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Figure 16.4 The Ac-Ds mutability system in corn. Shown is an ear of corn with purple and bronze kernels. The purple kernels have no transposons. The bronze kernels (light-colored) lack the purple pigment because they have a Ds element in both copies of the Bz2 locus, disrupting pigment production. Without an Ac element present, the kernel remains bronze. In the presence of the Ac element, the Ds element can leave its position, restoring the allele and producing a purple spot in a bronze kernel. Spots differ in size based on when the Ds element was excised during the development of the kernel: early excision yields large spots; late yields small spots. (Corn ear courtesy of Dr. Neelima Sinha; Photo by the author.)
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16. Gene Expression: Control in Eukaryotes
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Transposons determine the mating type in yeast. Haploid yeast cells exist in one of two mating types, a and , determined by the MATa and MAT alleles. Homothallic strains of yeast switch mating types, as often as every generation. (The term homothallic, a misnomer, means that every cell is alike each can mate with any other. The term was applied before scientists realized that the cells could change mating types.) Homothallism is determined by the dominant HO allele that codes for an endonuclease that initiates transposition. Strains that do not change mating type are heterothallic, determined by the recessive ho allele; no active endonuclease is present to allow transposition, and thus they undergo no change in mating type. The ability to switch mating types in a single cell implies that both forms of the mating-type gene are present in each cell. In 1971, Y. Oshima and I. Takano proposed that mating type was controlled by a transpositional event, similar to the Ac-Ds system in corn or the agellar phase in Salmonella. Later genetic and recombinant DNA studies revealed the exact mechanism. The third chromosome in yeast contains the matingtype locus (MAT). Silent (unexpressed) copies of the mating-type alleles are found on the left and right arms of the same chromosome ( g. 16.5). HML contains the silent allele and HMR contains the silent a allele. In transposition, a copy of one or the other (HMR or HML) moves to the MAT site, replacing whatever allele was there to begin with. This mechanism has been called a cassette
mechanism. The MAT site is analogous to a cassette player, with HMR and HML similar to cassette tapes. Transposition brings a new tape to the cassette player. MATa and MAT each begin a genetic cascade that activates certain genes and represses others. For example, MAT codes for two proteins. The MAT 1 protein activates the transcription of an -factor (a pheromone) gene and an a-factor (pheromone) receptor gene. (Pheromones are chemical signals, analogous to hormones, that convey information between individuals.) The MAT 2 protein represses the a-speci c genes. Conjugation requires the emission of one type of pheromone and the reception of the other type: An cell emits factor and is receptive to a factor; an a cell emits a factor and is receptive to factor. In summary, then, transposons can affect eukaryotic gene expression. However, with the exception of a few systems such as mating-type determination in yeast, transposons appear to have a random, disruptive effect on developmental processes.
PAT T E R N S I N D E V E L O P M E N T
Development is the orderly sequence of change that produces increasing complexity during the growth of an organism; it is controlled by the differential expression of genes. A central problem of development is explaining genomic equivalence, how cells with identical genetic material can give rise to different cell types. A favored approach to understanding the genetic control of development in higher organisms requires rst learning the details of the normal developmental process in an organism and then studying the disruption of this normal process by mutation and experimental manipulation. At one point, scientists believed that development might take place through permanent changes in chromosomes. The idea was that subtle changes might occur in chromosomes during development; these changes would not be observable by karyotyping a cell. Geneticists have explored this hypothesis by several methods. However, the cloning of a mammal, such as the sheep Dolly (see chapter 13), from the cell of an adult demonstrates that adult nuclei are totipotent: Any adult nucleus can give rise to the whole organism and all its cellular types, indicating the chromosomes are intact.
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