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P Pelle Cactus Dorsal Dorsal Cytoplasm Nuclear membrane Nucleus Dorsal Cactus
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The signal transduction pathway. In this mechanism, the Sp tzle protein outside of a cell interacts with the Toll receptor protein, freeing the Dorsal protein to act as a transcription factor in the nucleus. When Toll binds Sp tzle, spanning the cell membrane, it changes the con guration of the interior domain of Toll, which then interacts with Pelle, causing it to phosphorylate the Cactus protein. Previously, Cactus had been bound to Dorsal, making Dorsal inactive; phosphorylation of Cactus releases it from Dorsal. Dorsal is then free to cross the nuclear membrane and act as a transcription factor.
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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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16. Gene Expression: Control in Eukaryotes
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target gene is interleukin-1, a protein in the immune system that induces fever. In the mammalian pathway, the signal protein is called Toll-like receptor-4, and the speci c transcription factor is called NF- B.
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Transposons
We have already shown that transposons can affect gene expression in prokaryotes, as, for example, in controlling the agellar phase in Salmonella (see chapter 14). Here we show how transposons can alter or regulate eukaryotic gene expression. Barbara McClintock discovered transposons in eukaryotes in the 1940s, without the aid of the tools of modern molecular genetics. She won the Nobel Prize for her work in 1983. She observed corn kernels that were streaked or spotted, indicating a high mutation rate. After careful genetic analysis, she showed that the mutability was due to transposons, which she called controlling elements.
The Ac-Ds System
The Ac-Ds system consists of two transposons. McClintock referred to the Ac (activator) transposon as an autonomous element and to the Ds (dissociation) transposon as a nonautonomous element. Ds cannot transpose until Ac enters the genome. At that time, Ds can transpose, be excised, or cause the chromosome it occurs on to break. Ds affects the phenotype by blocking expression of the genes it transposes into, as well as by causing the loss of alleles in acentric chromosomal fragments lost when Ds breaks its chromosome. In gure 16.4, we see three kinds of corn kernels: purple, bronze (light-colored), and bronze with purple spots. The purple kernels result from dominant function-
ing alleles that provide enzymes in the pathway for purple pigment. In the kernels that are bronze without spots, Ds elements have transposed into both copies of the Bz2 locus, disrupting the pigment pathway. Without the Ac element, the Ds elements remain in place, and the kernels are a uniform bronze color. In the bronze kernels with purple spots, the Ac element has entered the genome in the genetic cross. In the presence of Ac, Ds leaves its site in some of the cells, restoring activity to the Bz2 locus. This restored activity creates purple spots in those cells and in their progeny with the functioning Bz2 allele (see g. 14.34a). Ds and Ac elements have been cloned and sequenced. They are typical transposons that are very similar to each other. As might be expected, however, Ds has a deletion that prevents it from producing transposase. For Ds to transpose, Ac must provide the transposase. Ds apparently arose from Ac by deletion. It is interesting to note that one of Mendel s original seven characteristics of pea plants, wrinkled peas (rr: see g. 2.3), is caused by a transposon that inserts in the gene for Starch-branching enzyme I. When this gene is functional, the cells produce both branch-chained amylopectins and straight-chained amylose. If the gene fails to produce this enzyme, more sugar is present in the seeds, leading to greater osmotic pressure and, therefore, greater water content. More water is lost from these seeds upon maturation, resulting in greater shrinkage and wrinkling than in the wild-type seeds (RR and Rr). The transposon that disrupts this gene is about eight hundred base pairs long and is very similar to the Ds transposon in maize. The Ac-Ds system disrupts transcription through an invasive element that seems harmful (or at best neutral) to the organism. Mating-type control in yeast, by contrast, is a highly evolved system whose alternative expressions are advantageous to the organism.
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