Lack of seasonal changes in mitochondrial DNA variability of a Drosophila subobscura population

Restriction site analysis of mtDNA of 550 isofemale lines corresponding to different seasonal samples of a single geographic population of Drosophila subobscura was carried out. The distribution pattern of haplotypes was similar to that observed for the entire range of the species on the European continent: two haplotypes were equally and highly frequent, and a set of sporadic haplotypes were almost never present in more than one seasonal sampling. No statistically significant evidence was found for between-population heterogeneity across time, and the mean within-population variation was similar to other mtDNA restriction site analyses previously reported for D. subobscura populations. These observations could be interpreted in terms of the neutral mutation hypothesis where the entire population has not yet reached an equilibrium. The causes of this non-equilibrium are most likely periodic winter bottlenecks, phenomena that particularly affect the distribution of mtDNA haplotypes across time.     

Evolutionary domestication in Drosophila subobscura

The domestication of plants and animals is historically one of the most important topics in evolutionary biology. The evolutionary genetic changes arising from human cultivation are complex because of the effects of such varied processes as continuing natural selection, artificial selection, deliberate inbreeding, genetic drift and hybridization of different lineages. Despite the interest of domestication as an evolutionary process, few studies of multicellular sexual species have approached this topic using well-replicated experiments. Here we present a comprehensive study in which replicated evolutionary trajectories from several Drosophila subobscura populations provide a detailed view of the evolutionary dynamics of domestication in an outbreeding animal species. Our results show a clear evolutionary response in fecundity traits, but no clear pattern for adult starvation resistance and juvenile traits such as development time and viability. These results supply new perspectives on the confounding of adaptation with other evolutionary mechanisms in the process of domestication.     

Stress response in Drosophila subobscura

The pattern of puffing and protein synthesis was determined in individuals of Drosophila subobscura subjected to heat shock. Depending on the extent of the heat treatment, the response at the puffing level varied. Some puffs were expressed at 31°–34°C, and others at 37° C. Considering the response as a whole the depression of gene activity after shock at 31°–34° C in individuals raised at 19° C was greater than with the other treatments. Six major heat shock proteins (Hsps) were found in this species. The properties of the high molecular weight proteins are conserved their electrophoretic characteristics and the range of temperatures over which they are synthesized are close to those in other Drosophila species. Remarkable heterogeneity was found in the small Hsps. In addition, an M_r=41000 Hsp was clearly identified in this species. A low level of variability in the patterns of protein synthesis compared with those of puffing activity was detected.     

The genetics of Drosophila subobscura populations XVIII. Multiple insemination and sperm displacement in Drosophila subobscura

The multiple insemination of Drosophila subobscura females in nature was studied by the analysis of single female offspring using two highly polymorphic enzyme systems. On qualitative grounds it was found that 23% of these females were inseminated at least twice. The power of detection of one enzyme system was estimated using the already known multi-inseminated females detected by the other system. This permitted the true frequency estimation of multi-inseminated females, in nature, which is 0.276 (for 95% confidence limits it ranges from 0.186 to 0.417). In a laboratory experiment it was found that sperm from the second inseminating male displaces the sperm from the first male.     

Microdistribution and habitat selection in Drosophila subobscura Collin

Microdistribution and habitat selection were investigated in a population of Drosophila subobscura near Leeds, Yorkshire. Microdistribution was examined along a transect running through a deciduous woodland. Two main types of habitat were recognized, those with dry soil and a closed canopy of mature trees (dry and dark) and those with wet soil and an open canopy of mature trees (wet and light). Flies were much more frequent, judged by their attendance at baited traps, in the dry/dark habitat. Flies taken from both habitat types, marked with different coloured fluorescent dust and released at a point mid-way between the original habitats, tended to return to their area of initial capture.     

