Broken chromosomal ends can be elongated by conversion in Drosophila melanogaster

The fate of the termini of X chromosomes broken in the regulatory region of the yellow gene was followed in heterozygotes with X chromosomes carrying a point mutation inactivating the yellow gene. Each generation had a loss of about 70 terminal base pairs from the broken chromosome. However, gene conversion restoring the correct sequence at the chromosomal terminus took place with a frequency of about 1×10^–2 per generation. The average length of the conversion track was 2.7 kb. No recombination events occurred. In addition, we found that the normal functioning of the yellow body and wing enhancers located at the tip of the chromosome required about 4 kb of additional upstream sequence. Received: 8 October 1998; in revised form: 18 January 1999 / Accepted: 18 January 1999     

Enhanced cost of mating in female sterile mutants of Drosophila melanogaster

In Drosophila females, mating is known to cause a reduction in life span, which is referred to as 'the cost of mating'. Since mating enhances oogenesis and oviposition, the cost of mating may be regarded as a trade-off between reproduction and longevity. We examined whether the cost of mating exists in mutant females that are unable to produce eggs. Three different mutant alleles of ovarian tumors (otu) and an allele of dunce (dnc(M11)) of Drosophila melanogaster were used to sterilize females. For all the female sterile mutants tested, mating dramatically decreased the life span of homozygous sterile females. Even more extreme shortening of life spans were observed when the sex peptide gene (Acp70A) was expressed in homozygous otu females, though they were virgin, indicating that the shortening in life span is due to seminal factors. These results indicate that the cost of mating is greater in females defective in oogenesis than that in normally fertile females.     

The effect of mating on starvation resistance in natural populations of Drosophila melanogaster

In nature, behavioural and physiological processes involved in mating may entail different costs and benefits for males and females. However, it has been hypothesized that sexual interactions may have additional costs for Drosophila females like decrease in receptivity to remating and shortening of lifespan. During mating, males transfer seminal fluid proteins to females that exert severe physiological changes that may compromise female’s lifespan and reproductive success. However, under specific stressful environmental conditions that organisms usually face in nature, mating may also confer benefits to females. In the present work, we examine the effect of mating on starvation resistance in wild Drosophila melanogaster. We demonstrate that mated females derived from different geographic locations have the benefit of a greater starvation resistance as compared to virgin females. Even though mating status did not affect mean starvation resistance, we detected a strong genotype-specific effect in males. Beyond the obvious advantage of mating, our study reveals that mating might not be perilous for females, as envisaged by sexual conflict theories, but advantageous for flies exposed to shifts in environmental conditions. Thus, our results highlight the importance of studying other ecologically relevant traits that may contribute to the evolution of male–female interactions.     

Influence of female age, sperm senescence and multiple mating on sperm viability in female Drosophila melanogaster

Sperm viability has been associated with the degree of promiscuity across species, as well as the degree of reproductive success within species. Thus, sperm survival within the female reproductive tract likely plays a key role in how mating systems evolve. In the fruit fly, Drosophila melanogaster, however, the extent and cause of sperm death has been the subject of recent debate. Here, we assess sperm death within the female reproductive tract of D. melanogaster following single and multiple matings in order to elucidate the extent of death and its potential mechanisms, including an acute female response to mating, female age and/or sperm senescence. We found no evidence that sperm viability was influenced by an acute female response or female age. We also found that rival ejaculates did not influence viability, supporting recent work in the system. Instead, the majority of death appears to be due to the aging of male gametes within the female, and that at least some dead resident sperm remain in the female after multiple mating. In contrast to earlier in vivo work, we found that overall sperm death was minimal (8.7%), indicating viability should have a negligible influence on female remating rates.     

