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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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a posterior tip, called the telson, that will give rise to the internal structures at the very posterior end of the y. The fates of these segments have been determined by treating them with various harmless dyes and tracing where the dyes end up. A projection of adult structures on embryonic tissue is called a fate map.
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The General Body Plan
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The role genes play in determining the general axes of the body plan has been worked out at several levels. First, mutations causing female sterility were isolated. (C. N sslein-Volhard and E. Wieschaus were instrumental in systematically isolating many of these mutants; they were awarded Nobel prizes for this work.) For example, among normal female ies that were sterile, some
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Christiane N sslein-Volhard (1942 ). (Courtesy of
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Christiane N sslein-Volhard.)
Eric F. Wieschaus (1947 ). (Courtesy of Dr.
Eric F. Wieschaus. Photograph by Denise Applewhite.)
The relationship between parasegments, segments, and the adult fruit y. The initial segments of the y are called parasegments; the nonsegmented parts of the embryo are called the acron at the head end (accounting for eyes and antennae) and the telson at the tail end (accounting for the end of the alimentary canal). Later segments are made up of the posterior end of one parasegment and the anterior portion of the next (p, a). The later segments map directly on the adult body, accounting for mouthparts (mandible, maxilla, and labium), thoracic segments (1 3), and abdominal segments (1 8). (H is for head.)
(From P. A. Lawrence, The Making of a Fly, Copyright 1992 Blackwell Science, Ltd., Oxford, England. Reprinted by permission.)
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Sixteen
Gene Expression: Control in Eukaryotes
produced embryos without heads or thoracic structures. The gene for this mutation, which has since been cloned and sequenced, is called bicoid (fig. 16.9). It codes for a speci c transcription factor, the Bicoid protein. (Remember that gene names are italicized, using the rst letter, lowercase for recessive and uppercase for dominant; the protein product of these genes is not italicized, but the rst letter is capitalized.) Pricking the anterior end of a normal embryo, causing the loss of cytoplasm from that end ( g. 16.10), can mimic these mutants. This experiment indicates there is some cytoplasmic localization determining the development of the anterior end of the y. To support that idea further, it was possible to get normal development from a bicoid y by injecting the anterior end with cytoplasm from a normal embryo ( g. 16.10b). This process of facilitating normal development by manipulating the embryo is termed a rescue experiment. By probing with a complementary oligonucleotide to the bicoid messenger RNA, researchers found that the bicoid messenger RNA is formed in the nurse cells and then passed into the oocyte, where it becomes localized at the anterior tip ( g. 16.11a). After fertilization, this messenger RNA is translated into Bicoid, which begins to diffuse from the anterior end of the egg, until it reaches about 50% of the length of the egg. The protein can be visibly located by treating the eggs with antibodies to the protein; these antibodies can then themselves be made visible ( g. 16.11b). The Bicoid protein is called a morphogen, a substance that diffuses through the egg and by its concentration determines the developmental fate of that part of the embryo. Although nurse cells are germ-line cells, they are of maternal origin and not from the embryo. Since
maternal cells, not the embryo itself, produce this morphogen, the gene responsible for its production is called a maternal-effect gene. Other maternal-effect genes are involved in formation of the anterior pattern that produces headless embryos. However, they don t appear to produce a morphogen. Rather, these genes seem to be involved in the transport, stabilization, and modi cation of the morphogen. In mutants of these other genes (swallow, exuperantia), Bicoid is found in the nurse cells but not in the embryo; cytoplasm from the nurse cells of these mutants can rescue bicoid mutants, indicating that the morphogen is present but not delivered to the oocyte. Only mutants of the bicoid gene itself cannot rescue the various headless mutants because only in bicoid mutants is the morphogen itself missing. Through experiments similar to the ones described for bicoid, four independent signaling pathways of maternal-effect genes have been isolated. These pathways determine the general body plan of the developing embryo: anterior, posterior, terminal, and dorso-ventral. The posterior pattern is controlled by the gradient of a protein, Nanos. Before the nanos gene is active, producing messenger RNA, the rst posterior gene active is oskar; the localization of oskar messenger RNA then de nes the localization of nanos messenger RNA. Mutant embryos can be rescued by wild-type cytoplasm; the nanos mes-
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