Morphogenetic fate map of prospective adult structures of the honey bee

Previously published data on 40 honey-bee gynandromorphs were used to develop a morphogenetic blastoderm fate map indicating the relative locations of several presumptive adult structures. Comparison of honey-bee blastoderm fate map with Drosophilia maps reveals a general similarlity between them.     

Entwicklungsgenetische Untersuchungen an Gynandern vonDrosophila melanogaster

Summary Gynandromorphs with female XX-and male XO-areas result from the loss of an unstable ring-X-chromosome in the early cleavage mitoses of ring/rod-X-chromosome heterozygotes. The phenotypes of the recessive alleles on the rod-X-chromosome are expressed in the XO-areas.377 larval gynandromorphs of the genotypeR(1)2, In(1)w ^vC /y w sn³Iz^50e mal were examined and scored for the phenotypes of 13 paired and 10 unpaired structures (Table 2, Fig. 2). This was possible mainly by the cell-autonomous expression of aldehyde oxidase activity in soft tissues and by the comparison of the distribution of enzyme activity in wildtype and gynander larvae. The distances between pairs of structures were calculated in sturt-units (Tables 3 and 4). A morphogenetic fate map with the presumptive areas of larval structures was constructed (Fig. 3). The relative positions of the structures agree well with Poulson's fate map (Fig. 4). In addition, the distribution of phenotypes was scored in 380 adult gynandromorphs Table (5). The fate map (Fig. 5) which was constructed from these data is very similar to the fate map of larval structures. This similarity becomes even more pronounced if fate maps are constructed which contain only structures analogous in larva and imago (Table 6, Fig. 6). Therefore an attempt was made to set up an integrated morphogenetic fate map containing the presumptive areas of both larval and imaginal structures (Fig. 7). The possibilities of further blastoderm mapping are discussed.

Herrn Prof. Dr. Dr. h. c. B. Rensch zum 75. Geburtstag gewidmet     

Behavior of gynandromorphs of the wasp Habrobracon juglandis.

A study was made of sex behavior for 276 gynandromorphs of the wasp Habrobracon juglandis. In general, gynandromorphs with male heads behaved like males, those with female heads behaved like females, and those with mosaic heads as either male or female or both. These observations corroborate those made by P. W. Whiting in 1932 and confirm his view that the site of sex behavior is in the brain. Correlations between sex behavior and the sex of various body structures showed clearly that sex behavior is unrelated to the thorax, abdomen and genitalia. The sex behavior of gynandromorphs does not seem to be related to the sex of the external sensors of the head but rather to the genotype of the brain.     

Analysis of morphogenetic movements in the development of the notum anlage of Drosophila melanogaster

Summary A comparison of the morphogenetic maps of the notum anlage of Drosophila melanogaster derived from the gynandromorph data and mosaics induced by somatic crossing-over during the first instar larval stage revealed that practically no major morphogenetic movements occur in the development of the anlage between the blastoderm and first instar larval stages and the adult stage. By comparing the morphogenetic map derived from gynandromorphs and the fate map derived from data on the transplantation of fragments of the mature wing imaginal disc, it was observed that no major morphogenetic movements occur in the notum anlage between the stages of the allocation of the disc and the mature disc. The results are consistent with the observations of other authors concerning the larval development of eye-antenna, wing and leg discs.     

A New Mutant Controlling Mitotic Chromosome Disjunction in DROSOPHILA MELANOGASTER

A new mutant, mit (mitotic loss inducer), is described. The mutant is recessive and maternal in action, producing gynandromorphs and haplo-4 mosaics among the progeny of homozygous mit females. Mosaic loss of maternal or paternal chromosomes can occur. The probabilities of either maternal or paternal X chromosome loss are equal. mit has been mapped to approximately 57 on the standard X chromosome map.-Using gyandromorphs generated by mit, a morphogenetic fate map, placing the origins of 40 cuticular structures on the blastoderm surface, has been constructed. This fate map is consistent with embryological data and with the two other fate maps generated in different ways.     

