Localization of ribosomal DNA insertion elements in polytene chromosomes of Drosophila simulans,Drosophila mauritiana and their interspecific hybrids

The locations of the ribosomal DNA (rDNA) insertion elements type I and type II along the polytene chromosomes of three Drosophila species of the melanogaster subgroup-D. simulans, D. mauritiana and D. melanogaster-have been compared. In situ hybridization has shown that the intragenomic distribution of type I as well as of type II insertions is different for these related species. In particular, we have revealed rDNA-free autosomal sites, containing type II element sequences within the D. simulans and D. mauritiana chromosomes. This finding confirms the ability of this type of insertion to transpose, as was demonstrated earlier for Bombyx mori. The appearance of the rDNA not associated with the nucleolar organizers, evident by additional nucleoli, occurred with species-specific frequency. At the same time, for all three species the pattern of such changes (an attachment of the nucleoti to varying sites of the chromosomes and the presence of ectopic contacts between them, a composition of the rDNA repeats in the nucleolar material not integrated at the nucleolar organizer) was similar. The number of additional nucleoti in the hybrid polytene nuclei corresponded to the value of the parental species exhibiting nucleolar replicative dominance.     

Phylogenetic incongruence in the Drosophila melanogaster species group

Drosophila melanogaster and its close relatives are used extensively in comparative biology. Despite the importance of phylogenetic information for such studies, relationships between some melanogaster species group members are unclear due to conflicting phylogenetic signals at different loci. In this study, we use twelve nuclear loci (eleven coding and one non-coding) to assess the degree of phylogenetic incongruence in this model system. We focus on two nodes: (1) the node joining the Drosophila erecta-Drosophila orena, Drosophila melanogaster-Drosophila simulans, and Drosophila yakuba-Drosophila teissieri lineages, and (2) the node joining the lineages leading to the melanogaster, takahashii, and eugracilis subgroups. We find limited evidence for incongruence at the first node; our data, as well as those of several previous studies, strongly support monophyly of a clade consisting of D. erecta-D. orena and D. yakuba-D. teissieri. By contrast, using likelihood based tests of congruence, we find robust evidence for topological incongruence at the second node. Different loci support different relationships among the melanogaster, takahashii, and eugracilis subgroups, and the observed incongruence is not easily attributable to homoplasy, non-equilibrium base composition, or positive selection on a subset of loci. We argue that lineage sorting in the common ancestor of these three subgroups is the most plausible explanation for our observations. Such lineage sorting may lead to biased estimation of tree topology and evolutionary rates, and may confound inferences of positive selection.     

Genetics of esterases in Drosophila. IV. Slow-migrating S-esterase in Drosophila of the virilis group

A slow-migrating -esterase (S-esterase) is described which has been detected in Drosophila montana, Drosophila imeretensis, and some stocks of Drosophila virilis when mixtures of - and -naphthyl acetate are used as substrates in histochemical reactions after electrophoresis. Sexual dimorphism for S-esterase has been demonstrated. This esterase is contained in male genitalia only, predominantly in the ejaculatory bulb (waxy plug). It appears 3–4 days after emergence of flies. In hybrids between S⁺ and S⁰ species, the activity of the slow esterase is either decreased or inhibited. An autonomous synthesis of the S-esterase in the ejaculatory bulb was established by transplantation of imaginal genital discs into larvae of different Drosophila stocks. Based on analysis of physicochemical and immunochemical properties, S-esterase is suggested to be an independent fraction of esterase, possibly dimeric, which does not cross-react with -esterase antiserum.     

Drosophila do not skew offspring sex ratio as predicted by trivers and willard (Diptera: Drosophilidae)

Based on both previously published literature and results reported here, it appears thatDrosophila melanogaster meet the explicit assumptions of the Trivers and Willard offspring sex allocation model. However, contrary to the model's predictions, offspring sex ratio was not significantly affected when we manipulated factors that influence offspring quality. We suggest that contrary to implicit predictions of offspring sex ratio models,Drosophila may lack the genetic plasticity to readily alter sex ratio.     

