Molecular organization of the Drosophila melanogaster Adh chromosomal region in D. replete and D. buzzatii, two distantly related species of the Drosophila subgenus

The molecular organization of a 1.944-Mb chromosomal region of Drosophila melanogaster around the Adh locus has been analyzed in two repleta group species: D. repleta and D. buzzatii. The extensive genetic and molecular information about this region in D. melanogaster makes it a prime choice for comparative studies of genomic organization among distantly related species. A set of 26 P1 phages from D. melanogaster were successfully hybridized using fluorescence in-situ hybridization (FISH) to the salivary gland chromosomes of both repleta group species. The results show that the Adh region is distributed in D. repleta and D. buzzatii over six distant sites of chromosome 3, homologous to chromosomal arm 2L of D. melanogaster (Muller's element B). This observation implies a density of 2.57 fixed breakpoints per Mb in the Adh region and suggests a considerable reorganization of this chromosomal element via the fixation of paracentric inversions. Nevertheless, breakpoint density in the Adh region is three times lower than that estimated for D. repleta chromosome 2, homologous to D. melanogaster 3R (Muller's element E). Differences in the rate of evolution among chromosomal elements are seemingly persistent in the Drosophila genus over long phylogenetic distances. This revised version was published online in July 2006 with corrections to the Cover Date.     

Taste and pheromone perception in the fruit fly Drosophila melanogaster

Taste is an essential sense for detection of nutrient-rich food and avoidance of toxic substances. The Drosophila melanogaster gustatory system provides an excellent model to study taste perception and taste-elicited behaviors. “The fly” is unique in the animal kingdom with regard to available experimental tools, which include a wide repertoire of molecular-genetic analyses (i.e., efficient production of transgenics and gene knockouts), elegant behavioral assays, and the possibility to conduct electrophysiological investigations. In addition, fruit flies, like humans, recognize sugars as a food source, but avoid bitter tasting substances that are often toxic to insects and mammals alike. This paper will present recent research progress in the field of taste and contact pheromone perception in the fruit fly. First, we shall describe the anatomical properties of the Drosophila gustatory system and survey the family of taste receptors to provide an appropriate background. We shall then review taste and pheromone perception mainly from a molecular genetic perspective that includes behavioral, electrophysiological and imaging analyses of wild type flies and flies with genetically manipulated taste cells. Finally, we shall provide an outlook of taste research in this elegant model system for the next few years.     

Ovulation and the Suppression of Mating in Drosophila melanogaster Females: Behavioral Basis

Virgin females of Drosophila melanogaster that are ectopically expressing the sex-pep-tide gene show a high level of ovulation and are unreceptive to males. However, if they are genetically deprived of eggs, receptivity is considerably restored (Fuyama, 1995). These females, whether they have eggs or not, extrude their ovipositors toward courting males as frequently as do fertilized females. However, this rejection behavior was ineffective in suppressing male courtship. Of females with eggs, about half of them could suppress male courtship. Females lacking eggs could not suppress male courtship and continued to elicit vigorous courtship. This difference seems to account for the increased mating frequency in sterilized females. Courtship behavior by mutant males defective in olfaction or learning suggested that females are capable of repelling males by emitting a volatile pheromone(s) with an inhibitory effect on male courtship.     

Pheromonal control of mating patterns inDrosophila melanogaster

Multiple-choice mating tests comparing the mating propensity among and within inbred lines were performed forDrosophila melanogaster. Courtship activity, time to copulation, and assortative mating were all directly correlated with the degree of inbreeding. By the eighth generation of sib mating, there was a 76% incidence of negative assortative mating in multiple-choice tests and a marked reduction of courtship behavior among sibs. Furthermore, absence of sperm in the females of nonreproductive pairs indicated that much of the attrition of inbred lines was due to failure of sibs to mate. When individuals of sterile cultures were allowed an opportunity to outcross, most were fertile and exhibited normal mating and courtship activity. Olfactometer tests with either unrelated flies or collected pheromone samples as source material indicated that airborne chemosignals are required for initiation of courtship inD. melanogaster. Visual and audio cues were found to be noncritical inD. melanogaster courtship and mating. Intraspecific qualitative pheromone variation, at one or a few loci, is thought to be functioning as the control mechanism in selective mating.The investigation was conducted as partial fulfillment of the requirements for the degree of Doctor of Philosophy by W. W. A. while he was supported by PHS Training Grant No. 5T01 GM-00337 from the National Institutes of General Medical Sciences. Other research support was by NSF Grant GB 36103X to R. H. R. The support of NIH Research Career Development Award 5 KO4-GM 47350 to R. H. R. also is acknowledged.     

