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Figure 16.13 The interaction of the oocyte nucleus and follicle cells early in development. (a) The oocyte nucleus, located posteriorly in the oocyte, activates posterior follicle cells. These cells will later provide the nanos messenger RNA to control posterior development of the embryo. After this interaction, a product of the follicle cells causes a rearrangement of microtubules in the oocyte, moving the oocyte nucleus anteriorly and dorsally. There the same interaction takes place, in this case activating follicle cells to control dorsal development. (b) The oocyte signal (red circles) is the product of the gurken gene; it interacts with a receptor (blue Y-shapes) on the surface of follicle cells, the product of the torpedo gene.
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III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Patterns in Development
ventral axis. As of yet, we don t know all of the signaling going on, nor why two similar cell types react differently to the same oocyte signal (Torpedo), but we do know that maternal-effect genes in the follicle cells are induced by the oocyte itself. Maternal-effect genes are the rst in a series controlling a cascade of gene expression that eventually determines the fates of individual cells in the developing y embryo.The rest of the genes are zygotic genes, genes active in the cells of the embryo itself. As we move down this cascade of genes, we go from broad patterns to more and more focused gene activity. We go from a single cell with gradients of morphogens to stripes of cells with different genes active.
Segmentation Genes
Once the general body plan of the y is in place, development continues in the formation of parasegments and then segments. The various organs of the y s body are produced from these segments. Further development is now under the control of the zygote s own genes, generally referred to as segmentation genes. These genes fall into three general categories: gap genes, pair-rule genes, and segment-polarity genes (table 16.2; g. 16.14).These genes are activated sequentially, each by the genes activated before it; each group controls a smaller and more focused domain of the y s development. In this discus-
sion, we will concentrate on the anterior-posterior system. The maternal-effect genes of the anterior-posterior system have created Bicoid and Nanos gradients.The segmentation genes increment, narrow, and focus these gradient signals until fourteen distinct bands form, corresponding to the fourteen parasegments that develop, creating compartments that the tissues of the y arise from (e.g., wings, legs, bristles). The gap genes were rst discovered as mutants that resulted in missing segments in the embryo ( g. 16.14). The Bicoid and Nanos gradients act on gap genes, specifically hunchback. Although the Hunchback protein is present in the egg from maternal production, the maternally supplied quantity is apparently not signi cant. Bicoid and Nanos independently create a Hunchback gradient that is maximal at the anterior end of the embryo, due to activation by Bicoid, and absent at the posterior end, due to Nanos repression. Bicoid is a speci c transcription factor that can bind to at least six sites in the promoter region of the hunchback gene. Three of these sites are strong binding sites and three are weak. Thus, depending on the concentration of Bicoid in the gradient, different levels of Hunchback are produced, creating the Hunchback gradient. Experiments with extra copies of the bicoid gene show that it is the actual quantity of Bicoid present at a particular point, and not the shape of the gradient, that actually determines the effect.
Table 16.2 Segmentation Genes in Drosophila
Class Gap Locus Kr ppel knirps hunchback Pair-rule paired even-skipped odd-skipped barrel* runt engrailed Segment-polarity cubitus interruptus wingless gooseberry hedgehog fused patch Allelic Designation Kr kni hb prd eve odd brr run en ci wg gsb hh fu pat Chromosome 2 3 3 2 2 2 3 1 2 4 2 2 3 1 2
Source: Reprinted with permission from Nature, Vol. 287, C. Nusslein-Volhard and E. Wieschaus. Copyright 1980 Macmillan Magazines Limited. * barrel is a synonym of the hairy gene.
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