The Drosophila sex determination hierarchy modulates wingless and decapentaplegic signaling to deploy dachshund sex-specifically in the genital imaginal disc

The integration of multiple developmental cues is crucial to the combinatorial strategies for cell specification that underlie metazoan development. In the Drosophila genital imaginal disc, which gives rise to the sexually dimorphic genitalia and analia, sexual identity must be integrated with positional cues, in order to direct the appropriate sexually dimorphic developmental program. Sex determination in Drosophila is controlled by a hierarchy of regulatory genes. The last known gene in the somatic branch of this hierarchy is the transcription factor doublesex (dsx); however, targets of the hierarchy that play a role in sexually dimorphic development have remained elusive. We show that the gene dachshund (dac) is differentially expressed in the male and female genital discs, and plays sex-specific roles in the development of the genitalia. Furthermore, the sex determination hierarchy mediates this sex-specific deployment of dac by modulating the regulation of dac by the pattern formation genes wingless (wg) and decapentaplegic (dpp). We find that the sex determination pathway acts cell-autonomously to determine whether dac is activated by wg signaling, as in females, or by dpp signaling, as in males.

Sex determination genes control the development of the Drosophila genital disc, modulating the response to Hedgehog, Wingless and Decapentaplegic signals

In both sexes, the Drosophila genital disc contains the female and male genital primordia. The sex determination gene doublesex controls which of these primordia will develop and which will be repressed. In females, the presence of Doublesex(F) product results in the development of the female genital primordium and repression of the male primordium. In males, the presence of Doublesex(M) product results in the development and repression of the male and female genital primordia, respectively. This report shows that Doublesex(F) prevents the induction of decapentaplegic by Hedgehog in the repressed male primordium of female genital discs, whereas Doublesex(M) blocks the Wingless pathway in the repressed female primordium of male genital discs. It is also shown that Doublesex(F) is continuously required during female larval development to prevent activation of decapentaplegic in the repressed male primordium, and during pupation for female genital cytodifferentiation. In males, however, it seems that Doublesex(M) is not continuously required during larval development for blocking the Wingless signaling pathway in the female genital primordium. Furthermore, Doublesex(M) does not appear to be needed during pupation for male genital cytodifferentiation. Using dachshund as a gene target for Decapentaplegic and Wingless signals, it was also found that Doublesex(M) and Doublesex(F) both positively and negatively control the response to these signals in male and female genitalia, respectively. A model is presented for the dimorphic sexual development of the genital primordium in which both Doublesex(M) and Doublesex(F) products play positive and negative roles.

VISUAL DETERMINATION OF SEX OF DIAMONDBACK MOTH LARVAE

The diamondback moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), is a major worldwide pest of cruciferous vegetables (Talekar and Shelton 1993). Male and female adults of this species can be accurately identified by visible differences in their external genitalia. According to Robertson (1939), the sex of pupae can also be identified, but, in our experience, this approach is more difficult. For some types of research, determination of larval sex would be useful. For example, one could determine sex-specific responses of larvae to a treatment by observing mortality, development rate, or behavior of male and female larvae separately. In studies requiring hybrid mating between different strains of diamondback moth (Tabashnik et al. 1992), one could isolate groups of male larvae separately from female larvae to avoid mating within a strain. Some lepidopteran larvae can be sexed by the gonads being visible through the cuticle of the fifth abdominal segment or by the position of external pits marking the location of the genital discs in the eighth and ninth abdominal segments (Stehr 1987). But, because of their small size, it is not practical to sex diamondback moth larvae by external pits and we have found no studies in which male and female larvae were identified by gonadal form. Herein we report that third- and fourth-instar larvae of diamondback moth can be sexed efficiently and reliably.

orthodenticle activity is required for the development of medial structures in the larval and adult epidermis of Drosophila

Lethal alleles of orthodenticle (= otd) cause abnormalities in the embryonic head that reflect an early role in anterior pattern formation. In addition, otd activity is required for the development of the larval and adult epidermis. Clonal analysis of both viable and lethal alleles shows that the adult requirement for otd is restricted to medial regions of certain discs. When otd activity is reduced or removed, some medial precursor cells produce bristles and cuticle characteristic of more lateral structures. Similar medial defects are observed in the larval epidermis of embryos homozygous for lethal otd alleles. Antibodies to otd recognize a nuclear protein found at high levels in the medial region of the eye antennal discs, the leg discs, the genital discs and along the ventral midline of the ventral epidermis of the embryo. These results suggest that the otd gene product is required to specify medial cell fates in both the larval and adult epidermis.

TEMPERATURE-SENSITIVE CELL-LETHAL MUTANTS OF DROSOPHILA: ISOLATION AND CHARACTERIZATION

ABSTRACT One hundred and twenty-one temperature-sensitive (ts) sex-linked lethals were screened by means of X-ray-induced somatic crossing over to determine if any were ts cell-lethal mutants. Cell-lethal mutations were identified by their ability to block the development of homozygous clones when raised under restrictive conditions (29°). Twenty-two ts cell-lethal mutants were isolated and categorized into three classes, depending upon the patterns of damage observed in larval and imaginal tissues. The phenotypes produced by these mutations ranged from those which affected only a limited set of structures (i.e., genital discs only) to those which affected diverse tissues at all stages of the life cycle. Each mutation has its own characteristic time-dependent pattern, frequency, and type of damage. All the mutations affect imaginal tissue, but only one-third of the mutations affect both larval and imaginal tissue. The fastest-acting lethals need 15 hours at the restrictive temperature to kill the cells and the slowest-acting lethals require at least 48 hours. By choosing the appropriate mutant and by manipulating the times of exposure to the restrictive temperature, it has proven possible to produce duplications and deficiencies in specific structures of the adult. A mechanism by which lethality might yield such structures is suggested. In addition, 15 of the mutants are ts female sterile mutants. Only one of these 15 mutants can recover its fertility when shifted back down to the permissive temperature (22°).

The larval development of Dacus tryoni (Frogg,) (Diptera : Trypetidae) 1. larval instars, Imaginal discs, and haemocytes.

In D. tryoni three larval instars occur, distinguishable by their mouth hooks and anterior and posterior spiracles. In the first two instars, the imaginal discs established during embryonic development grow slowly, only the cephalic discs showing marked change. Paired labial and lateral genital discs invaginate during the second half of the 2nd instar. During the 3rd instar, the imaginal discs increase further in size, first rapidly, then more slowly. Each shows attenuation of the lateral wall as a peripodial membrane, thickening and folding of the median wall without histodifferentiation and, in the wing, haltere, leg, and genital discs, formation of mesoderm from cells in the outer part of the median wall. Disc morphogenesis is largely completed by 120 hr after hatching. A single pair of lymph glands and 13-15 pairs of pericardial cells lie along the heart. Both show cell enlargement during larval life but no proliferation. Haemocytes in the blood arise directly from embryonic mesoderm as two main types, numerous spheroids and less numerous hexagons. They circulate freely in the living larva, but tend to occur in irregular clumps in fixed material, especially after 120 hr. No epigenetic relationship between haemocytes, imaginal discs, and lymph glands such as that described for Drosophila melanogaster by el-Shatoury and Waddington can be discerned in D. tryoni. The haemocytes do not arise from the lymph glands. The disc mesoderm does not arise from the haemocytes. The larval development of the discs is largely completed before the haemocytes become numerous.