Evolution of the Drosophila melanogaster-sigma virus system in a natural population from Tübingen

Summary In natural populations of D. melanogaster, usually, a minority of individuals are infected by a Rhabdovirus called sigma. This virus is not contagious but is vertically transmitted through the gametes. In D. melanogaster, a polymorphism for two alleles (O, permissive and P, restrictive) of a gene responsible for resistance to the virus is regularly observed in the wild. On the virus side two types are found, which differ in their sensitivity to the P allele: Type I is very sensitive, and Type II more resistant. Previous findings had led to the hypothesis that an invasion of Type II clones, starting from central France, might be spreading over European populations. This replacement of viral Type I by viral Type II in natural populations could be observed in Languedoc (southern France), where it led to a dramatic increase in the frequency of infected flies. The invasion hypothesis is confirmed by the data from samples collected at Tübingen, where the frequency of Type II clones increased from 0.27 to 0.93 over a 6-year period (1985–1991). However, over the same period, no increase in the frequency of infected flies was observed. The evolution of other viral characteristics is discussed.     

Evolution of natural populations in the Drosophila melanogaster sigma system II. Northern and Central France

A survey of French natural populations of Drosophila melanogaster has been systematically performed, concerning their status of infection by the sigma virus and the characteristics of viral clones. These investigations, which were not as extensive as those performed in the Languedoc region (Fleuriet et al., 1990) nevertheless give a good representation of the evolution of this system because of the long period involved (almost 20 years). Some trends were observed in all French populations such as (1) a decrease in the high efficiency of transmission by males (which is an important parameter for the viral invading ability); (2) high frequency of a best adapted viral Type. These high frequencies might be due to a recent invasion which is expected to spread to other European populations. However, the frequency of infected flies remained low in northern and central France, unlike in Languedoc. The complexity of this, apparently simple, system of two well-known coevolving organisms should once again be stressed. It is impossible with the known parameters to arrive at a general interpretation of observations made in Languedoc and the rest of France. These data may also throw some light on the structure of French wild populations of D. melanogaster which appear to be subdivided into local populations between which gene flow might be low.     

Evolution of the proportions of two sigma viral types in experimental populations of Drosophila melanogaster in the absence of the allele that is restrictive of viral multiplication

A minority of flies in natural populations of Drosophila melanogaster are endemically infected by a rhabdovirus, sigma. The virus is vertically transmitted through male and female gametes. Two alleles of a fly locus, the ref(2)P locus, are present as a polymorphism in all populations: O permissive, and P restrictive for viral multiplication and transmission. Two viral types are known, Type I, which is very sensitive to the P allele, and Type II, which is more resistant. Previous observations have shown that, in presence of the P allele, viral Type II is selected for, in both natural and experimental populations. The aim of the present study was to determine whether, in the absence of P, Type I is selected for, or whether the two types are equivalent. For this purpose, experimental populations deprived of the P allele and differing in the initial proportions of the two viral types were established. After several generations, and despite a possible bias toward Type I, the frequencies of Type I and Type II clones differed in the various populations, depending on their initial values. These findings do not rule out selective advantage of viral Type I in the absence of P, but suggest that, if any, this advantage is in no way comparable to that displayed by viral Type II in the presence of P.     

Comparative evolution of P-M system and infection by the sigma virus in French and Spanish populations of Drosophila melanogaster.

In 1983, an extensive survey of populations of D. melanogaster was started in a southern French region (Languedoc) in two non-Mendelian systems: the P-M system of transposable elements and the hereditary Rhabdovirus sigma. Unexpectedly fast-evolving phenomena were observed and interesting correlations were noted, giving similar geographical pattern to the region in both systems. For these reasons, the analysis was continued and extended towards the north (Rh?ne Valley) and the south (Spain). In the P-M system, all the Languedoc populations evolved from 1983 to 1991 towards the Q type which is characteristic of the Rh?ne Valley populations. In contrast, M' strains are currently observed in the southernmost French populations and in all Spanish ones, so that there is a clear pattern in their geographical distribution. The frequency of flies infected by the sigma virus dramatically increased from 1983 to 1988 in Languedoc; this increase was clearly correlated with some viral characteristics. But, in northern France, similar characteristics did not trigger any increase in the frequency of infected flies. The data presented here show that the distinctive features of Languedoc extend northwards through the Rh?ne Valley up to Lyon and disappears southwards before the Spanish border.     

