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(c) Figure 16.5
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Role of transposition in controlling the mating type in yeast. (a) Mating-type loci on the third chromosome. MAT is the active mating-type locus. HML and HMR are silent loci, carrying the two mating-type alleles, and a, respectively. (b) Transposition of HML to MAT results in the MAT allele at the MAT site and the mating type. (c) Transposition of HMR to MAT results in an active MATa allele, yielding the a mating type.
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Drosophila Development
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The fruit y, Drosophila melanogaster, has emerged as an excellent model organism for the study of development. The zygote develops from the egg, in maternal cytoplasm. Maternal messenger RNAs and proteins are the first expressed in the embryo. These substances
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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
Sixteen
Gene Expression: Control in Eukaryotes
rst determine the broad pattern of the embryo. Then, through signal pathways involving numerous speci c transcription factors, they initiate a cascade of gene expression that eventually determines the fate of each cell. As we will see, many parallels exist between the fruit y and higher organisms. We will concentrate on two overall patterns of development here: the formation of the basic body plan (anterior-posterior and dorsal-ventral polarity, which results in a segmented embryo that has a front, back, top, and bottom) and the determination of gene expression within segments. Drosophila development begins within a follicle that contains the oocyte surrounded by follicle and nurse cells. The fteen nurse cells, along with the oocyte, were derived from four divisions of an earlier germ-line cell ( g. 16.6). The nurse cells maintain connections to each other and to the oocyte by cytoplasmic bridges, openings in the membranes surrounding the cells. Thus, the nurse cells can readily pass materials (messenger RNAs and proteins) into the oocyte. After fertilization, the diploid nuclei divide thirteen times in the space of about 3.5 hours, forming a syncitium a group of nuclei without cell membranes. During this time, most of the nuclei migrate to the inner surface of the developing embryo, where cell membranes eventually form, producing a cellular blastoderm. During the syncitial period, materials can move freely through the cytoplasm. At the posterior end of the embryo, several cells, called pole cells, that will eventually form the germ cells of the developing y are set aside ( g. 16.7). Development then proceeds through gastrulation, in which cells grow inward, forming the basic germ layers of the embryo (mesoderm, endoderm, and ectoderm). From these layers, various adult struc-
tures will arise. At about six hours of development, furrows become visible in the embryo, delineating segments. The rst segments visible are called parasegments. They do not give rise to the later segments of the embryo, but rather overlap the later segments in a simple fashion: Each later segment is made up of the anterior end of one parasegment and the posterior end of the next ( g. 16.8). This distinction is meaningful since, as we shall see later, some genes express themselves within the borders of parasegments rather than segments. The fully segmented embryo has an anterior region, destined to be the head; three thoracic segments, which will give rise to the thorax (the middle region of the y containing wings and legs); and eight abdominal segments that will give rise to the abdomen. The embryo also has an anterior tip, the acron, that will give rise to structures at the very head end eyes, and antennae; and
0.5 hours
1.5 hours
Pole cells Nucleus 2.5 hours
Oocyte (egg) 3.25 hours
Nurse cells
Follicle cells
Figure 16.6 The follicle from a fruit y, Drosophila, consisting of the oocyte, fteen nurse cells arising from four divisions of a germ-line cell that also gave rise to the oocyte, and follicle cells.
Figure 16.7 Development of the fertilized Drosophila egg after laying. Pole cells, which will be future germ cells, are set apart at about 2 hours. A syncitial blastoderm forms at about 2.5 hours, followed by a cellular blastoderm, consisting of about ve thousand cells, at about 3.25 hours.
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