Interspecific competition between Drosophila melanogaster and Drosophila simulans: Effects of adult density on adult viability

Survival (viability) of newly eclosed adults of D. simulans st and D. melanogaster Or R-C, which had excess quantities of dead yeast available throughout life, was measured daily for 7 days in an experiment where adult density (6 levels), species frequency (6 levels) and ³²P tissue content (2 levels) were varied factorially. A separate experiment comparing viability in different types of experimental unit also was done. Similar experiments were done for D. simulans st competing against D. melanogaster y w. The entire data were subjected to a least-squares (unbalanced, missing-plot) analysis of variance. D. melanogaster Or R-C had a higher average viability than D. melanogaster y w or D. simulans st, which were equal. The competing strain of D. melanogaster influenced D. simulans st viability — viability being higher when y w was the competitor. Viability decreased over the seven day period but at different rates for the three strains. Increasing density reduced viability for all three strains, but species frequency effects, although significant, were generally not consistent. Females had higher viability than males in both D. melanogaster strains, but the reverse was true in D. simulans st. ³²P lowered viability and experimental unit type altered viability. Numerous interactions were significant.Adult density was shown to have a delayed effect on viability — the delay before the appearance of the effect (an increase in death rate) being decreased as density rose. The term variably delayed density dependent has been adopted to describe the fitness component, adult viability. Some high density populations showed a readjustment (a decrease) in their death rate as a reaction to the effect of reduced density caused by high early mortality.     

Interspecific competition in Drosophila II. Competitive outcome in some 2–resource environments

Populations of Drosophila metanogaster and D. simulans were made to compete in a number of 2– resource environments. The resources used were all based on a standard Drosophila medium and differed from each other only in their concentrations of ethanol. Each experiment comprised three cages started at a species–frequency of 0.8 D. melanogaster and three at 0.2 D. melanogaster. This enabled screening for a wide range of equilibria.
Reversal of competitive superiority was achieved (unintentionally) by raising the alcohol level of one of the resources. Thus while D. simulans won in an environment with media containing 0 and 896 ethanol, D. melanogaster won if the ethanol concentrations were 0 and 10%, or 0 and 12%. This parallels the reversal of dominance reported earlier for single–resource systems. However, in none of the 2–resource systems studied was there a stable equilibrium in species–frequency–in fact there was no significant frequency–dependence at all.
An artificial selection experiment produced a stock of D. melanogaster ovipositing a higher proportion of eggs on the alcoholic resource. Thus, this stock was more divergent from D. simulans , in at least one aspect of resource use, than was the original stock of D. melanogaster. However, competition between stock D. simulans and selected D. melanogaster showed no frequency–dependence. Again, reversal of competitive dominance was unintentionally achieved.
The implications of these results are discussed, and the need for a comprehensive study of resource–utilization in a pair of competing species is stressed.     

The LSP1-α gene is not dosage compensated in the Drosophila melanogaster species subgroup

The X-linked subunit of larval serum protein 1 (LSP1-) is shown to lack dosage compensation in six members of the melanogaster species subgroup, viz., Drosophila melanogaster, D. simulans, D. mauritiana, D. erecta, D. yakuba, and D. teissieri, by quantitative filter hybridization and by electrophoretic and autoradiographic analyses of fat body proteins. These results support the hypothesis that there is little selection pressure on the LSP1- gene to acquire dosage compensation.H.W.B. acknowledges a Commonwealth Scholarship. This work was supported by an SRC grant to D.B.R.     

The impact of anaesthetic technique on survival and fertility in Drosophila

Abstract The consequences of ice and carbon dioxide anaesthetics on the survival and reproductive success of Drosophila simulans and Drosophila melanogaster were investigated after observations of high levels of mortality in D. simulans, possibly associated with brief chill coma. The association between brief chill coma and death is confirmed in female D. simulans but there is no mortality in male D. simulans or D. melanogaster of either sex. Mortality is unlikely to be associated with a strain specific cold intolerance because two geographically distinct populations of D. simulans were examined. In addition to the effect of ice anaesthesia, anaesthetizing newly‐eclosed male D. simulans with CO₂ causes a reduction in fertility, which is evident 9–13 days after anaesthesia. This finding is important given that CO₂ anaesthesia is a standard technique used in Drosophila and other insect cultures, and may have important consequences for studies of male fertility and sperm competition.     