Mitochondrial genome sequences and molecular evolution of the Irish potato famine pathogen, Phytophthora infestans

The mitochondrial genomes of haplotypes of the Irish potato famine pathogen, Phytophthora infestans, were sequenced. The genome sizes were 37,922, 39,870 and 39,840 bp for the type Ia, IIa and IIb mitochondrial DNA (mtDNA) haplotypes, respectively. The mitochondrial genome size for the type Ib haplotype, previously sequenced by others, was 37,957 bp. More than 90% of the genome contained coding regions. The GC content was 22.3%. A total of 18 genes involved in electron transport, 2 RNA-encoding genes, 16 ribosomal protein genes and 25 transfer RNA genes were coded on both strands with a conserved arrangement among the haplotypes. The type I haplotypes contained six unique open reading frames (ORFs) of unknown function while the type II haplotypes contained 13 ORFs of unknown function. Polymorphisms were observed in both coding and non-coding regions although the highest variation was in non-coding regions. The type I haplotypes (Ia and Ib) differed by only 14 polymorphic sites, whereas the type II haplotypes (IIa and IIb) differed by 50 polymorphic sites. The largest number (152) of polymorphic sites was found between the type IIb and Ia haplotypes. A large spacer flanked by the genes coding for tRNA-Tyr (trnY) and the small subunit RNA (rns) contained the largest number of polymorphic sites and corresponds to the region where a large indel that differentiates type II from type I haplotypes is located. The size of this region was 785, 2,666 and 2,670 bp in type Ia, IIa and IIb haplotypes, respectively. Among the four haplotypes, 81 mutations were identified. Phylogenetic and coalescent analysis revealed that although the type I and II haplotypes shared a common ancestor, they clearly formed two independent lineages that evolved independently. The type II haplotypes diverged earlier than the type I haplotypes. Thus our data do not support the previous hypothesis that the type II lineages evolved from the type I lineages. The type I haplotypes diverged more recently and the mutations associated with the evolution of the Ia and Ib types were identified. Electronic Supplementary Material Supplementary material is available for this article at     

Sexual differences for emigration behavior in natural populations of Drosophila melanogaster

Evolutionary biology considers migration behavior as central in genetic structure of populations and speciation. Here we report on emigration patterns in Drosophila melanogaster behavior under laboratory conditions. For this study, a special apparatus was employed that includes a few important changes in its design and size compared with other known systems. The results presented in this paper were obtained on flies derived from natural populations of two contrasting climatic and geographical regions, from mesic northern and xeric southern parts of Israel. Highly significant difference between sexes in emigration activity was found for both localities. Emigration activity of females appeared to be higher than that of males. We also found that the flies' geographic origin affects emigration behavior (flies from a relatively closed natural system seem to display lower emigration ability than those from an open habitat), although broader sampling from various habitats is needed to confirm these results.     

Male sexual signaling is defective in mutants of theapterous gene ofDrosophila melanogaster

Theapterous (ap) gene ofDrosophila melanogaster exhibits extreme pleiotrophy: its functioning is essential for life, normal wing structure, juvenile hormone production, female fertility, and normal development of female sexual receptivity. Four mutantap alleles (ap ⁴,ap ^56f,ap ^c, andap ^blt) were characterized for three additional phenotypes: male mating success, courtship behavior, and immature male sex appeal (the ability of males to stimulate homosexual cortship). Mating success with mature wild-type virgin females is reduced in males mutant for theap gene, the extreme case beingap ⁴/ap ⁴ males, which are behaviorally sterile. Inap mutants, nonwing courtship elements are qualitatively like those ofap ⁺/ap ⁺ males. However, the mean rate of nonwing courtship directed toward virgin wild-type females (i.e., the mean temporal frequency of these displays) is reduced in males homozygous forap ⁴,ap ^56f, orap ^c alleles. In contrast, theap ^blt allele makes for wild-type rates of nonwing courtship. Immature male sex appeal persists for at least 3 days in males homozygous forap ^c and, to a lesser extent, inap ^56f orap ⁴ homozygotes;ap ^blt/ap ^blt and wild-type males lose immature male sex appeal after 1 day. All three male phenotypes map to theap locus, which is therefore essential for the development of normal levels of male courtship and male mating success and for the timely loss of immature male sex appeal. For each phenotype,ap ⁺ is dominant toap alleles making for behavioral abnormalities, with a single exception (for rate of nonwing courtship,ap ⁺/ap ^c was low). For mating success and frequency of nonwing courtship, each allele pair exhibits at least partial complementation, except forap ⁴ andap ^56f, which fail to complement. For immature male sex appeal,ap ^c,ap ⁴, andap ^56f fall into the same complementation group. Juvenile hormone production is not correlated with effects on male reproductive behavior.This work was supported in part by The Scheinfeld Center for Human Genetics in the Social Sciences (J.R.), The National Science Foundation (BNS-882 1339, to J.R.), BARD (IS-1664-89R, to D.S.), and The Israel Cancer Research Fund (grant to D.S.).     