Perpetuation of the hereditary sigma virus in populations of its host,Drosophila melanogaster. Geographical analysis of correlated polymorphisms

In natural populations of Drosophila melanogaster, about 10% of the individuals are infected by a virus, sigma, which is not contagious but is transmitted through gametes. These populations are also regularly polymorphic for two alleles, O and P, of a locus ref(2)P; the P allele interferes with the multiplication of the virus. Two viral Types are found in populations, differing in their sensitivity to the P allele. Many samples of flies have been collected in different parts of the world and for each of them, the P frequency has been measured and the viral Type determined. A clear geographical differentiation appears for both these traits; they present a mutual adaptation leading to relatively low frequencies of infected flies in natural populations. Most viruses are only known from highly selected laboratory strains. The observations reported in this paper give evidence of the self restraint exercised by the sigma virus at the population level; they indicate that the characteristics of wild viral clones are likely to differ from those of laboratory strains and also from one population to another.The sigma virus is comparable to other genetical elements, that can be more efficiently transmitted than a mendelian allele, such as transposable elements. The discussion illustrates some of the factors involved in the perpetuation of such elements in a population and points out the difficulty of taking them all into consideration in theoretical models dealing with their perpetuation.     

Flies on the move: an inherited virus mirrors Drosophila melanogaster's elusive ecology and demography

Vertically transmitted parasites rely on their host's reproduction for their transmission, leading to the evolutionary histories of both parties being intimately entwined. Parasites can thus serve as a population genetic magnifying glass for their host's demographic history. Here, we study the fruitfly Drosophila melanogaster's vertically transmitted sigma virus DMelSV. The virus has a high mutation rate and low effective population size, allowing us to reconstruct at a fine scale how the combined forces of the movement of flies and selection on the virus have shaped its migration patterns. We found that the virus is likely to have spread to Europe from Africa, mirroring the colonization route of Drosophila. The North American DMelSV population appears to be the result of a recent single immigration from Europe, invading together with its host in the late 19th century. Across Europe, DMelSV migration rates are low and populations are highly genetically structured, likely reflecting limited fly movement. Despite being intolerant of extreme cold, viral diversity suggests that fly populations can persist in harsh continental climates and that recolonization from the warmer south plays a minor role. In conclusion, studying DMelSV can provide insights into the poorly understood ecology of D. melanogaster, one of the best‐studied organisms in biology.     

On the maintenance of the polymorphism at the ref(2)P locus in populations of Drosophila melanogaster

Polymorphism for two alleles of the ref(2)P locus is a very constant feature of French natural populations of Drosophila melanogaster. One of these alleles interferes with the multiplication of the hereditary sigma virus in the fly. An equilibrium, quite similar to the natural one, has been observed previously in experimental populations, whether the sigma virus is present or not. Evidence is given that one of the selection components involved in the maintenance of this equilibrium affects adult stages when flies have not suffered severe larval competition. In conditions of severe larval competition, a maternal effect seems to be involved in the differential egg-to-adult viability of heterozygotes.     