Transplantation of nuclei between eggs of different species ofDrosophila

Summary Interspecific chimeras have been produced by nuclear transplantation inDrosophila. The following species were used:Drosophila melanogaster, D. simulans, D. mauritiana, D. teissieri, D. yakuba, D. erecta andD. ananassae.Nuclei transplantated into fertilized eggs were able to multiply in a foreign cytoplasm and heterologous cells become integrated into the embryo to give viable adult chimeras.The morphological pattern of differentiation was autonomous both from that of the host and donor. In some cases, a possible non-compatibility between nuclei and cytoplasm has been postulated to explain the lack of chimeras.     

High level of divergence of male-reproductive-tract proteins, between Drosophila melanogaster and its sibling species, D. simulans

We compared male-reproductive-tract polypeptides of Drosophila melanogaster and D. simulans by using two-dimensional gel electrophoresis. Approximately 64% of male-reproductive-tract polypeptides were identical between two randomly chosen isofemale lines from these two species, compared with 83% identity for third-instar imaginal wing-disc polypeptides. Qualitatively similar differences were found between reproductive tracts and imaginal discs when D. sechellia was compared with D. melanogaster and with D. simulans. When genic polymorphism was taken into account, approximately 10% of male- reproductive-tract polypeptides were apparently fixed for different alleles between D. melanogaster and D. simulans; this proportion is the same as that found for soluble enzymes by one-dimensional gel electrophoresis. Strikingly, approximately 20% of male-reproductive- tract polypeptides of either D. melanogaster or D. simulans had no detectable homologue in the other species. We propose that proteins of the Drosophila male reproductive tract may have diverged more extensively between species than have other types of proteins and that much of this divergence may involve large changes in levels of polypeptide expression.      

Antifungal activity of cysteine,its effect on C-21 oxygenated lanosterol derivatives and other lipids in Inonofus obliquus,in vitro

The antifungal activity of cysteine (2-amino-3-mercaptopropionic acid) on mycelial growth and the production of lanosterol, 3,-hydroxy-lanosta-8,24-dien-21-al, 3,-21,dihydroxy-lanosta-8,24-diene, trametenolic acid and ergosterol were investigated in solid and liquid cultures ofInonotus obliquus at cysteine concentrations of 0.9 and 9.0 mm by means of thinlayer chromatography densitometry. Cysteine elicited the production of ergosterol in all the cultures and at a concentration of 0.9 mm distinctly elicited the production of 3,-hydroxy-lanosta-8,24-dien-21-al. At 9.0 mm, cysteine had greater antifungal activity than at 0.9 mm, decreasing growth and the production of metabolities.     

Microsatellite analysis of Drosophila melanogaster populations along a microclimatic contrast at lower Nahel Oren canyon, Mount Carmel, Israel

Drosophila melanogaster populations collected at the south-facing slope (SFS) and north-facing slope (NFS) of lower Nahel Oren canyon, Mount Carmel, Israel display significant differences in survival and longevity at temperature, drought, and starvation stresses. Furthermore, significant assortative mating was previously observed between populations of the two slopes. We used a set of 48 microsatellite markers to analyze patterns of genetic differentiation between D. melanogaster populations from both slopes and D. simulans. Consistent with previous reports, we found D. simulans to be well differentiated from D. melanogaster. Genetic differentiation between SFS and NFS D. melanogaster populations was low (F(ST) = 0.0012). Also a tree of individuals based on the proportion of shared alleles and a model-based clustering method provided no evidence for population substructuring.     

Courtship inDrosophila sechellia: Its structure,functional aspects,and relationship to those of other members of theDrosophila melanogaster species subgroup

The courtship behavior of Drosophila sechelliais described. Male wing displays are mainly vibration and scissoring, with low levels of rowing. As courtship proceeds the proportion of courtship spent in male wing vibration and licking increases, whereas female movement decreases. The male courtship song of sechelliacontains pulse song but no sine song. This species also shows a distinctive copulation song associated with mounting and copulation. The main cuticular hydrocarbon in females is 7,11-heptacosadiene. The number of copulations increased when flies were placed in the presence of food. Visual and acoustic stimuli appear to be important for mating. A multidimensional comparison was used to compare members of the melanogaster species subgroup, based upon courtship behavior, song characteristics, and cuticular hydrocarbons. A multidimensional comparison of courtship sequences in sechellia, melanogaster, simulans,and mauritianashowed differences in variability between the two island species as compared to the two cosmopolitan species. The courtship song of D. orenais described: it shows both sine and pulse song; there is also a copulation song in this species.     