Phototransduction and retinal degeneration in Drosophila

Drosophila visual transduction is the fastest known G-protein-coupled signaling cascade and has therefore served as a genetically tractable animal model for characterizing rapid responses to sensory stimulation. Mutations in over 30 genes have been identified, which affect activation, adaptation, or termination of the photoresponse. Based on analyses of these genes, a model for phototransduction has emerged, which involves phosphoinoside signaling and culminates with opening of the TRP and TRPL cation channels. Many of the proteins that function in phototransduction are coupled to the PDZ containing scaffold protein INAD and form a supramolecular signaling complex, the signalplex. Arrestin, TRPL, and Gα_q undergo dynamic light-dependent trafficking, and these movements function in long-term adaptation. Other proteins play important roles either in the formation or maturation of rhodopsin, or in regeneration of phosphatidylinositol 4,5-bisphosphate (PIP₂), which is required for the photoresponse. Mutation of nearly any gene that functions in the photoresponse results in retinal degeneration. The underlying bases of photoreceptor cell death are diverse and involve mechanisms such as excessive endocytosis of rhodopsin due to stable rhodopsin/arrestin complexes and abnormally low or high levels of Ca²⁺. Drosophila visual transduction appears to have particular relevance to the cascade in the intrinsically photosensitive retinal ganglion cells in mammals, as the photoresponse in these latter cells appears to operate through a remarkably similar mechanism.     

Diversification of the expression patterns and developmental functions of the dishevelled gene family during chordate evolution

Dishevelled (Dvl) proteins are key transducers of Wnt signaling encoded by members of a multi‐gene family in vertebrates. We report here the divergent, tissue‐specific expression patterns for all three Dvl genes in Xenopus embryos, which contrast dramatically with their expression patterns in mice. Moreover, we find that the expression patterns of Dvl genes in the chick diverge significantly from those of Xenopus. In addition, in hemichordates, an outgroup to chordates, we find that the one Dvl gene is dynamically expressed in a tissue‐specific manner. Using knockdowns, we find that Dvl1 and Dvl2 are required for early neural crest specification and for somite segmentation in Xenopus. Most strikingly, we report a novel role for Dvl3 in the maintenance of gene expression in muscle and in the development of the Xenopus sclerotome. These data demonstrate that the expression patterns and developmental functions of specific Dvl genes have diverged significantly during chordate evolution. Developmental Dynamics 238:2044–2057, 2009. © 2009 Wiley‐Liss, Inc.     

Female sexual receptivity is defective in juvenile hormone-deficient mutants of theapterous gene ofDrosophila melanogaster

During reproductive maturation of female insects, the acquisition of sexual receptivity is coordinated with ovarian development. Juvenile homone regulates vitellogenesis in the ovaries, but the action of this hormone in the development of sexual behavior is less well-understood. A strain ofDrosophila melanogaster carrying a mutation in theapterous gene(ap ⁴) was known to exhibit arrested vitellogenesis (rescuable by applying exogenous juvenile hormone), sterility of both sexes, and a deficiency of juvenile hormone. In this study, we examined the effects of mutations ofap on female receptivity and its relationship to juvenile hormone. We observed abnormally low female receptivity in homozygousap strains, and heteroallelic combinations ofap mutations exhibited low receptivity. For female receptivity,ap showed no dominance (i.e.,ap/ap ⁺ was intermediate betweenap/ap andap ⁺/ap ⁺). Low receptivity mapped genetically to theap locus. The reduction in female receptivity in these mutants is positively correlated with levels of juvenile hormone synthesized by their corpora allata.This work was supported in part by The Scheinfeld Center for Humans Genetics in the Social Sciences (J.R.), The National Science Foundation (BNS-882 1339 to J.R.), BARD (No. IS-1664-89R to D.S.), The Israel Cancer Research Fund (grant to D.S.), The Rekanati Foundation of Tel Aviv University (grant to D.S.), and The Israeli Fruit Council (award to M.A.)     

Restriction of amino acids extends lifespan in Drosophila melanogaster

Dietary restriction extends adult Drosophila melanogaster life span when the concentration of dietary yeast is diluted in a media with abundant carbohydrates. Here we vary the concentration of casein as a source of amino acids in adult diet to uncover a quality of nutrient yeast responsible for longevity control. Longevity is maximized upon diet with intermediary levels of casein. Differences in survival are not caused by elevated age-independent mortality; the longevity maximum at intermediate casein does not arise because casein is non-specifically harmful at higher concentrations. Furthermore, fecundity increases when the level of dietary casein is elevated. The demographic phenotypes of adult Drosophila maintained on intermediate levels of casein resemble their response to limited dietary yeast. Dietary restriction through dilution of yeast may extend longevity because this limits the intake of amino acids.     

Selection for geotaxis inDrosophila melanogaster: Heritability,degree of dominance and correlated responses to selection

Selection for geotaxis was carried out with flies from a natural population ofDrosophila melanogaster; geotactic behavior was measured by means of a Hirsch classification maze. The population was initially almost neutral to gravity, and it responded to both positive (downward) and negative (upward) selection with a realized heritability of about 0.13. Stabilizing selection toward neutral gravity was carried out simultaneously. At generations 6, 9, and 10, all possible hybrid crosses between pairs of the selected populations were generated and tested. The geotactic scores of hybrids in generations 6 and 9 were not significantly different from the midparent values, while the scores of hybrids in generation 10 deviated significantly from the midparent values in the direction of positive geotaxis. The frequencies of polymorphic inversions declined in every population during selection, but the population under neutral selection seemed to maintain a higher chromosomal polymorphism than those under positive or negative selection. There was no significant depression of productivity, measured as number of progeny, in any population during nine generations of selection.This work was supported by USPHS Grant HD-07180 to W. W. A.