Drosophila C virus and Drosophila hosts: a good association in various environments

To discover whether the beneficial effects observed when Drosophila C virus (DCV_C) infects Drosophila larvae should be considered as a specific case or, as a general rule, the DCV_C-Drosophila association was studied in different environments (3 host-populations, 2 temperatures, 2 viral dose). We measured pre-adult mortality, developmental time and number of ovarioles per fly. In all experiments, and, in all contaminated populations, DCV_C could be considered, either a parasite since it increased pre-adult mortality, or a symbiont since it decreased developmental time and increased mean number of ovarioles. The viral dose necessary for the appearance of beneficial effects varied with population, temperature and life-cycle parameters. A sufficient number of viral particles appears necessary at a specific point in the host life-cycle for a beneficial DCV_C effect on the host. DCV_C does not affect the variance of any of these parameters. The observed changes in DCV_C-contaminated populations were thus not a response to the selection pressure of DCV_C. Two arguments support the association as an example of the “cohabitation-without-struggle model” and will be discussed.     

Viviparity in the halfbeak genera Dermogenys and Nomorhamphus (Teleostei: Hemiramphidae)

Gravid ovaries were examined histologically from two species of Nomorhamphus and 21 populations of Dermogenys. In addition, changes in dry-weight throughout gestation are provided for 15 populations. The ovaries are paired organs running along the lateral body wall and are separated along most of their length. In all specimens examined, embryos are fertilized within the ovarian follicle. Viviparity in these species is divided herein into five categories designated types I–V. In types I and II the entire gestation period is intrafollicular, whereas in types III–V only the early stages of gestation are intrafollicular with the major period of development occurring in the ovarian lumen (intraluminal). Type I is characterized by the retention of a large amount of yolk throughout gestation. Superfetation is not observed. Populations of D. pusilla from Vietnam and Thailand decrease in dry-weight throughout gestation. This, coupled with the slight vascularization of the yolk sac, suggests strict lecithotrophy. Populations of D. pusilla from Singapore and Bangladesh undergo an increase in dry weight and exhibit an increased vascularization of the yolk sac, suggesting a form of unspecialized matrotrophy. Type II is characterized by a small amount of yolk, an expansion of the coelomic cavity and pericardial sac, and a simple cuboidal epithelium on the general body surfaces. Superfetation occurs with up to three broods present within a single ovary. Dermogenys pusilla from Sabah, D. orientalis and Dermogenys sp. (Sulawesi) exhibit the type II form of viviparity. Dermogenys vivipara from the eastern Philippine islands of Culion and Busuanga exhibit characteristics considered intermediate between type I and II. These results are compared with those from other viviparous species exhibiting intrafollicular gestation. In species with types III–V (intraluminal gestation), developing oocytes are restricted to a distinct ridge of ovigerous tissue extending along the entire length of the ovary. Two species, D. viviparus (Luzon, Philippines) and Dermogenys sp. (Luzon) have the type III form of viviparity. In this form, oocytes are small (0.8–1.0 mm) with little yolk reserves and embryos, covered with a simple cuboidal epithelium and possessing an expanded belly sac, are retained within the follicles until a late fin-bud stage. Type III embryos found within the ovarian lumen have a greatly expanded belly sac and remain covered by a simple cuboidal epithelium until parturition. Superfetation is present in these species with two broods observed simultaneously within a single ovary. Five species, D. megarrhamphus, D. weberi, D. viviparus (Jolo, Philippines), Nomorhamphus sp. (Sulawesi), and N. towoetii, were observed with the type IV form of viviparity. Embryos in this category are evacuated into the ovarian lumen prior to a fin-bud stage and retain a large yolk mass throughout development. Superfetation is absent in these species. A differentform of viviparity (type V) is present in D. ebrardtii in which embryos appear to obtain nutrients through a form of oophagy and aldelphophagy (feeding on developing oocytes or less-developed siblings). In all specimens with intraluminal development, atretic oocytes within the ovigerous ridge are abundant. These findings support the hypothesis that current species and generic limits may be artificial and underscores the potential of histological evidence for phylogenetic analysis of this group. J. Morphol. 234:295–317, 1997. © 1997 Wiley-Liss, Inc.     