Stage-specific sex differences in Drosophila immunity to parasites and pathogens

Arguments from life-history theory predict that other things being equal females are likely to invest more in defence against parasites and pathogens than males. This is either because males and females differ in behaviour or, more importantly, because the variance in mating success is typically higher in males than in females. Such effects are likely to be most pronounced in those developmental stages where sex differences are greatest. In most organisms, but especially in holometabolous insects, this will be the adult stage. We explored sex-specific resistance to four natural enemies of Drosophila melanogaster that attack the insect at different developmental stages: the larval parasitoid Asobara tabida, the pupal parasitoid Pachycrepoideus vindemiae; and the adult pathogens Beauvaria bassiana (a fungus) and Tubulinosema kingi (a microsporidian). Measures of resistance were designed to reflect the four species’ natural history. Female larvae were able to defend themselves more strongly against A. tabida than males and there was weak evidence that adult females suffered less from microsporidian attack than males. No differences were found for the other two species. Our results provide some support for lower investment in defences in males, and we discuss why the strongest effect was found at the larval rather than the adult stage contrary to our prediction.     

Autosomal control of lampbrush-loop formation during spermatogenesis in Drosophila hydei by a gene also affecting somatic sex determination

Summary The Y chromosome of Drosophila hydei carries information that is necessary for the development of the spermatozoa. In primary spermatocytes Y chromosomal genes become active: five of the male fertility factors form giant lampbrush loops. Our prior work indicated interactions between the Y chromosomal genes and autosomal loci. It is of interest to identify loci regulating the activity of the Y chromosomal genes. We, therefore, screened a total of about 14,000 chromosomes (X, 2, 3 and 4) for mutations that interfere with the expression of the lampbrush loops. Two mutations with substantial effects on the loop morphology were recovered. One of them, a recessive male sterile mutation (ms (3) 5) on chromosome 3, is described in this paper. Its homozygous state results in a complete absence of all Y chromosomal lampbrush loops at 26° C; at 18° C the loops are formed. Temperature shifts with homozygous males indicate that the function early during the spermatogonial stage is crucial for the development of lampbrush loops in the primary spermatocyte. Meiosis is entirely absent in the male, but normal in females. Females homozygous for ms (3) 5 display a maternal effect, which reduces the viability and fertility of homozygous daughters and produces sons with signs of intersexuality. Linkage studies indicated that the effect on the male germ line and the maternal effects cannot be separated and may hence be induced by a single gene.     

Basal relationships in the Drosophila melanogaster species group

The Drosophila melanogaster species group is a popular model for evolutionary studies due to its morphological and ecological diversity and its inclusion of the model species D. melanogaster. However, phylogenetic relationships among major lineages within this species group remain controversial. In this report, the phylogeny of 10 species representing each of the well-supported monophyletic clades in the melanogaster group was studied using the sequences of 14 loci that together comprise 9493 nucleotide positions. Combined Bayesian analysis using gene-specific substitution models produced a 100% credible set of two trees. In the strict consensus of these trees, the ananassae subgroup branches first in the melanogaster species group, followed by the montium subgroup. The remaining lineages form a monophyletic clade in which D. ficusphila and D. elegans branch first, followed by D. biarmipes, D. eugracilis, and the melanogaster subgroup. This strongly supported phylogeny resolves most basal relationships in the melanogaster species group, and provides a framework that can be extended in the future to encompass more species.     

Evolution of the transposable element mariner in the Drosophila melanogaster species group

The population biology and molecular evolution of the transposable element mariner has been studied in the eight species of the melanogaster subgroup of the Drosophila subgenus Sophophora. The element occurs in D. simulans, D. mauritiana, D. sechellia, D. teissieri, and D. yakuba, but is not found in D. melanogaster, D. erecta, or D. orena. Sequence comparisons suggest that the mariner element was present in the ancestor of the species subgroup and was lost in some of the lineages. Most species contain both active and inactive mariner elements. A deletion of most of the 3 end characterizes many elements in D. teissieri, but in other species the inactive elements differ from active ones only by simple nucleotide substitutions or small additions/deletions. Active mariner elements from all species are quite similar in nucleotide sequence, although there are some-species-specific differences. Many, but not all, of the inactive elements are also quite closely related. The genome of D. mauritiana contains 20–30 copies of mariner, that of D. simulans 0–10, and that of D. sechellia only two copies (at fixed positions in the genome). The mariner situation in D. sechellia may reflect a reduced effective population size owing to the restricted geographical range of this species and its ecological specialization to the fruit of Morinda citrifolia.     