Defence strategies against a parasitoid wasp in Drosophila: fight or flight?

Hosts may defend themselves against parasitism through a wide variety of defence mechanisms, but due to finite resources, investment in one defence mechanism may trade-off with investment in another mechanism. We studied resistance strategies against the parasitoid wasp Leptopilina boulardi in two Drosophila species. We found that D. melanogaster had significantly lower physiological resistance against L. boulardi than D. simulans, and hypothesized that D. melanogaster might instead invest more heavily in other forms of defence, such as behavioural defence. We found that when given a choice between clean oviposition sites and sites infested with wasps, both D. melanogaster and D. simulans detected and avoided infested sites, which presumably limits later exposure of their offspring to infection. Unlike D. simulans, however, D. melanogaster laid significantly fewer eggs than controls in the forced presence of wasps. Our findings suggest that D. melanogaster relies more heavily on behavioural avoidance as defence against wasp parasitism than D. simulans, and that this may compensate for a lack of physiological defence.     

Effects of aggregation pheromone on individual behaviour and food web interactions: a field study on Drosophila

Abstract.  1. The effects of an aggregation pheromone on individual behaviour and food web interactions were investigated in two ecological communities, using Drosophila melanogaster and D. simulans as focal species.
2. Fruit substrates with aggregation pheromone were significantly more attractive to adult D. melanogaster and D. simulans than control fruit substrates, and the response was positively dose dependent. Competing species and natural enemies were also significantly attracted to substrates with the aggregation pheromone of D. melanogaster and D. simulans .
3. Significantly more eggs were deposited on pheromone-treated fruits than on control fruits, and the microdistribution of eggs within fruits was correlated to the microdistribution of the pheromone. The aggregation pheromone induced more females to share the breeding site.
4. The extremely high densities of fruit flies in the large aggregations appeared to reduce the oviposition rate of females. Physical interactions with conspecific and heterospecifics were frequently observed in the aggregations, and often led to patch leaving of the fruit flies.
5. Competition for food among larvae occurred at high densities and parasitism was density dependent. Aggregation pheromones can be directly responsible for these patterns through their effects on the con- and heterospecific behaviour.
6. The combined results show that aggregation pheromones affect a multitude of aspects in the ecology of interacting animals. The importance of incorporating the communication signals in ecological theory of aggregations is discussed.     

Nucleotide variation in the triosephosphate isomerase (Tpi) locus of Drosophila melanogaster and Drosophila simulans

DNA sequence variation in a 1.1-kb region including the coding portion of the Tpi locus was examined in 25 homozygous third-chromosome lines of Drosophila melanogaster, nine lines of Drosophila simulans, and one line of Drosophila yakuba. Our data show that the widespread allozyme polymorphism observed in cosmopolitan D. melanogaster is due to a glutamic acid substitution occurring in a phylogenetically conserved lysine that has been identified as part of the "hinged-lid" active site of the enzyme. This observation suggests that the replacement polymorphism may have important functional consequences. One replacement polymorphism was also observed in D. simulans, although its functional relevance is more difficult to assess, since it affects a site that is not strongly conserved. This amino acid change in D. simulans is associated with a single lineage possessing seven unique silent substitutions, which may be indicative of balancing selection or population subdivision. The absence of fixed amino acid differences between D. melanogaster and D. simulans and only a single difference with D. yakuba suggests that triose phosphate isomerase is under strong functional constraint. Silent variation is slightly higher for D. melanogaster than for D. simulans. Finally, we outline the general lack of evidence for old balanced polymorphisms at allozyme loci in D. melanogaster.      

