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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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Non-Mendelian Inheritance
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Electron micrograph of lettuce chloroplasts. The chloroplast consists of an outer membrane, stacks of grana, lamellae, and stroma. Magni cation 3,570 . ( Dr. J.
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knowledge of mitochondrial genetics: We should nd both chromosomal and plastid mutants of chloroplast functions. Simple segregation should occur in the chromosomal mutations, and cytoplasmic patterns of inheritance should occur with the chloroplast DNA mutations. Investigation of these inheritance patterns is complicated by the fact that plant cells have both mitochondria and chloroplasts. Since both have prokaryotic af nities, it is sometimes dif cult to determine whether a genetic trait is due to a defect in the genetic system of the chloroplast or the mitochondrion. Like mitochondria, chloroplasts generally show homoplasmy and maternal inheritance, although, as in mitochondria, there are exceptions. For example, gymnosperms usually have paternal inheritance of chloroplasts. Lesions in the photosystems of the chloroplast result in proplastid formation, with a loss of green color. When proplastid formation occurs in a particular tissue of a plant, variegation results.That is, there are both green and white parts, often as stripes. Some interesting genetic studies have focused on the inheritance of variegation, especially in the interaction of chloroplast and chromosomal genes.
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Marcus M. Rhoades (1903 1991). (Courtesy of
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Zea mays
M. Rhoades worked on the variegation in corn (Zea mays) controlled by the iojap chromosomal locus, which, when homozygous, prevents proplastids from developing into chloroplasts and thus results in variegation. The iojap-affected plastids do not contain ribosomes or ribosomal RNA; they therefore lack protein synthesis.
The interaction of chromosomal and extrachromosomal inheritance is shown in the reciprocal crosses depicted in gure 17.9. One cross produces results exactly as would be predicted on the basis of simple Mendelian inheritance, with the homozygous recessive genotype (ijij) inducing variegation. When the reciprocal cross is carried out, blotch variegation is seen in both the F1 and F2 that carry the dominant Ij allele. This inheritance pattern is caused by the fact that the pollen grain in corn does not carry any chloroplasts, whereas the ovule does. Thus, the rst cross in gure 17.9 deals with the passage of normal chloroplasts only into the F2 generation. In the F2, the ijij genotype then induces variegation. The chloroplasts of the pollen parent are unimportant because they do not enter the F1. In the reciprocal cross, however, because the stigma parent is variegated, the F1 is heterozygous but carries proplastids from the ovule that remain proplastids even under the dominant normal (Ij) allele. Therefore, regions of colorless cells produce white spots (blotchy variegation). Once the ij allele induces chloroplasts to become pro-
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Cytoplasmic Inheritance
Reciprocal crosses involving the chromosomal gene iojap in corn. The homozygous recessive condition (ijij) induces variegation (representative corn leaves are shown). (Blotch variegation consists of irregularly shaped white areas rather than striping.) However, plants with the dominant allele (IjIj, Ijij) can still be variegated if their mothers were variegated, since mothers pass on their chloroplasts to their offspring; males (pollen parents) do not pass on their chloroplasts. Iojap homozygotes induce variegation. The defective chloroplasts are then inherited in a cytoplasmic fashion.
plastids, they do not revert to normal type even under the Ij allele. Thus, we see the interaction of a chromosomal gene and the chloroplast itself, which inherits a changed condition. There is some evidence that iojap may suppress the chloroplast rather than cause a mutation of some function. There are loci in corn and in other species that can induce back mutation in the chloroplasts. Removal of suppression rather than an actual reversion is more likely to occur because the reversion rate is too high to be due to simple back mutation.
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