High mortality and sex ratio imbalance in a critically declining Oriental White-backed Vulture population (Gyps bengalensis) in Pakistan

Populations of Oriental White-backed Vulture (Gyps bengalensis) and Long-billed Vulture (Gyps indicus) declined dramatically by 95–100% on the Indian subcontinent in during the mid-1990s. The study reported here was conducted in Pakistan to compare the population size, breeding success, patterns of mortality and sex ratios among dead vultures and newly hatched nestlings of G. bengalensis and G. indicus at Toawala (TW) and Nagar Parkar (NP), respectively, during the breeding seasons 2005/2006 and 2006/2007. At TW, diclofenac poisoning was most likely responsible for the high mortality and sex ratio imbalance among dead G. bengalensis, where vulture counts and breeding success declined quickly during the study period. However, at NP no significant difference in population size, breeding success and annual mortality of G. indicus was recorded during the study period. A sex ratio imbalance was detected among nestlings of G. bengalensis, with 68% males and 32% females. In contrast, the sex ratio did not differ significantly in G. indicus.     

A study of wing morphology and fluctuating asymmetry in interspecific hybrids between Drosophila buzzatii and D. koepferae

In this work we investigate the effect of interspecific hybridization on wing morphology using geometric morphometrics in the cactophilic sibling species D. buzzatii and D. koepferae. Wing morphology in F₁ hybrids exhibited an important degree of phenotypic plasticity and differs significantly from both parental species. However, the pattern of morphological variation between hybrids and the parental strains varied between wing size and wing shape, across rearing media, sexes, and crosses, suggesting a complex genetic architecture underlying divergence in wing morphology. Even though there was significant fluctuating asymmetry for both, wing size and shape in F₁ hybrids and both parental species, there was no evidence of an increased degree of fluctuating asymmetry in hybrids as compared to parental species. These results are interpreted in terms of developmental stability as a function of a balance between levels of heterozygosity and the disruption of coadaptation as an indirect consequence of genomic divergence.     

Nucleotide divergence of the rp49 gene region between Drosophila melanogaster and two species of the Obscura group of Drosophila

Summary A 2.1-kb SStI fragment including the rp49 gene and the 3 end of the -serendipity gene has been cloned and sequenced in Drosophila pseudoobscura. rp49 maps at region 62 on the tip of chromosome II of this species. Both the coding and flanking regions have been aligned and compared with those of D. subobscura. There is no evidence for heterogeneity in the rate of silent substitution between the rp49 coding region and the rate of substitutions in flanking regions, the overall silent divergence per site being 0.19. Noncoding regions also differ between both species by different insertions/deletions, some of which are related to repeated sequences. The rp49 region of D. pseudoobscura shows a strong codon bias similar to those of D. subobscura and D. melanogaster. Comparison of the rates of silent (K _S ) and nonsilent (K _a ) substitutions of the rp49 gene and other genes completely sequenced in D. pseudoobscura and D. melanogaster confirms previous results indicating that rp49 is evolving slowly both at silent and nonsilent sites. According to the data for the rp49 region, D. pseudoobscura and D. subobscura lineages would have diverged some 9 Myr ago, if one assumes a divergence time of 30 Myr for the melanogaster and obscura groups.Offprint requests to: C. Segarra     

Genetics and the environment in interspecific competition: a study using the sibling species Drosophila melanogaster and Drosophila simulans

