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Chlamydomonas
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The single-celled green alga, Chlamydomonas reinhardi, has been used in the study of extrachromosomal inheritance for several reasons. First, it has a single, large chloroplast; it can survive by culture technique even when the chloroplast is not functioning; and nally, it shows some interesting non-Mendelian inheritance patterns related to mating type. R. Sager has done extensive work on the inheritance of streptomycin resistance in Chlamydomonas. Streptomycin resistance can be selected for in Chlamydomonas in several ways. Normal cells, sensitive to the antibiotic, are killed in its presence. If cells are grown in low levels of the antibiotic (100 g/ml), some cells show resistance to it. When these cells are crossed
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Ruth Sager (1918 1997).
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The rst work with corn variegation was done by Carl Correns, one of Mendel s rediscoverers. Correns also found maternal inheritance of variegation in the fouro clock plant, Mirabilis jalapa. He could predict color and variegation of offspring solely on the basis of the region of the plant on which the stigma parent was located. A ower from a white sector, when pollinated by any pollen, would produce white plants; a ower on a green sector or a variegated sector produced green or variegated plants, respectively, when pollinated by pollen from any region of a plant. We thus see the simple maternal nature of the inheritance of the variegation. A chromosomal gene, like iojap, induces variegation. Inheritance of this induced variegation follows the maternal pattern of chloroplast inheritance.
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III. Molecular Genetics
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17. Non Mendelian Inheritance
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Seventeen
Non-Mendelian Inheritance
with the wild-type, the resistance segregates in a 1:1 ratio, indicating that streptomycin resistance is controlled by a chromosomal locus.The same experiment can be repeated using high levels of the antibiotic in the medium (500 1,600 g/ml). Again, resistant colonies grow. If they are crossed with the wild-type, a 1:1 ratio does not ensue. Chlamydomonas does not have sexes but does have mating types mt and mt . Only individuals of opposite type can mate. Mating type is inherited as a single locus with two alleles. When two haploid cells of opposite mating type fuse, they form a diploid zygote, which then undergoes meiosis to produce four haploid cells, two of mt and two of mt .The high-level resistance always segregates with the mt parent ( g. 17.10). It is as if the mt parent were contributing the cytoplasm to the zygote in a manner similar to maternal plastid inheritance in plants. The mt parent acts like a pollen parent by making a chromosomal contribution but not a cytoplasmic one. The mechanism of the extrachromosomal inheritance pattern of Chlamydomonas is the preferential di-
gestion of the DNA of the chloroplast from the mt parent. Currently, we believe that streptomycin s target is the chloroplast. More recent work has shown that the mt inheritance is only 99.98% effective that is, 0.02% of the offspring in crosses of the type shown in gure 17.10 have the streptomycin phenotype of the mt parent.Thus, we have the possibility of studying recombination in chloroplast genes. Although most of the evidence is only indirect and plagued by the previously mentioned problems of separating chloroplast and mitochondrial effects, some initial mapping studies have been done.
Infective Particles
Paramecium
Tracy Sonneborn discovered the killer trait in Paramecium. Before analyzing this trait, we must digress a moment to look at the life cycle of Paramecium, a ciliated
Figure 17.10 Inheritance pattern of streptomycin resistance in Chlamydomonas is dependent on the genotype of the mt parent. (The n and 2n refer to the ploidy of the cells.) If the mt parent is streptomycin resistant (red), then the diploid heterozygote, as well as the meiotic products, will be streptomycin resistant. If, however, the mt parent is streptomycin sensitive (green), the diploid heterozygote, as well as the meiotic products, will be streptomycin sensitive.
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