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Human Mitochondrial Inheritance
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Petite yeasts categorized on the basis of segregation patterns. Three types of petites are recognized (segregational, neutral, and suppressive), depending on the meiotic segregation pattern of petite wild-type diploids. Segregational petite heterozygotes segregate a 1:1 ratio of spores; neutral petites are lost when heterozygous; and suppressive petites act in a dominant fashion under the same circumstances.
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In human beings, certain diseases trace their dysfunction to mitochondrial pathologies. The rst such disease, Luft disease, characterized by excessive sweating and general weakness, was reported in 1962. In 1988, Douglas Wallace and his colleagues showed that Leber optic atrophy is a cytoplasmically inherited disease. This disease causes blindness, with a median age of onset of twenty to twenty-four years. The onset age and phenotype are variable, depending on the degree of heteroplasmy in the in-
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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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Cytoplasmic Inheritance
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dividual. Apparently, defects in mitochondria are not tolerable in the optic nerve, which demands a great deal of energy. The disease also does some damage to the heart. Pedigrees showed that Leber optic atrophy is transmitted only maternally. Sequencing of mitochondrial DNAs in affected families pinned down the disease to a point mutation, a change in nucleotide 11,778, which is in the gene for NADH dehydrogenase subunit 4 (see g. 17.4). A guanine is changed to an adenine at codon 340, which converts an arginine to a histidine. This is the rst human disease traced to a speci c mitochondrial DNA mutation. Since 1962, over one hundred diseases, including some of the general symptoms of aging and cancer, have been attributed to mitochondrial pathology.
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Antibiotic In uences
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Since the machinery of mitochondrial protein synthesis is prokaryotic in nature, antibiotics such as chloramphenicol and erythromycin can inhibit it. These antibiotics elicit a petite-type growth response in yeast. Antibiotic-resistant strains can be obtained by growing yeast on the antibiotic; only resistant mutants will grow. The resistance appears to be inherited in the mitochondrial, not the cellular, DNA. A mitochondrial inheritance pattern results, with crosses between a resistant and a sensitive (wild-type) yeast, as shown in gure 17.7. The resulting diploid colonies segregate both resistant and sensitive cells. Although not expected on the basis of a chromosomal gene, the random sorting of mitochondria through cell division could result in a wild-type cell containing only sensitive mitochondria. Since some yeast have only one to ten mitochondria per cell, this random assortment of sensitive mitochondria can be expected to occur at a relatively high rate.
Inheritance of antibiotic (chloramphenicol) resistance in yeast. Resistant and sensitive cells are produced by a diploid cell that resulted from a cross of resistant and sensitive haploids. The segregation is not in a simple Mendelian ratio, but depends on the random assortment of mitochondria. Sensitive cells have no resistant mitochondria. Resistant cells have resistant mitochondria.
Chloroplasts
The chloroplast is the chlorophyll-containing organelle that carries out photosynthesis and starch-grain formation in plants ( g. 17.8). Chloroplasts are referred to as plastids before chlorophyll develops. However, when grown in the dark (and under some other circumstances), plastids do not develop into chloroplasts, but remain reduced in size and complexity. These undeveloped plastids, referred to as proplastids, are each about the size and shape of a mitochondrion. Like mitochondria, chloroplasts contain DNA and ribosomes, both with prokaryotic af nities. The DNA of chloroplasts (cpDNA) is a circle that ranges in size from 85 kilobases (kb) in the green alga Codium to as large as 2,000 kilobases in the green alga Acetabularia. Thus, chloroplast DNA is minimally about ve times the size of an animal mitochondrial DNA. The chloroplast DNA, like mitochondrial DNA, controls the production of transfer RNAs, ribosomal RNAs, and some of the proteins found within the organelle. From the more than nineteen chloroplast DNAs that have been sequenced, there seem to be about one hundred genes in the chloroplast genome. About thirty code for the subunits of the ve photosynthetic protein complexes: photosystem I, photosystem II, ribulose bisphosphate carboxylase-oxygenase, cytochrome b6-f complex, and ATP synthase. About sixty genes code for the protein synthesis apparatus of the chloroplast. Scientists believe that the chloroplast evolved from symbiotic cyanobacteria (blue-green algae), which have many af nities with the chloroplast: The ribosomal RNA of cyanobacteria will hybridize with the DNA of chloroplasts. The similarities between mitochondria and chloroplasts make it possible to predict the inheritance patterns of chloroplast mutations on the basis of existing
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