Expression of a cystine-rich fish antifreeze in transgenicDrosophila melanogaster

We have usedDrosophila melanogaster as a model system for the transgenic expression of cystine-rich Type II antifreeze protein (AFP) from sea raven. This protein was synthesized and secreted into fly haemolymph where it migrated as a larger species (16 kDa) than the mature form of the protein (14 kDa) as judged by immunoblotting.Drosophila-produced Type II AFP demonstrated antifreeze activity both in terms of thermal hysteresis (0.13 °C) and inhibition of ice recrystallization. Recombinant AFP was purified and N-terminal sequencing revealed a 17 aa extension that began at the predicted signal peptide cleavage point. The expression of all three AFP types in transgenicDrosophila has now been achieved. We conclude that the globular Type II and Type III AFPs are better choices for antifreeze transfer to other organisms than is the more widely used linear Type I AFP.     

Constraints to genetic exchange support gene coadaptation in a tripartite RNA virus

Genetic exchange by recombination, or reassortment of genomic segments, has been shown to be an important process in RNA virus evolution, resulting often in important phenotypic changes affecting host range and virulence. However, data from numerous systems indicate that reassortant or recombinant genotypes could be selected against in virus populations and suggest that there is coadaptation among viral genes. Little is known about the factors affecting the frequency of reassortants and recombinants along the virus life cycle. We have explored this issue by estimating the frequency of reassortant and recombinant genotypes in experimental populations of Cucumber mosaic virus derived from mixed infections with four different pairs of isolates that differed in about 12% of their nucleotide sequence. Genetic composition of progeny populations were analyzed at various steps of the virus life cycle during host colonization: infection of leaf cells, cell-to-cell movement within the inoculated leaf, encapsidation of progeny genomes, and systemic movement to upper noninoculated leaves. Results indicated that reassortant frequencies do not correspond to random expectations and that selection operates against reassortant genotypes. The intensity of selection, estimated through the use of log-linear models, increased as host colonization progressed. No recombinant was detected in any progeny. Hence, results showed the existence of constraints to genetic exchange linked to various steps of the virus life cycle, so that genotypes with heterologous gene combinations were less fit and disappeared from the population. These results contribute to explain the low frequency of recombinants and reassortants in natural populations of many viruses, in spite of high rates of genetic exchange. More generally, the present work supports the hypothesis of coadaptation of gene complexes within the viral genomes.     

Genetics of organophosphate resistance in field populations of the house fly (Diptera: Muscidae)

The genetics of resistance to the organophosphate insecticide diazinon were investigated in four populations of the house fly, Musca domestica L., collected in the southern United States. Crosses were made between individual females of lines derived from each population and males of a susceptible strain with three recessive mutants on chromosome II. Individual F1 females were crossed to mutant males, and the progenies were scored for resistance to diazinon and for the presence of mutant phenotypes. A major chromosome II gene for resistance to diazinon was present in all populations at an overall frequency of 83%. Map distances between the resistance gene and the mutant aristapedia and between the mutants aristapedia and stubby wing were highly variable in all populations. Recombination among the visible mutants was usually reduced in resistant progenies relative to susceptible progenies. The data suggest that a single major gene for resistance to diazinon was present on chromosome II in all test populations at variable map positions and is usually associated with a chromosome rearrangement, probably an inversion. The results are similar to those obtained earlier with house fly populations selected for resistance to insecticides in the laboratory; therefore, they seem to be characteristic of field and laboratory populations of the house fly. Overall, the data offer an explanation for previous results suggesting the existence of multiple, closely linked genes for metabolic resistance to insecticides on house fly chromosome II.     