Genes regulated by mating, sperm, or seminal proteins in mated female Drosophila melanogaster

In Drosophila melanogaster, sperm and accessory gland proteins ("Acps," a major component of seminal fluid) transferred by males during mating trigger many physiological and behavioral changes in females (reviewed in ). Determining the genetic changes triggered in females by male-derived molecules and cells is a crucial first step in understanding female responses to mating and the female's role in postcopulatory processes such as sperm competition, cryptic female choice, and sexually antagonistic coevolution. We used oligonucleotide microarrays to compare gene expression in D. melanogaster females that were either virgin, mated to normal males, mated to males lacking sperm, or mated to males lacking both sperm and Acps. Expression of up to 1783 genes changed as a result of mating, most less than 2-fold. Of these, 549 genes were regulated by the receipt of sperm and 160 as a result of Acps that females received from their mates. The remaining genes whose expression levels changed were modulated by nonsperm/non-Acp aspects of mating. The mating-dependent genes that we have identified contribute to many biological processes including metabolism, immune defense, and protein modification.     

The effect of sexual harassment on lethal mutation rate in female Drosophila melanogaster

The rate by which new mutations are introduced into a population may have far-reaching implications for processes at the population level. Theory assumes that all individuals within a population have the same mutation rate, but this assumption may not be true. Compared with individuals in high condition, those in poor condition may have fewer resources available to invest in DNA repair, resulting in elevated mutation rates. Alternatively, environmentally induced stress can result in increased investment in DNA repair at the expense of reproduction. Here, we directly test whether sexual harassment by males, known to reduce female condition, affects female capacity to alleviate DNA damage in Drosophila melanogaster fruitflies. Female gametes can repair double-strand DNA breaks in sperm, which allows manipulating mutation rate independently from female condition. We show that male harassment strongly not only reduces female fecundity, but also reduces the yield of dominant lethal mutations, supporting the hypothesis that stressed organisms invest relatively more in repair mechanisms. We discuss our results in the light of previous research and suggest that social effects such as density and courtship can play an important and underappreciated role in mediating condition-dependent mutation rate.     

The frequency dependence of mating success in Drosophila melanogaster

In several competition experiments between various strains of Drosophila melanogaster, a rare female advantage as well as a rare male advantage has been observed. This phenomenon was detected by the use of a method described earlier (Lichtenberger et al., Behav. Genet. 17: 203–210, 1987). Often, the use of less refined indices does not lead to the same conclusion.This paper is dedicated to the memory of Dr Marcel Lichtenberger, M.D., fifteen years medical missionary in China, and seventeen years Professor of Human Genetics in the State University of Saigon, Vietnam.     

The developments between gametogenesis and fertilization: ovulation and female sperm storage in Drosophila melanogaster

In animals with internal fertilization, ovulation and female sperm storage are essential steps in reproduction. While these events are often required for successful fertilization, they remain poorly understood at the developmental and molecular levels in many species. Ovulation involves the regulated release of oocytes from the ovary. Female sperm storage consists of the movement of sperm into, maintenance within, and release from specific regions of the female reproductive tract. Both ovulation and sperm storage elicit important changes in gametes: in oocytes, ovulation can trigger changes in the egg envelopes and the resumption of meiosis; for sperm, storage is a step in their transition from being "movers" to "fertilizers." Ovulation and sperm storage both consist of timed and directed cell movements within a morphologically and chemically complex environment (the female reproductive tract), culminating with gamete fusion. We review the processes of ovulation and sperm storage for Drosophila melanogaster, whose requirements for gamete maturation and sperm storage as well as powerful molecular genetics make it an excellent model organism for study of these processes. Within the female D. melanogaster, both processes are triggered by male factors during and after mating, including sperm and seminal fluid proteins. Therefore, an interplay of male and female factors coordinates the gametes for fertilization.     

Multiple mating in wild Drosophila melanogaster revisited by microsatellite analysis

The occurrence of multiple mating in Drosophila melanogaster is of particular interest to evolutionary biologists, as seminal fluid has some toxic effects for females. Thus, it has been predicted that the number of matings per females should be low. We have tested this prediction with seven highly polymorphic microsatellite loci in inseminated females from a Viennese D. melanogaster population. In contrast to the predicted low number of matings and previous studies in natural populations, we identified the genotypes of four to six different males fertilizing the offspring of each female tested. Potential causes and consequences are discussed.     