A DETAILED FATE MAP OF THE WING DISC OF SARCOPHAGA PEREGRINA

In order to construct an anlageplan of the imaginal disc, various fragments of the wing disc from mature larvae of Sarcophaga peregrina were implanted into host larvae of the same age. The implants were metamorphosed with the host and the differentiated adult structures were analyzed.
On the basis of the results obtained from the transplanted disc fragments, a fate map of the wing disc was satisfactorily constructed.
The fate map of S. peregrina is similar to an earlier study of Drosophila , but the PAA, PWP, and AP are located in different positions from those of Drosophila .     

Control of Female Reproduction in Drosophila: Genetic Dissection Using Gynandromorphs

The sexual behavior of Drosophila melanogaster gynandromorphs was studied to analyze the relationship between different steps in the female reproductive pathway. It was assumed that, in some gynandromorphs, certain female functions are missing because the corresponding control sites (foci) are either composed of male tissue or did not develop. A given gynandromorph can show elements of both male and female reproductive pathways. None of the steps of the female reproductive pathway appeared to be dependent on any other, in contrast to male behavior where, for example, following of females is a prerequisite for attempted copulation. By correlating each of the behaviors with the genotype of the cuticle, we confirmed previous findings that the focus for the female sex appeal is located in the abdomen, but receptivity to copulation is controlled by a site in the head. Many of the gynandromorphs did not lay eggs, presumably because either the focus controlling egg transfer from the ovaries to the uterus or the one controlling egg deposition was composed of male tissue. Many of the nonovipositing gynandromorphs laid eggs while dying or could be induced to deposit eggs after implantation of hormone-producing glands or topical application of a juvenile hormone analog. Some of the noninseminated gynandromorphs laid eggs at the rate characteristic for inseminated females, suggesting that an oviposition focus (mapping in the head region) suppresses oviposition in virgin females, but not in gynandromorphs whose focus is composed of male tissue. Some of the inseminated gynandromorphs oviposited eggs at a low rate, possibly because the focus responsible for detection of insemination could not function properly. Some of the inseminated gynandromorphs laid unfertilized eggs, revealing the importance of the focus controlling sperm release from the seminal receptacle. Foci controlling egg transfer, egg deposition and sperm release are located in the thorax, according to mosaic fate mapping results and studies on the reproductive behavior of decapitated females. The location of egg deposition in the culture vial seems to be controlled by a brain site. Sexual behavior in Drosophila does not depend on the presence (or absence) of the ovary or germ line.     

Developmental compartments in the Drosophila melanogaster wing disc

Distribution of the enzyme aldehyde oxidase (AO) within the pouch of the mature wing disc is precise and differential. General locations of compartmental boundaries have been identified by fate mapping and studies of AO distribution. The suspected locations of the boundaries were verified by analyzing the distribution of AO-negative cells within an AO-stained background in gynandromorphs and in X-ray-induced clones of AO-negative cells. The anterior/posterior border appeared slightly anterior to the junction of the AO⁺ anterior presumptive wing surfaces and AO^? posterior wing surfaces. A narrow band of AO⁺ cells extending proximodistally on both presumptive wing surfaces belongs to the posterior compartment. Two dorsal/ventral (dor./vent.) restrictions were found. The dor./vent. restriction equivalent to the dor./vent. border found in the adult wing was located at the ventral most edge of the AO-stained presumptive wing margin. A second restriction which was less strictly obeyed was found on the dorsal edge of the wing margin. We conclude that the whole presumptive wing margin is part of the dorsal compartment. Within the anterior wing margin an intensively stained oval was also found to be clonally restrictive. Therefore, territories were found within the prospective wing margin for which no such features have been identified in the adult Drosophila melanogaster wing.     

Adult abnormalities resulting from a localized blastoderm defect in a maternal-effect mutant of Drosophila.