Interaction between a picornavirus and a wild population ofDrosophila melanogaster

Summary Drosophila C virus (DCV) has a considerable impact on ovarian morphogenesis inDrosophila melanogaster host populations. This virus also affects the developmental time and the fresh weight of infected females. In order to investigate the hypothesis that DCV may play a role in the dynamics ofDrosophila populations, the fertility and embryonic and larvo-pupal death rates of a host population and that of five DCV-free populations were determined. A comparison of two populations, one of them DCV-free, the other infected, suggested that the fertility of the DCV-infected flies was higher than that of uninfected flies, despite a greater larvo-pupal death rate. Fertility of the infected flies was greater among the infected population than for the DCV-free populations. The DCV-free populations originated from five different localities. The virus clearly does have an impact on the biotic potential of its host population. This paper reports for the first time a positive interaction between a viral population and a host population as it increases certain parameters of host population dynamics.     

Isolation and characterization of recombinant DNA clones of avian retroviruses: size heterogeneity and instability of the direct repeat

Unintegrated proviral DNA of Schmidt-Ruppin B Rous sarcoma virus was cloned in the bacteriophage lambda vector Charon 21A. A total of 12 independent recombinant lambda SRBtd clones which were derived from the transformation-defective component in the viral preparation were analyzed with restriction endonucleases and molecular hybridization techniques. Three classes of clones were observed. Type I clones contained a 5.0-megadalton insert of viral DNA, type II clones contained phage with two size classes of inserts (5.0 and 5.2 megadaltons), and one type III clone contained only a 5.2-megadalton insert. The smaller insert present in type II clones appeared to be derived by deletion of one copy of a directly repeated sequence which was present in the larger insert. Mapping data indicated that the deletion includes all or part of the terminal repeat found in linear double-stranded proviral DNA. Similar results were obtained from lambda RAV2 recombinant clones derived from Rous-associated virus type 2. Analysis of DNA from type II and type III clones of lambda SRBtd and lambda RAV2 revealed limited heterogeneity in the size of the direct repeat.     

Tracing the history and ecological context of Wolbachia double infection in a specialist host (Urophora cardui)—parasitoid (Eurytoma serratulae) system

The endosymbiotic bacterium Wolbachia is the most widespread bacteria in insects, yet the ecology of novel acquisitions in natural host populations is poorly understood. Using temporal data separated by 12 years, I tested the hypothesis that immigration of a parasitoid wasp led to transmission of its Wolbachia strain to its dipteran host, resulting in double‐strain infection, and I used geographic and community surveys to explore the history of transmission in fly and parasitoid. Double infection in the fly host was present before immigration of the parasitoid. Equal prevalence of double infection in males and females, constant prevalence before and after immigration in two regions, and increase in one region of immigration indicate little if no competition between strains. Double infection was present throughout the fly's distribution range, but proportions varied highly (0–0.71, mean = 0.26). Two fly‐specific MLST strains, observed in Eastern and Western Europe, respectively, differed at hcpA only. Flies with either fly‐strain could be double infected with the parasitoid's strain. The geographic distribution of double infection implies that it is older than the fly host's extent distribution range and that different proportions of double infection are caused by demographic fluctuations in the fly. The geographic data in combination with community surveys of infections and strains further suggest that the parasitoid strain was the fly's ancestral strain that was transmitted to the parasitoid, that is, the reverse transmission route as first hypothesized. Based on these findings together with a comparison of oviposition strategies of other hosts harboring related Wolbachia strains, I hypothesize that trans‐infection during an insect host's puparial metamorphosis might be important in promoting horizontal transmission among diverse holometabolic taxa.     