Reproductive effort of winkles (Littorina spp.) with contrasted methods of reproduction

Summary Reproductive effort was compared in Littorina rudis (ovoviviparous), L. nigrolineata (benthic eggs with direct development), L. neritoides and L. littorea (planktonic eggs and larvae). Three indices of reproductive effort were used: the proportion of total production committed to reproduction per unit time, the cumulative proportion of total production committed to reproduction up to a given age, and the ratio of reproductive production per unit time to somatic biomass at the beginning of the unit time interval. The indices were plotted against age and the snails ranked in order of their reproductive efforts at equivalent ages. When plotted on axes of absolute time, all three indices ranked the snails in the order L. neritoides. The rank order of reproductive efforts on a time axis standardized for growth rate was L. littorea, and on a time axis standardized for generation time was L. neritoidesL. rudis. The implications and relative merits of the different expressions of reproductive effort are discussed. No general relationship was found between the magnitude of reproductive effort and either reproductive type or population ecology among the Littorina spp., or among other intertidal prosobranchs reviewed from the literature. Semelparous species had among the highest reproductive efforts, but there was no clear separation in this respect from certain iteroparous species. Parental investment, measured as the energy content of an egg plus any accompanying jelly or capsular material, was one to two orders of magnitude greater in the species with direct development than in those with indirect development. There were significant differences between egg and hatchling sizes of L. rudis from three contrasted types of shore and these differences were thought to be correlated with the intensity of desiccation and predation on hatchlings rather than with demographic factors.     

Inter‐specific competition and competition‐free space in the tephritid parasitoids Utetes anastrephae and Doryctobracon areolatus (Hymenoptera: Braconidae: Opiinae)

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Chemical cues from competitors change the oviposition preference of Drosophila suzukii

Although Drosophila suzukii Matsumura (Diptera: Drosophilidae) infests fresh and ripening fruits, it is attracted to fermented fruits as well. Because fermented fruits attract other flies too, if D. suzukii utilizes fermented fruits as oviposition substrates, competition can be more intense on them. To avoid such competition, D. suzukii may change oviposition preference when particular species of competitor flies are present, but the effect of odor cues associated with competitors on the oviposition preference of D. suzukii is still unknown. To examine such an effect, we investigated the oviposition preference of D. suzukii in the presence of four competitor fly species – Drosophila melanogaster Meigen, Drosophila lutescens Malloch, Drosophila rufa Kikkawa & Peng, and Drosophila auraria Peng – and D. suzukii itself. We prepared artificial substrates with yeast treatment (Y⁺: yeast supplementation, Y⁻: control) and competitor fly treatment (F⁺: pre-inoculated with competitor fly odor, F⁻: control), and performed two-choice experiments using the substrates with various Y and F treatments. Our results showed that D. suzukii oviposited more eggs on Y⁺ substrates than on Y⁻ substrates when no competitor flies were present and the presence of competitor flies influenced D. suzukii’s oviposition preference for yeast-supplemented substrates and its effect changed depending on the competitor fly species. If the presence of competitors around fallen fruits on the ground suppresses D. suzukii’s oviposition on the fallen fruits and facilitates the oviposition on non-fermenting substrates in nature, it may drive D. suzukii to use ripening fruits on the tree. Such selective pressure may facilitate the evolution of morphological traits such as a serrated ovipositor in D. suzukii.     

African Drosophila melanogaster and D. simulans populations have similar levels of sequence variability, suggesting comparable effective population sizes

Drosophila melanogaster and D. simulans are two closely related species with a similar distribution range. Many studies suggested that D. melanogaster has a smaller effective population size than D. simulans. As most evidence was derived from non-African populations, we readdressed this question by sequencing 10 X-linked loci in five African D. simulans and six African D. melanogaster populations. Contrary to previous results, we found no evidence for higher variability, and thus larger effective population size, in D. simulans. Our observation of similar levels of variability of both species will have important implications for the interpretation of patterns of molecular evolution.     