In the mutant mat(3)3 of Drosophila melanogaster, there is a temperature-sensitive maternal effect on blastoderm formation. When oogenesis occurs in homozygous mat(3)3 females at the fully restrictive temperature of 29°C, the embryonic progeny form a defective cellular blastoderm in which cells are either completely or partially missing from a posterior-dorsal region, and the embryos die before hatching. Transplantation tests for the presence in the embryos of primordial imaginal cells capable of developing into adult structures showed a relatively high yield of eye and antenna structures, an intermediate yield of labium structures, and low or zero yields of wing, haltere, and leg structures. These results are consistent with the fate mapping of the primordial imaginal cells by analysis of gynandromorph mosaics; the eye and antenna map in the fully cellular region of the mutant blastoderm, the labium near the border of the defective region, and the wing, haltere, and legs within the defective region. When oogenesis oocurs at a lower temperature, the lethal maternal effect in mat(3)3 is reversed, but there is a nonlethal effect on larval and adult progeny of the mat(3)3 females. Many of the adults are missing one or more cuticular structures, usually a leg, haltere, or abdominal segment, and many of the larvae are missing the corresponding imaginal discs from which the thoracic structures are derived. These selective effects on imaginal development appear to be caused by maternally induced blastoderm defects that are less extensive at the lower temperature of oogenesis.     

Pattern Regulation of the Leg Disc of the Fleshfly, Sarcophaga peregeina

A fate map of the hind leg disc of Sarcophaga peregrina was constructed by examining the adult structures of implanted disc fragments. The locations of presumptive adult structures in the disc were similar to those of fore leg disc of Drosophila and Sarcophaga ruficornis . However, the concentric borderlines of the segments could not be ascertained in the present case.
Pattern regulation of disc fragments was studied by culturing them either in adult females for several days or for 3 days in mature larvae placed on wet condition. Cultured disc fragments regenerated or duplicated as in Drosophila , with some exceptions. For instance, the region with a high density of positional values, the upper medial quarter, of the fore leg disc of Drosophila was not found. A characteristic difference in the rate of regeneration or duplication was observed in the implanted fragments, when cultured in larvae or adult hosts. This variable pattern regulation in larval and adult hosts could be due to different compositions of the hemolymph in which would healing of the implanted disc fragments takes place.     

Developmental fates of larval tissues after metamorphosis in the ascidian, Halocynthia roretzi

Cell lineages during ascidian embryogenesis are invariant. Developmental fates of larval mesodermal cells after metamorphosis are also invariant with regard to cell type of descendants. The present study traced developmental fates of larval endodermal cells after metamorphosis in Halocynthia roretzi by labeling each endodermal precursor blastomere of larval endoderm. Larval endodermal cells gave rise to various endodermal organs of juveniles: endostyle, branchial sac, peribranchial epithelium, digestive organs, peripharyngeal band, and dorsal tubercle. The boundaries between clones descended from early blastomeres did not correspond to the boundaries between adult endodermal organs. Although there is a regular projection from cleavage stage and larval stage to juvenile stage, this varies to some extent between individuals. This indicates that ascidian development is not entirely deterministic. We composed a fate map of adult endodermal organs in larval endoderm based on a statistical analysis of many individual cases. Interestingly, the topographic position of each prospective region in the fate map was similar to that of the adult organ, indicating that marked rearrangement of the positions of endodermal cells does not occur during metamorphosis. These findings suggest that fate specification in endoderm cells during metamorphosis is likely to be a position-dependent rather than a deterministic and lineage-based process. Received: 16 June 1999 / Accepted: 16 August 1999     

A new species ofTrichogramma [T. Cordubensis n. sp.] [Hym.: Trichogrammatidae], parasitoid ofHeliothis eggs in cotton crops in the sw of spain

Trichogramma cordubensis n. sp. is a parasite ofHeliothis armigera Hb. eggs in cotton crops in western Andalusia. Reproduced onEphestia kuehniella Zeller eggs, it is described morphologically and compared withT. chilonis Ishii andT. evanescens Westwood as both present similarities with this species.T. cordubensis can be distinguished in the male genitalia by the shape of the dorsal expansion of the gonobase and the size, shape and relative position of the median ventral projection. Male antennal setae are longer inT. cordubensis than in the other 2 species. In addition, unlikeT. chilonis andT. evanescens, T. cordubensis is thelytokous at 20°C. in our laboratory, though males and gynandromorphs can be obtained by altering the rearing temperature, this being another difference of this species.     