Recrudescent Infection Supports Hendra Virus Persistence in Australian Flying-Fox Populations

Zoonoses from wildlife threaten global public health. Hendra virus is one of several zoonotic viral diseases that have recently emerged from Pteropus species fruit-bats (flying-foxes). Most hypotheses regarding persistence of Hendra virus within flying-fox populations emphasize horizontal transmission within local populations (colonies) via urine and other secretions, and transmission among colonies via migration. As an alternative hypothesis, we explore the role of recrudescence in persistence of Hendra virus in flying-fox populations via computer simulation using a model that integrates published information on the ecology of flying-foxes, and the ecology and epidemiology of Hendra virus. Simulated infection patterns agree with infection patterns observed in the field and suggest that Hendra virus could be maintained in an isolated flying-fox population indefinitely via periodic recrudescence in a manner indistinguishable from maintenance via periodic immigration of infected individuals. Further, post-recrudescence pulses of infectious flying-foxes provide a plausible basis for the observed seasonal clustering of equine cases. Correct understanding of the infection dynamics of Hendra virus in flying-foxes is fundamental to effectively managing risk of infection in horses and humans. Given the lack of clear empirical evidence on how the virus is maintained within populations, the role of recrudescence merits increased attention.     

杂交水稻中超显性效应的分析

本文提出亚群体分析法,在二个衍生于杂交稻推广组合的F2群体中,挑选均匀分布于连锁图谱的DNA标记作为固定因子,分别根据每个固定因子的基因型将F2群体分成三类亚群体：母本型（I型）、父本型（II型）和杂合型（III型）。在大量III型亚群体中,杂合度与产量和穗数呈显著正相关。在表现这种相关性的III型亚群体内分析产量QTL和穗数QTL,发现超显性作用是研究组合杂种优势的主要遗传基础。 Abstract:Two F2 populations were each derived from a commercial hybrid and used for identification of genetic factors contributing to heterosis of rice.DNA markers distributing evenly in the linkage maps were selected as fixing factors.The F2 populations were divided into three types of sub-populations based on the genotypes of each fixing factor:maternal type (Type I),paternal type (Type II) and heterozygous type (Type III).In a large number of Type III sub-populations,significant correlations were observed between heterozygosity and grain yield,and between heterozygosity and number of panicle.QTL analysis in Type III sub-populations showing such correlations indicated that over-dominance was the major genetic basis of heterosis in the two crosses.     

Viral infectious and chromosome aberrations in Bank Vole from natural and laboratory populations]

The frequency of chromosome damage was studied in the carriers of virus of the hemorrhagic fever with renal syndrome (Puumala virus) and in noninfected animals from two laboratory colonies and two natural populations of bank vole. In the laboratory colony, where Puumala virus persisted for three years, multiaberrant ("rogue") cells were found in the bone marrow; the mean frequencies of both structural and numeral chromosome abnormalities were significantly enhanced. In the other laboratory colony, no Puumala virus was detected during all 30 years of its existence, but the mean frequencies of structural chromosome damage were increased to the same degree probably due to the prolonged breeding under laboratory conditions, which resulted in suppression of immunity and DNA repair. The voles from the natural populations were more resistant to the clastogenic viral effect, but they also had multiaberrant cells which served as indicators of viral infection. The data obtained support the hypothesis that viral infections increase mutation rate, contributing thereby to the evolution process.     

Virus population extinction via ecological traps

Populations are at risk of extinction when unsuitable or when sink habitat exceeds a threshold frequency in the environment. Sinks that present cues associated with high-quality habitats, termed ecological traps, have especially detrimental effects on net population growth at metapopulation scales. Ecological traps for viruses arise naturally, or can be engineered, via the expression of viral-binding sites on cells that preclude viral reproduction. We present a model for virus population growth in a heterogeneous host community, parameterized with data from populations of the RNA bacteriophage Φ6 presented with mixtures of suitable host bacteria and either neutral or trap cells. We demonstrate that viruses can sustain high rates of population growth in the presence of neutral non-hosts as long as some host cells are present, whereas trap cells dramatically reduce viral fitness. In addition, we demonstrate that the efficacy of traps for viral elimination is frequency dependent in spatially structured environments such that population viability is a nonlinear function of habitat loss in dispersal-limited virus populations. We conclude that the ecological concepts applied to species conservation in altered landscapes can also contribute to the development of trap cell therapies for infectious human viruses.