The outcome of interspecific competition of two closely related species may depend upon genetic variation in the two species and the environment in which the experiment is carried out. Interspecific competition in the two sibling species, Drosophila melanogaster and D. simulans, is usually investigated using longterm laboratory stocks that often have mutant markers that distinguish them. To examine competition in flies that genetically more closely resemble flies in nature, we utilized freshly caught wildtype isofemale lines of the two species collected at the same site in San Carlos, Mexico. Under ordinary laboratory conditions, D. melanogaster always won in competition. However, in hotter and drier conditions, D. simulans competed much more effectively. In these environmental conditions, there were genetic differences in competitive ability among lines with the outcome of competition primarily dependent upon the line of D. melanogaster used but in some cases also influenced by the line of D. simulans used. Differences in the measures of productivity and developmental time did not explain the differences in competitive ability among lines. This suggests that the outcome of competition was not due to differences in major fitness components among the isofemale lines but to some other attribute(s) that influenced competitive ability. When lines of flies were combined, the outcome of competition was generally consistent with competitive outcomes between pairs of lines. In several cases, the combination of lines performed better than the best of the constituent lines, suggesting that competitive ability was combined heterotically and that the total amount of genetic variation was important in the outcome of interspecific competition.     

Parental Imprinting in Drosophila

Genetic imprinting is a form of epigenetic silencing. But with a twist. The twist is that while imprinting results in the silencing of genes, chromosome regions or entire chromosome sets, this silencing occurs only after transmission of the imprinted region by one sex of parent. Thus genetic imprinting reflects intertwined levels of epigenetic and developmental modulation of gene expression. Imprinting has been well documented and studied in Drosophila, however, these studies have remained largely unknown due to nothing more significant than differences in terminology. Imprinting in Drosophilais invariably associated with heterochromatin or regions with unusual chromatin structure. The imprint appears to spread from imprinted centers that reside within heterochromatin and these are, seemingly, the only regions that are normally imprinted in Drosophila. This is significant as it implies that while imprinting occurs in Drosophila, it is generally without phenotypic consequence. Hence the evolution of imprinting, at least in Drosophila, is unlikely to be driven by the function of specific imprinted genes. Thus, the study of imprinting in Drosophilahas the potential to illuminate the mechanism and biological function of imprinting, and challenge models based solely on imprinting of mammalian genes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Heterosis in crosses between lines of Drosophila melanogaster selected for adaptation to different environments

Summary Experiments were designed to examine whether heterosis would occur in crosses of Drosophila melanogaster populations adapted to 18 °C or 28 °C environments. Crosses were examined in parental environments, an intermediate environment (23 °C) and a mixed environment (alternating 18°/28°C). Parental populations did not show divergence for larval viability, cold shock or high temperature mortalities when tested in a common environment. However, the 28 °C population was less fecund than the 18 °C population, but had higher larval competitive ability and higher adult longevity. Heterosis for viability, cold shock mortality and high temperature mortality occurred in crosses between a population adapted to 18 °C and another adapted to 28 °C, but not in crosses between two populations adapted to the same temperature. The results suggest that, in the absence of drift, heterosis is expected in crosses between lines or populations with different histories of selection but not between lines with the same selection histories.     

Energy storage during reproductive diapause in the Drosophila melanogaster species group

Summary Temperate species of the Drosophila melanogaster group enter reproductive diapause for overwintering in response to short daylength. During the prediapause period they accumulate triacylglycerols, but not glycogen, as energy resources. The capacity for storing triacylglycerols differs between species, and appears to be closely correlated with diapause and cold-hardiness; cool-temperate species, such as those of the auraria species complex, which enter a deep diapause and are highly cold-hardy, accumulate larger quantities of triacylglycerols than warm-temperate species, such as D. rufa and D. lutescens, which enter a weak diapause and are less cold-hardy. Among the cool-temperate spcies, D. subauraria occurs at a higher latitude and has the greatest capacity for accumulating triacylglycerols. A subtropical species, D. takahashii, which has no diapause in nature and is not cold-hardy, is unable to store the same quantities of triacylglycerols as temperate species.     