Fine cytogenetical analysis of the band 10A1-2 and the adjoining regions in the Drosophila melanogaster X chromosome. III. The fate mapping of the lethal focus of the l(1)BP4 gene

Summary Focuses of the l(1)BP4, vermilion and sevenless genes located in the same band 10A1-2 of Drosophila melanogaster X-chromosome, have been compared. For this purpose first the fate map of the blastoderm for 60 pairs of adult cuticular landmarks has been built based on scoring 612 gynandromorphs. The location of the legs and antenna primordia has been defined more exactly as compared to analogous maps by other authors. The location of new landmarks has been carried out: mesosternal bristle, sex-comb and first tergite primordia. The focus of the l(1)BP4 gene has been determined with the help of this map on the basis of analysis of 321 mosaics. The data obtained show that the lethal focus of the gene belongs to the bilateral domineering type and is located in the blastula area giving rise to the nervous system, behind the region of the third thoracic ganglia origin. Beside, there is a pair of autonomous nonlethal focuses of the same gene limited by the wings' zone. The focuses of the l(1)BP4 gene do not coincide with the known focuses of vermilion and sevenless genes, which means that all three genes are active in different tissues.     

The three leg imaginal discs of Drosophila: "Vive la différence"

The imaginal discs of Drosophila are the larval primordia for the adult cuticular structures of the adult fly. Fate maps of different discs have been generated that show the localization of prospective adult structures. Even though the three legs differ in their morphology, only the fate map for the T1 (prothoracic) leg disc has been generated. Here we present fate maps for the T2 (meso-) and T3 (metathoracic) leg discs. We show that there are many similarities to the map of the T1 leg disc. However, there are also significant differences in the contributions of each disc to the thorax, in the morphology of joints connecting the legs to the thorax, in bristle patterns, and in the positioning of some sensory organs. We also tested the developmental potential of disc fragments and observed that T2 and T3 leg discs have more limited plasticity and are unable to transdetermine. The differences in the cuticle patterns between legs are robust and conserved in many species of dipterans. While most previous analyses of imaginal disc development have not distinguished between the different leg discs, we believe that the underlying differences of the three leg discs demonstrated here cannot be ignored when studying leg disc development.     

Requirements for integrins during Drosophila development

The common beta subunit of the PS antigens of Drosophila is homologous with vertebrate integrins and is encoded by the lethal(1)myospheroid gene. We have generated flies mosaic for wild-type and mutant alleles of lethal(1)myospheroid using adult gynandromorphs and radiation-induced somatic crossing over. The defects observed in the gynandromorphs demonstrate widespread requirements for PS integrins during development especially in ventrally derived structures, which also show strong expression of PS beta integrin. Smaller lethal(1)myospheroid clones induced during larval development result in blister and vein defects in the wings and aberrant development of photoreceptor cells, demonstrating roles for PS integrins during development of both wings and eyes. PS integrins are required for the close apposition of the dorsal and ventral wing epithelia and for the proper arrangement of photoreceptor cells. However, many other adhesive and morphogenetic processes proceed normally in the absence of integrins containing the beta subunit encoded by lethal(1)myospheroid.     

