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Two Drosophila larvae, with cuticular patterns visible on the ventral surfaces. On the top is the wild-type with the cuticular pattern coinciding with thoracic and abdominal segments. On the bottom is a bicoid mutant, lacking head and thoracic structures. (Courtesy of Christiane N sslein-Volhard.)
Experiments to demonstrate that a cytoplasmic localization at the anterior end of the fruit y egg determines anterior structures. (a) A wild-type egg has anterior cytoplasm removed, resulting in a larva lacking anterior structures, similar to a bicoid mutant. (b) A bicoid mutant egg has anterior cytoplasm from a wild-type egg injected into the anterior of the egg, resulting in a larva indistinguishable from the wild-type.
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Patterns in Development
bicoid mRNA
the developing embryo. (Tyrosine kinases phosphorylate the amino acid tyrosine in speci c proteins.) Apparently, other genes in follicle cells located only at the poles of the egg produce a substance that activates the torso tyrosine kinase receptor, making it active in only the poles of the egg ( g. 16.12). A maternal-effect gene, Toll, that also produces a membrane receptor, controls the dorsoventral axis.Thus, we see that four pathways of maternaleffect genes determine the major body plan of the egg. Two of the pathways are determined by genes that result in diffusion of a morphogen (bicoid and nanos), and two are determined by genes for membrane receptors (torso and Toll ). About thirty maternal-effect genes are known (table 16.1).
(b) Bicoid protein
The bicoid morphogen rst appears in the fruit y egg as (a) messenger RNA at the anterior end of the egg. After fertilization, the messenger is translated into (b) Bicoid protein that diffuses toward the posterior end of the embryo.
(Courtesy of Daniel St. Johnston.)
senger RNA is localized at the posterior tip of the embryo and produces a protein that diffuses from that tip. Maternal-effect genes that act in a somewhat different manner control the other two pattern systems in the developing embryo. The terminal pattern controls development of both ends of the embryo; a key gene is torso. This gene codes for a membrane-bound tyrosine kinase receptor protein that is found evenly distributed on the outer surface of
The Drosophila follicle, showing follicular cells (green) at the tip of the oocyte that secrete a substance that activates the Torso (torso gene) tyrosine kinase at the areas marked by red lines; the inactivated kinase is located around the surface of the oocyte.
Table 16.1 Maternal-Effect Genes in Drosophila (Allelic Designations in Parentheses)
Anterior bicoid (bcd) swallow (swa) exuperantia (exu) bicaudal (bic) Bicaudal-D (BicD) Bicaudal-C (BicC) Posterior nanos (nos) oskar (osk) vasa (vas) tudor (tud) stauffen (stau) valois (val) pumilio (pum) Terminal torso (tor) trunk (trk) torsolike (tsl) polehole [fs(1) ph] Nasrat [fs(1) N] Dorso-Ventral Toll (Tl) nudel (ndl) pipe (pip) windbeutel (wbl) snake (snk) easter (ea) cactus (cact) sp tzle (spz) tube (tub) pelle (pll)
Source: Reprinted with permission from C. Nusslein-Volhard, et al., Science, 238:1675 81, 1987. Copyright 1987 American Association for the Advancement of Science.
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
Activity of maternal-effect genes in the follicle cells is controlled by an interaction between the oocyte itself and the follicle cells. Follicle cells at the anterior of the oocyte produce bicoid messenger RNA as a default condition. At the posterior of the oocyte, the follicle cells produce nanos messenger RNA, along with several other gene products. These follicle cells are induced to action by the product of the gurken gene in the oocyte; the oocyte nucleus is located posteriorly at this point, and its gene products can be directed to the posterior of the
Nurse cell Follicle cell
oocyte, where they diffuse to adjacent follicle cells. These cells have a receptor on their surfaces, the product of the torpedo gene, that recognizes the gurken gene product. Through signal transduction, these follicle cells are induced to express the nanos gene ( g. 16.13). At this point, some product of these follicle cells induces a reorganization of the microtubules in the oocyte, causing the oocyte nucleus to move anteriorly and dorsally. Now, the same gurken-torpedo interaction takes place, causing these follicle cells to induce the dorso-
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