Evolution of a desaturase involved in female pheromonal cuticular hydrocarbon biosynthesis and courtship behavior in Drosophila

Drosophila species exhibit polymorphism in female pheromonal cuticular hydrocarbons, with 7-monoenes produced in Drosophila simulans and 7,11-dienes in most populations of Drosophila melanogaster (5,9-dienes in several African populations). A female-biased desaturase, desatF, expressed only in D. melanogaster is involved in the synthesis of 7,11-dienes. We investigated the role of desatF in 5,9-diene flies. We constructed a 5,9-diene strain knock-down for desatF and showed that desatF is involved in 5,9-diene formation. We also studied D. melanogaster/D. simulans hybrids. These hybrid females produced dienes and received normal courtship from D. melanogaster males, but copulation success was reduced. With D. simulans males, courtship was decreased and no copulation occurred. Hybrids with a chromosomal deletion of the D. melanogaster desatF gene had no dienes and received normal courtship from D. simulans males; depending on the D. simulans parental strain, 7-19% of them succeeded in mating. D. simulans desatF promoter region shows 21-23% gaps and 86-89% identity with D. melanogaster promoter region, the coding region 93-94% identity, depending on the strain. These differences could explain the functional polymorphism of desatF observed between both species, contributing to different cuticular hydrocarbon profiles, that constitute an effective barrier between species.     

Wolbachia-mediated sperm modification is dependent on the host genotype in Drosophila.

Estimates of Wolbachia density in the eggs, testes and whole flies of drosophilid hosts have been unable to predict the lack of cytoplasmic incompatibility (CI) expression in so-called mod(-) variants. Consequently, the working hypothesis has been that CI expression, although related to Wolbachia density, is also governed by unknown factors that are influenced by both host and bacterial genomes. Here, we compare the behaviour of the mod(-) over-replicating Wolbachia popcorn strain in its native Drosophila melanogaster host to the same strain transinfected into a novel host, namely Drosophila simulans. We report that (i) the popcorn strain is a close relative of other D. melanogaster infections, (ii) the mod(-) status of popcorn in D. melanogaster appears to result from its inability to colonize sperm bundles, (iii) popcorn is present in the bundles in D. simulans and induces strong CI expression, which demonstrates that the bacterial strain does not lack the genetic machinery for inducing CI and that there is host-species-specific control over Wolbachia tissue tropism, and (iv) infection of sperm bundles by the mod(-) D. simulans wCof strain indicates that there are several independent routes by which a strain can be a CI non-expressor.     

DNA variability and divergence at the notch locus in Drosophila melanogaster and D. simulans: a case of accelerated synonymous site divergence

DNA diversity in two segments of the Notch locus was surveyed in four populations of Drosophila melanogaster and two of D. simulans. In both species we observed evidence of non-steady-state evolution. In D. simulans we observed a significant excess of intermediate frequency variants in a non-African population. In D. melanogaster we observed a disparity between levels of sequence polymorphism and divergence between one of the Notch regions sequenced and other neutral X chromosome loci. The striking feature of the data is the high level of synonymous site divergence at Notch, which is the highest reported to date. To more thoroughly investigate the pattern of synonymous site evolution between these species, we developed a method for calibrating preferred, unpreferred, and equal synonymous substitutions by the effective (potential) number of such changes. In D. simulans, we find that preferred changes per "site" are evolving significantly faster than unpreferred changes at Notch. In contrast we observe a significantly faster per site substitution rate of unpreferred changes in D. melanogaster at this locus. These results suggest that positive selection, and not simply relaxation of constraint on codon bias, has contributed to the higher levels of unpreferred divergence along the D. melanogaster lineage at Notch.     

Chromosome Localization of the λ20p1.4 clone of the Drosophila melanogaster Nuclear Lamina DNA in the melanogaster Species Subgroup of the Genus Drosophila (Sophophora)

Chromosome localization of sequences homologous to 20p1.4 of the Drosophila melanogaster nuclear lamina DNA (nlDNA) was established by in situ hybridization in species of the melanogastersubgroup. DNA of the 20p1.4 clone was shown to be located in the chromocenter in all the species examined. Laboratory strains of D. simulans, D. mauritiana, andD. sechellia exhibited interspecific differences in localization of 20p1.4 nlDNA on chromosome arms. In eight natural populations, intraspecific polymorphism of 20p1.4 nlDNA chromosome localization was shown to be present in D. simulans but absent in D. melanogaster. The possible participation of transposable elements in 20p1.4 nlDNA relocation is discussed.