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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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The McGraw Hill Companies, 2001
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Gene Expression: Control in Eukaryotes
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Segmentation genes of Drosophila fall into three categories: gap, pair-rule, and segment polarity. To the left of each pair is the wild-type larva with cuticular pattern that indicates segment position; to the right is the mutant larva. An example of a gap mutation is Kr ppel, which eliminates the three thoracic and ve of the eight abdominal segments (shaded in the wild-type larva). Pair-rule genes are shown that eliminate even (even-skipped ) or odd (odd-skipped ) segments (counting from the abdominal segments). An example of a segment polarity gene is gooseberry, in which the posterior portion of each segment behaves like a mirror image of the anterior portion of the segment. (Reprinted with permission from Christiane N ssleinFigure 16.14
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Volhard and Eric Wieschaus, Mutations affecting segment number and polarity in Drosophila, Nature, 287:795 80, 1980. Copyright 1980 Macmillan Magazines, Ltd., London, England.)
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Presumably, as more Bicoid is present, it binds to more of the hunchback promoter sites, resulting in greater transcriptional activity. At least three gap genes are controlled by the concentrations of the speci c transcription factor hunchback: Kr ppel, knirps, and giant. In response to the Hunchback gradient, these three genes are expressed in discrete stripes in the embryo ( g. 16.15). Both anterior and posterior edges of the Kr ppel stripe are controlled by Hunchback concentration; Hunchback concentration also controls the anterior edges of the Knirps and Giant stripes. The posterior edges of the Knirps and Giant stripes are controlled by the gradient of the Tailless protein, which is controlled in turn by the terminal maternal-effect gene, torso ( g. 16.15). We know the distributions of these proteins by antibody studies, and we know the limits of the protein distributions from studies of various mutants that lack the clear edges of the stripes. For example, the borders of the Kr ppel stripe are changed in hunchback mutants in accordance with the number of copies of the genes. We have thus gone from very broad and fuzzy regions of maternal-effect gene products to more de ned bands of gap gene products. Interaction of the gap gene proteins then controls transcription of the pair-rule genes (see g. 16.14). These genes affect alternate sets of segments, even and odd. For example, mutants of the even-skipped gene cause the loss of the even-numbered segments, counting by the abdominal segments (loss of two thoracic segments as well
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Kruppel knirps
giant
Posterior
Concentration
Hunchback protein
Tailless protein
Three discrete bands of gene expression (Kr ppel, knirps, and giant) in the developing Drosophila embryo. These bands come about because of the gradients of Hunchback and Tailless proteins. The Hunchback protein level controls the anterior edge of gene expression of Kr ppel, knirps, and giant, as well as the posterior edge of the Kr ppel gene expression. The Tailless protein level controls the posterior end of knirps and giant gene expression. The nature of these border edges is veri ed in mutations of the hunchback and tailless genes that result in different limits. The three genes (Kr ppel, knirps, and giant) are transcription factors, further controlling gene expression in these regions of the embryo.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Patterns in Development
as abdominal segments 2, 4, 6, and 8). Finally, the segmentpolarity genes are controlled by the pair-rule genes, resulting in genes that affect all segments (see g. 16.14). For example, mutants of the gooseberry gene modify the posterior half of each segment, making it the mirror image of the anterior half. As development continues, and different classes of segmentation genes are activated, the borders of stripes of activation for these various genes become sharper and sharper, until cell-cell interactions focus the expression of different genes to neighboring cells. For example, we see in gure 16.16 the narrowing and sharpening of the even-skipped and fushi tarazu bands in the developing embryo. (The gene fushi tarazu, meaning not enough segments in Japanese, is a pair-rule gene.) Most segmentation genes are speci c transcription factors, genes that interact with DNA to activate or repress transcription. Thus, pattern formation in develop-
ment is a process of activating different genes in sequence, gradually narrowing the scope of which cells express a particular gene. There is one nal group of genes we will discuss in this developmental cascade in Drosophila. At this early stage of development, these genes, the homeotic genes, take control of the development of the segments.
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