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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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Tracy M. Sonneborn (1905 1981). (Courtesy: Indiana
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protozoan familiar to most biologists. Ciliates have two types of nuclei: macronuclei and micronuclei. In Paramecium, there are two micronuclei, which are primarily reproductive nuclei, and one macronucleus, which is a polyploid nucleus concerned with the vegetative functions of the cell. During cell division, termed binary ssion, the micronuclei divide by mitosis and the macronucleus constricts and is pulled in half. Paramecium undergoes two types of nuclear rearrangements, during conjugation and autogamy. In conjugation, individuals of two mating types come to-
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gether and form a connecting bridge. The nuclear events are shown in gure 17.11. Brie y, the macronucleus of each cell disintegrates while the micronuclei undergo meiosis. Of the resulting eight micronuclei per cell, seven disintegrate and one remains; this one undergoes mitosis to form two haploid nuclei per cell. A reciprocal exchange of nuclei across the bridge then occurs. Each cell now has two haploid nuclei, one original and one migrant. The two nuclei fuse to form a diploid nucleus. The diploid nuclei in the two conjugating cells are genetically identical because of the reciprocity of the process. These nuclei then undergo two mitoses each to form four diploid nuclei per cell. Two nuclei become macronuclei, which separate at the next cell division; two remain as micronuclei that divide by mitosis at the next cell division. The two cells that separate are known as exconjugants. Depending primarily on the amount of time conjugating cells remain united, an exchange of cytoplasm may occur along with the exchange of nuclei. In the second type of process, autogamy, only one Paramecium is involved ( g. 17.12). The nuclear events are the same as in conjugation except that, at the point where a reciprocal exchange of nuclei would take place, the two haploid nuclei within the cell fuse. All cells after autogamy are homozygous.
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Figure 17.11 Conjugation in Paramecium. The letters K and k represent alleles of a gene in each micronucleus. When a KK and a kk individual conjugate, the exconjugants have the identical Kk genotype.
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Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Seventeen
Non-Mendelian Inheritance
Figure 17.12 Autogamy in Paramecium. The letters K and k represent alleles of a gene in each micronucleus. If a heterozygote undergoes autogamy, it becomes homozygous for one of the alleles (KK or kk).
Killer Paramecium and Kappa Particles
Sonneborn and his colleagues found that when certain stocks of Paramecium were mixed together, one stock had the ability to cause the individuals of the other stock to die. Those individuals causing death were called killers and those dying were referred to as sensitives. During conjugation, the sensitives are temporarily resistant to the killers. If cytoplasm is not exchanged during the conjugation, the exconjugants retain their original phenotypes so that killers stay killers and sensitives stay sensitives. When an exchange of cytoplasm occurs between sensitive and killer cells, both exconjugants are killers. The transfer of some cytoplasmic particle seems to be implied. Indeed, Sonneborn observed such particles in the cytoplasm of killers and called them kappa particles ( g. 17.13). Although the occurrence of killer Paramecium does not appear to involve chromosomal genes, Sonneborn reported one case in which exconjugant killer paramecia of hybrid origin underwent autogamy. He found that half of the resulting cells had no kappa particles and had become sensitives. He concluded that a gene is required for the presence of kappa particles, which has subsequently
been veri ed by numerous crosses. Figure 17.14 illustrates the sequence of genetic events that would produce a heterozygous killer Paramecium that, upon autogamy, would have a 50% chance of becoming sensitive. Although not yet cultured outside of a Paramecium, kappa is presumably a bacterium because it has many bacterial attributes including size, cell wall, presence of DNA, and presence of certain prokaryotic reactions ( g. 17.15). J. Preer and his colleagues, who studied kappa itself, named it Caedobacter taeniospiralis. Kappa occurs in at least two forms. The N form, the infective form that passes from one Paramecium to another, does not confer killer speci city on the host cell. The N form is attacked by bacteriophages that induce formation of inclusions, called R bodies, inside the kappa particle and thus convert it to the B form. These R bodies are visible under the light microscope as refractile bodies ( g. 17.15). In the B form, kappa can no longer replicate; it is often lysed within the cell. It confers killer speci city on the host cell, however. The sensitives are killed by the toxin paramecin, which is released by the killer Paramecium into the environment. Precisely what steps are involved in its formation are not known, although it is
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