The quantitative genetic basis of male mating behavior in Drosophila melanogaster

Male mating behavior is an important component of fitness in Drosophila and displays segregating variation in natural populations. However, we know very little about the genes affecting naturally occurring variation in mating behavior, their effects, or their interactions. Here, we have mapped quantitative trait loci (QTL) affecting courtship occurrence, courtship latency, copulation occurrence, and copulation latency that segregate between a D. melanogaster strain selected for reduced male mating propensity (2b) and a standard wild-type strain (Oregon-R). Mating behavior was assessed in a population of 98 recombinant inbred lines derived from these two strains and QTL affecting mating behavior were mapped using composite interval mapping. We found four QTL affecting male mating behavior at cytological locations 1A;3E, 57C;57F, 72A;85F, and 96F;99A. We used deficiency complementation mapping to map the autosomal QTL with much higher resolution to five QTL at 56F5;56F8, 56F9;57A3, 70E1;71F4, 78C5;79A1, and 96F1;97B1. Quantitative complementation tests performed for 45 positional candidate genes within these intervals revealed 7 genes that failed to complement the QTL: eagle, 18 wheeler, Enhancer of split, Polycomb, spermatocyte arrest, l(2)05510, and l(2)k02206. None of these genes have been previously implicated in mating behavior, demonstrating that quantitative analysis of subtle variants can reveal novel pleiotropic effects of key developmental loci on behavior.     

Seminal proteins but not sperm induce morphological changes in the Drosophila melanogaster female reproductive tract during sperm storage

In most insects, sperm transferred by the male to the female during mating are stored within the female reproductive tract for subsequent use in fertilization. In Drosophila melanogaster, male accessory gland proteins (Acps) within the seminal fluid are required for efficient accumulation of sperm in the female's sperm storage organs. To determine the events within the female reproductive tract that occur during sperm storage, and the role that Acps and sperm play in these events, we identified morphological changes that take place during sperm storage in females mated to wild-type, Acp-deficient or sperm-deficient males. A reproducible set of morphological changes occurs in a wild-type mating. These were categorized into 10 stereotypic stages. Sperm are not needed for progression through these stages in females, but receipt of Acps is essential for progression beyond the first few stages of morphological change. Furthermore, females that received small quantities of Acps reached slightly later stages than females that received no Acps. Our results suggest that timely morphological changes in the female reproductive tract, possibly muscular in nature, may be needed for successful sperm storage, and that Acps from the male are needed in order for these changes to occur.     

Control of female pheromones in Drosophila melanogaster by homeotic genes

We have investigated the role of the Antennapedia and Bithorax complexes (ANT-C and BX-C) on the production of cuticular hydrocarbons in Drosophila melanogaster. In males, there is little, if any, influence of these complexes on the hydrocarbon pattern. In females, there are large and opposite effects of these complexes on diene production: two ANT-C mutations cause an increase in diene production and a reduction of monoenes, whereas most BX-C mutations result in a decrease in dienes and an increase in monoenes, although their sum remains constant. The effect is the highest in Mcp and iab6 females. It is suggested that a factor originating from the prothorax might activate the conversion of monoenes to dienes in females. The abdomen seems to have a crucial role in the production or control of pheromones: abdominal segments four to seven have the main effects, with a most dramatic effect for segments four and five.     

