, and 6); CCO refers to cytochrome-c oxidase (subunits I III). (Source: Data from
V. McKusick, Mendelian Inheritance in Man, 7th edition, 1986.)
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