In Situ Hybridisation Analysis of the X-Linked Genes in the Species of the Virilis Group of Drosophila

When estimating the level of DNA sequence variation within and between populations or when planning QTL analysis, it is essential to know the location of the genes under study. In the present work, five X chromosomal genes, earlier localised in Drosophila virilis and D. littoralis, were mapped by in situ hybridisation on the larval polytene chromosomes of four other virilis group species, D. a. americana, D. flavomontana, D. lacicola and D. montana. Conjugation of X chromosomes of the most interesting species pairs was studied in interspecific hybrids. Three of the marker genes were used as RFLP markers to examine the occurrence of recombination in D. flavomontana and D. montana hybrid females. The gene arrangement of all species studied, appeared to be different at the proximal end of the X chromosome, which prevented normal conjugation along the most part of the X chromosome. The data illustrating the locations of five X chromosomal marker genes are presented for D. a. americana, D. flavomontana, D. lacicola and D. montana.     

Clonal analysis in hybrids between Drosophila melanogaster and Drosophila simulans

We have analysed the viability of cellular clones induced by mitotic recombination in Drosophila melanogaster/D. simulans hybrid females during larval growth. These clones contain a portion of either melanogaster or simulans genomes in homozygosity. Analysis has been carried out for the X and the second chromosomes, as well as for the 3L chromosome arm. Clones were not found in certain structures, and in others they appeared in a very low frequency. Only in abdominal tergites was a significant number of clones observed, although their frequency was lower than in melanogaster abdomens. The bigger the portion of the genome that is homozygous, the less viable is the recombinant melano-gaster/simulans hybrid clone. The few clones that appeared may represent cases in which mitotic recombination took place in distal chromosome intervals, so that the clones contained a small portion of either melanogaster or simulans chromosomes in homozygosity. Moreover, Lhr, a gene of D. simulans that suppresses the lethality of male and female melanogaster/simulans hybrids, does not suppress the lethality of the recombinant melanogaster/simulans clones. Thus, it appears that there is not just a single gene, but at least one per tested chromosome arm (and maybe more) that cause hybrid lethality. Therefore, the two species, D. melanogaster and D. simulans, have diverged to such a degree that the absence of part of the genome of one species cannot be substituted by the corresponding part of the genome of the other, probably due to the formation of co-adapted gene complexes in both species following their divergent evolution after speciation. The disruption of those coadapted gene complexes would cause the lethality of the recombinant hybrid clones.     

Natal dispersal in relation to population density and sex ratio in the field vole,Microtus agrestis

Summary In a sample of 240 juvenile field voles 8% of the males and 22% of the females reached sexual maturity within their natal home range. Among individuals retrapped as adults, 58% of males and 23% of females had dispersed, i.e. had moved more than one home range diameter. The mean distance moved for males (58.5 m) exceeded that for females (28.6 m). Male movement distances were negatively associated with total density, and with density of adult females, but not with male density. Female movements were not related to population density. There were no relation between sex ratio and distance moved. The distribution of distances moved for both males and females fit a geometrical distribution, suggesting the importance of competitive processes.     

Growth and sex ratio of nestlings in two species of crows: how important is hatching asynchrony?

Summary In experimental studies of avian hatching paterns offspring sex has been neglected. This may be a problem if nestling growth and mortality is sex biased, and if this bias is influenced by hatching spread. In a field study of two crow species, the magpie Pica pica and the hooded crow Corvus corone cornix, we manipulated hatching spread. Both species have asynchronous hatching, and adult males are larger than females by 12–14%. The sex ratios obtained from the different experimental groups on day 24 post-hatch (total sample n = 403) did not deviate significantly from unity, nor did the sex ratios obtained among young newly hatched in an incubator (total sample n = 305). Male and female offspring were of similar size at hatching but males were larger on day 24 post-hatch. Males seemed to be more costly to rear than females, judging by the 20% difference in the mean amounts of food found in the gizzards of the young on day 24 post-hatch. Dimorphism in body size did not seem to be influenced by degree of hatching spread. Asynchronous hatching did not seem to be needed to produce high quality offspring of the larger sex (i.e. males), nor did asynchronous hatching help to ensure equal parental investment in male and female progeny. One reason for the latter negative results may be that the size dimorphism of the two crow species studied were relatively small.