How to overwinter and be a founder: egg-retention phenotypes and mating status in Drosophila melanogaster

In temperate regions, Drosophila melanogaster has perennial overwintering populations. These populations present seasonal variations, under the influence of developmental temperature, for several genetically determined physiological traits. The capacity of virgin females to control ovulation is one of these characteristics. Phenotypes able to postpone egg-laying in the absence of insemination are favored under low temperature development and are numerous in Autumn generations. Moreover, a shift between Autumn and the following Spring has often been observed in favor of these phenotypes. The aim of the present study is to determine the characteristics and situations which confer an advantage on long as compared with short-retention phenotypes, during this non-reproductive period. Several traits were studied: resistance to cold shocks, resistance to long cold periods, developmental duration and viability, longevity and starvation resistance. Long-retention phenotypes (LL) had a longer life expectancy than short-retention phenotypes (ss) under virgin or mated status and greater resistance to starvation, by avoiding waste material (eggs). At 14 °C, flies that had mated once survived for several months on normal substrate with live spermatozoa, and flies on deficient medium (without proteins) survived for more than 3 months varying with phenotype. Flies with the best chance of overwintering are the long-retention phenotypes and some hybrids. The most favorable situation for population restoration is when flies are inseminated once in Autumn rather than when they remain virgin until Spring, because males die sooner than females.     

The genetic analysis of meiosis in female Drosophila melanogaster.

The three major features of meiosis are first synapsis, then exchange, and finally, disjunction of homologous chromosomes; these phenomena occur before pachytene, during pachytene, and after pachytene respectively. The effects of meiotic mutants, or other perturbations, either endogenous or exogenous, on the meiotic process may be assigned tentatively to one of these intervals, based on the earliest discernible abnormality. Thus mutants exhibiting abnormal disjunction and normal exchange affect post-pachytene functions; mutants exhibiting abnormal disjunction and exchange but with ultrastructurally normal appearing synaptonemal complex affect pachytene functions; and mutants with abnormal disjunction, exchange, and synaptonemal complex affect prepachytene functions. This rationale is applied to the temporal seriation of effects of meiotic mutants and chromosomal abnormalities on the meiotic programme.     

FB elements can promote exon shuffling: a promoter-less white allele can be reactivated by FB mediated transposition in Drosophila melanogaster

Foldback (FB) elements are transposable elements found in many eukaryotic genomes; they are thought to contribute significantly to genome plasticity. In Drosophila melanogaster, FBs have been shown to be involved in the transposition of large chromosomal regions and in the genetic instability of some alleles of the white gene. In this report we show that FB mediated transposition of w ^67C23, a mutation that deletes the promoter of the white gene and its first exon, containing the start codon, can restore expression of the white gene. We have characterized three independent events in which a 14-kb fragment from the w ^67C23 locus was transposed into an intron region in three different genes. In each case a local promoter drives the expression of white, producing a chimeric mRNA. These findings suggest that, on an evolutionary timescale, FB elements may contribute to the creation of new genes via exon shuffling.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by G. P. Georgiev     

The effect of ventral nerve cord severance and male castration on female mating behavior, clutch size, and maternal care in the ring-legged earwig

Mating is critical for the expression of oviposition and maternal care in the earwig, Euborellia annulipes; additionally, mating diminishes receptivity to additional mating and promotes a decline in juvenile hormone synthesis at the end of the gonadotrophic cycle (in contrast to most insect species wherein mating stimulates juvenile hormone production). We report here that severance of the ventral nerve cord of virgin females similarly promoted egg deposition and maternal care of eggs, diminished mating receptivity, and elicited a timely decline in juvenile hormone biosynthesis. Mating of intact females to adult males that were castrated as larvae did not abolish oviposition; however, clutch size was reduced, and no eggs developed. Such castrated males had smaller seminal vesicles than did intact males, presumably attributable to lack of sperm in castrated males. In contrast, mating of intact females to males castrated on day 1 of adult life did not reduce clutch size compared with those of sham-operated animals and did not abolish fertilization; in fact, these castrated males produced viable offspring after six matings. These results are consistent with the notion that ventral nerve cord severance mimicked mating in intact animals. Following mating, the ventral nerve cord likely is a conduit to release the brain from inhibiting oviposition and maternal care. The presence of sperm in the spermatheca is not necessary for release of this inhibition but may modulate clutch size.