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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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(a) Normal (sensitive) Paramecium. (b) Kappa-containing (killer) Paramecium. A Paramecium is about 200 m long. (Source: T. M. Sonneborn, gure 29.3,
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p. 373 in I. H. Herskowitz, Genetics, 2nd ed. [Boston: Little, Brown, 1965]. Reproduced by permission.)
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KK Killer
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plain that the virus plays an integral role. Whether the viral DNA or the kappa DNA codes the toxin is also not known at present.
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Kappa Conjugation with cytoplasmic exchange
Mate-Killer Infection and Mu Particles
Kappa is not the only infective agent known in Paramecium. Another agent is the mate-killer infection. Here again, killer cells have visible, bacterialike particles, called mu particles, in the cytoplasm. Preer and his colleagues have named them Caedobacter conjugatus. Mate-killers do not release a toxin into the environment, but instead kill their mates during conjugation. One of two unlinked dominant genes, M1 and M2, is required for the presence of mu particles. An interesting phenomenon occurs when a mate-killer becomes homozygous m1m1 m2m2 by autogamy. Although the offspring eventually lose their mu particles, virtually no loss of particles occurs until about the eighth generation, when some offspring lose all their mu. Up to this generation, all the cells maintain a full complement of mu. In the fteenth generation, only about 7% of the cells still have mu particles. This phenomenon is explained as the diluting out not of the mu themselves, but of a factor called metagon, which is necessary for the maintenance of mu in the cell. Once the cell becomes homozygous recessive, no further metagon production occurs. The verification that metagon is subsequently diluted out is evident in fteenth-generation cells that still have their mu. We would expect that after fission, one daughter cell would have a metagon and the other would not. What we expect, in fact, happens. The rate of dilution is consistent with an original number of about one thousand metagons per cell. The metagon appears to be
Kk All killer exconjugants
Autogamy
KK Killer
kk Sensitive
1:1 Figure 17.14 Autogamy in a heterozygous (Kk) killer Paramecium (formed by conjugation, with cytoplasmic exchange, of a KK killer and a kk sensitive cell). Upon autogamy, the heterozygote has a 50% chance of becoming a homozygous (KK) killer or a homozygous (kk) sensitive cell that loses its kappa particles.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Seventeen
Non-Mendelian Inheritance
Electron micrograph of a sectioned kappa particle (Caedobacter taeniospiralis). Phage particles appear as dark inclusions. The plane of the section cuts through a rolled-up R body. Magni cation 61,200 . (Reproduced by
permission of J. R. Preer, Jr.)
messenger RNA because it is destroyed by RNase. Its protein product is presently unknown. We thus see several instances of infective particles that interact with the Paramecium genome to produce interesting phenotypic results. Similar interactions are known in other organisms for example, the killer trait in yeast.
Electron micrograph of the spirochete associated with the extrachromosomal sex-ratio trait in Drosophila. Magni cation 22,700 .
(K. Oishi and D. F. Poulson, A virus associated with SR-spirochetes of Drosophila nebulosa, Proceedings of The National Academy of Sciences, USA, 67 [1970]:1565 72.
Drosophila
Several infective particles mimic patterns of inheritance in insects. In Drosophila, we nd forms of the sex-ratio phenotype in which females produce mostly, if not exclusively, daughters. One form is inherited as a chromosomal gene; another form, however, is not chromosomal. In the nonchromosomal form, females usually produce a few sons. These sons do not pass on the sex-ratio trait, but the daughters of sex-ratio females do. Because the trait persisted even after all the chromosomes had been substituted out of the stock by appropriate crosses, it was proven to be extrachromosomal. In addition, about half the eggs of a sex-ratio female fail to develop. Cytoplasm can be withdrawn from the undeveloped eggs and used to infect other females. The trait, then, is caused by some cytoplasmic factor that could infect other females and is not passed on by sperm. Detailed cytological examination of the cytoplasm of sexratio females has revealed a spirochete ( g. 17.16) that has been isolated and used to infect other female Drosophila with the sex-ratio trait; it is, therefore, the causal agent of this phenotype.
Reproduced by permission of the authors.)
in prokaryotes. The autonomous segments of DNA known as plasmids are, for the most part, known from bacteria, in which they occur as circles of DNA within the host cell (noncircular DNA is soon degraded). When plasmids become integrated into the chromosomes, they become indistinguishable from chromosomal material.
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