Maternal-effect genes as the recording genes of Turing-Child patterns: sequential compartmentalization in Drosophila

The early embryo is often a two-dimensional surface. The fate map is the subdivision of this surface into regions which give rise to parts of the phenotype. It is shown for Drosophila that the fate map is generated by the spontaneous and sequential formation of Turing-Child (TC) eigenfunction patterns. These patterns are recorded by the maternal-effect genes. The addition of the nodal lines of the TC patterns yields the correct number, positions, sequences and symmetries of regional boundaries. A simplest nontrivial 'homeotic transformation' is suggested and explained. A single mutation converts a region in one end of the fate map to a mirror-symmetric image of a nonadjacent region in the other end of the fate map, and this is attributed to the geometry of the TC patterns. This geometry also determines the initial shape of the zygotic gene expression. The vision of William Bateson that biological form is shaped like Chladni's patterns in acoustics and music is justified. A similar sequence of TC patterns occurs in the normal development of all organisms, and it is suggested that artificial intervention which completes the full sequence of TC patterns can be useful in the context of regenerative medicine and this is illustrated with the sea urchin.     

Two gynandromorphs of Sympetrum striolatum (Charpentier, 1840) (Odonata: Libellulidae)

Two gynandromorphs of the dragonfly Sympetrum striolatum , with different features, are described here, with special emphasis on the genitalic structures.     

Localized ultraviolet laser microbeam irradiation of early Drosophila embryos: fate maps based on location and frequency of adult defects.

Drosophila embryos were locally irradiated with a 257-nm laser microbeam during blastoderm and germ band stages. Depending on stage and beam diameter (10–30 μm), from 0 to 45 nuclei were exposed to the uv radiation. The doses used, 5 or 10 erg, did not eliminate nuclei or cells at once, but up to 50% of the adult survivors from irradiated eggs carried defects in the thorax. These were scored with reference to the imaginal discs from which the affected structures derive. For each thoracic disc a “target center” was calculated as the weighted mean value of all beam locations affecting the respective adult derivatives. The target centers for the germ band stage map within the respective germ band segments. The pattern of target centers for the blastoderm stage is comparable to the thoracic region of published fate maps, and the distances between adjacent leg centers (approximately three cell diameters) agree with recent evidence based on mosaic flies. We discuss the question whether the target centers mark the position of the respective disc progenitor cells at the stages of irradiation and conclude that these positions are rendered rather correctly at least with reference to the longitudinal egg axis.     

Prospective fates and regulative capacities of fragments of the female genital disc of Drosophila melanogaster.

A fate map of the female genital disc of Drosophila melanogaster was established by examining the derivatives of fragments transplanted into host larvae for metamorphosis. The fate map is presented as a two-dimensional projection, but for several reasons it is proposed that the anal plates originate from the dorsal epithelial layer whereas the genitalia are produced from the ventral layer. Fragments produced by cuts parallel to the axis of symmetry of the disc undergo regeneration during culture in adult hosts if the fragments comprise more than half of the disc, or duplication if they comprise less than half. Most of the fragments generated by bilaterally symmetrical cuts across the line of symmetry of the disc undergo neither regeneration nor duplication during culture, but with some such fragments there is a low frequency of regeneration. It is argued that the usual lack of regeneration in these fragments results from wound healing which confronts identical positions from right and left sides, giving no growth stimulation. The fragments which regenerate might do so as a result of healing between dorsal and ventral surfaces, providing the discontinuity in positional information which is thought to be involved in growth stimulation.     

Morphogenetic and cellular movements that shape the mouse cerebellum; insights from genetic fate mapping

We used the cerebellum as a model to study the morphogenetic and cellular processes underlying the formation of elaborate brain structures from a simple neural tube, using an inducible genetic fate mapping approach in mouse. We demonstrate how a 90 degrees rotation between embryonic days 9 and 12 converts the rostral-caudal axis of dorsal rhombomere 1 into the medial-lateral axis of the wing-like bilateral cerebellar primordium. With the appropriate use of promoters, we marked specific medial-lateral domains of the cerebellar primordium and derived a positional fate map of the murine cerebellum. We show that the adult medial cerebellum is produced by expansion, rather than fusion, of the thin medial primordium. Furthermore, ventricular-derived cells maintain their original medial-lateral coordinates into the adult, whereas rhombic lip-derived granule cells undergo lateral to medial posterior transverse migrations during foliation. Thus, we show that progressive changes in the axes of the cerebellum underlie its genesis.