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Suggested Readings for chapter 16 are on page B-15.
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III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
NON-MENDELIAN INHERITANCE
STUDY OBJECTIVES
1. To analyze the inheritance patterns of maternal effects
2. To analyze the patterns of cytoplasmic inheritance 3. To analyze the patterns of imprinting 524 511
STUDY OUTLINE
Determining Non-Mendelian Inheritance 509 Maternal Effects 509 Snail Coiling 509 Moth Pigmentation 510 Cytoplasmic Inheritance 511 Mitochondria 511 Chloroplasts 515 Infective Particles 518 Prokaryotic Plasmids 522 Imprinting 524 Summary 524 Solved Problems 525 Exercises and Problems 525 Critical Thinking Questions 527
Arti cially colored scanning electron micrograph of a mitochondrion in the cytoplasm of an intestinal epithelial cell. ( Professors P. Motta & T. Naguro/
SPL/Photo Researchers, Inc.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
17. Non Mendelian Inheritance
The McGraw Hill Companies, 2001
Maternal Effects
he phenotype can be controlled by chromosomal genes behaving according to Mendelian rules and the environment. In this chapter, we deal with another mode of inheritance, nonMendelian inheritance (also called extrachromosomal, cytoplasmic, and nonchromosomal inheritance; maternal effects; or imprinting). Maternal effects are the in uences of a mother s genotype on the phenotype of her offspring; examples include snail coiling and moth pigmentation (we started a discussion of maternal effects in chapter 16, when we looked at development in Drosophila). Cytoplasmic inheritance is controlled by nonnuclear genomes found in chloroplasts, mitochondria, infective agents, and plasmids. And imprinting is a process in which gene expression depends on the parent from which the gene came. None of these modes of inheritance follow the usual Mendelian rules and ratios. Maternal effects result from the asymmetric contribution of the female parent to the development of zygotes. Although both male and female parents contribute equally to the zygote in terms of chromosomal genes (with the exception of sex chromosomes), the sperm rarely contributes anything to development other than chromosomes. The female parent usually contributes the zygote s initial cytoplasm and organelles. Zygotic development, therefore, usually begins within a maternal milieu, so that the maternal cytoplasm directly affects zygotic development (see chapter 16). Cytoplasmic inheritance refers to the inheritance pattern of organelles and parasitic or symbiotic particles that have their own genetic material. Chloroplasts, mitochondria, bacteria, viruses, and, of course, plasmids all have their own genetic material. These genomes are open to mutation. As we shall see, their inheritance pattern does not follow Mendel s rules for chromosomal genes. Imprinting occurs in more than twenty genes and is responsible for several human diseases.
DETERMINING NONM E N D E L I A N I N H E R I TA N C E
How does one determine that a trait is inherited The question does not have as obvious an answer as we might expect. Environmentally induced traits can mimic inherited phenotypes, as with the phenocopies we discussed in chapter 5. For example, the inheritance of vitamin D-resistant rickets is mimicked by lack of vitamin D in the diet. It is possible to determine that the rickets is not inherited by simply administering adequate quantities of vitamin D. Inherited rickets does not respond to vitamin D until about 150 times the normally adequate amount is administered.
Some environmentally induced traits persist for several generations. For example, a particular Drosophila strain that normally grows at 21 C was exposed to 36 C for twenty-two hours. Dwarf progeny were produced. When they were mated among themselves, fewer and fewer dwarfs appeared in each generation, but smallerthan-normal ies were produced as late as the fth generation. The appearance of an environmentally induced trait that persists for several generations has been termed dauermodi cation. Extrachromosomal inheritance is usually identi ed by the odd results of reciprocal crosses. If the progeny of reciprocal crosses are not followed for several generations, the results can be misleading when extrachromosomal inheritance is involved. Where feasible, nuclear transplantation has proved useful in identifying extrachromosomal inheritance. In this technique, the nucleus of a cell, such as an amoeba or frog egg, is removed by microsurgery or destroyed by radiation, and another nucleus substituted. Thus, not only can a nucleus be isolated from its cytoplasm, but various nuclei can be implanted in the same cytoplasm. A similar experiment, called a heterokaryon test, can be done with various fungi such as Neurospora and Aspergillus: Mycelia can fuse and form a heterokaryon, a cell containing nuclei from different strains. Thus, nuclei of both strains exist in the mixed cytoplasm. Subsequently, spores (conidia) that have one or the other nucleus in the mixed cytoplasm can be isolated. The phenotype of the colonies produced from these isolated conidia show whether the trait under observation is controlled by the nucleus or the cytoplasm. Chromosomal genes in a particular cytoplasm can also be isolated by repeated backcrossing of offspring with the male-parent type. In each cross, the content of the female chromosomal genes is halved, but, presumably, the cytoplasm remains similar to the female line. Thus, after several generations, male genes can be isolated in female cytoplasm. The phenotypic results of the nal cross will indicate whether inheritance was chromosomal or extrachromosomal.
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