Genetic and environmental factors in the resistance of Drosophila subobscura adults to high temperature shock

Summary We have carried out two equivalent selection experiments to increase and decrease heat shock resistance of Drosophila subobscura adults, using an indirect selection method that avoids excessive consanguinity. Heat shock was 33±0.5 °C at saturation humidity. Control lines showed a rapid change of the physiological trait as a consequence of laboratory culture conditions, expressed as a decrease both in heat shock resistance and in the initial population variability for heat shock resistance. Thus, this reduction of variability seems to consist in the loss of those combinations of genes that confer high resistance to heat shock. After eight generations of selection, the selected lines were differentiated from their respective control lines, and the selection response obtained was similar in resistant and sensitive lines. Differences in survival of progeny of reciprocal crosses between selected lines suggest that inheritance of heat resistance may depend in part on the origin of egg cytoplasm.     

Adaptation to the laboratory environment in Drosophila subobscura

Adaptation to a novel environment is expected to have a number of features. Among these is a temporal increase in fitness and some or all of its components. It is also expected that additive genetic variances for these fitness characters will fall. Finally, it is expected that at least some additive genetic correlations will decrease, from positive toward negative values. In a study of several life‐history variables in a Drosophila subobscura population sampled from the wild and then cultured in the laboratory, we did not find any such longitudinal trends over the first 29 generations. However, a temporal comparison (over 14 generations) of the later generations of this laboratory‐adapted population with a new population, derived from a more recent wild‐caught sample, indicated clearly that laboratory adaptation was nonetheless occurring. This study suggests the need for extensive replication and control in studies of the features of adaptation to a novel environment.     

Mitochondrial DNA effects on fitness in Drosophila subobscura

We tested different fitness components on a series of conspecific mtDNA haplotypes, detected by RFLPs in Drosophila subobscura. Additionally, haplotype VIII, endemic to the Canary Islands, was tested upon its own native nuclear DNA background and upon that of the rest of mtDNAs tested herein. We found that both nuclear and mitochondrial DNA can have a significant effect upon their hosts' fitness, and that negative selection is one of the mechanisms that can intervene in this species' mtDNA haplotype pattern. We discuss the importance of this mechanism in relation to genetic drift, in the form of periodic population bottlenecks, and how the latter can enhance the former. We also detected a significant positive effect of haplotype VIII upon fitness that could explain in part the dominance of this endemic haplotype on some of the Canary Islands, and a mitochondrial heterosis involving this haplotype when on a foreign nuclear DNA background.     

Patterns of puffing activity and chromosomal polymorphism in Drosophila subobscura

Summary The effect of inbreeding on the puffing patterns of polytene chromosomes of Drosophila subobscura was analysed. Puffing activity was studied in two strains of D. subobscura: one which had been subject to inbreeding for 288 generations, and in the hybrids from a cross between them. A strong overall decrease in puffing activity was found in the inbred line. In general, hybrids behaved in a similar way to the inbred line or showed activity intermediate between the two lines. The fertility and viability of the two homozygous lines and of the hybrids were also determined. These parameters of fitness are very low in the inbred line. Hybrids displayed intermediate behaviour.Publication No. 109, Departamento de Genética, Universidad de Valencia, Spain     

The ontogeny and cost of migration in Drosophila subobscura Collin

The ontogeny of migratory potential of male and female Drosophila subobscura has been quantified using tethered flight in the laboratory. In both sexes there is an increase in the few days immediately after eclosion, and a decline before the flies cease reproductive activity. There is an intervening peak in the potential of males, which may result from an increased rejection-rate by inseminated females. The migratory potential generally appears greater than that usually utilized in the field. Such extended migration extracts a significant cost from female flies in terms of reduced fecundity.     

Patterns of puffing activity and chromosomal polymorphism in Drosophila subobscura

The puffing patterns in polytene E chromosomes of Drosophila subobscura were followed in third-instar larvae and throughout the prepupa period. Two gene arrangements, E_st and E_1·2+9–12 were studied. A majority of puffs exhibit a similar pattern, but the puffs 61AC and 67AB behave differently in the two chromosomal arrangements, both in homozygotes and in heterozygotes. These two puffs are located at the end of the E₁₂ inversion. This position effect is an interesting phenomenon that probably is not due to a heterochromatinization effect.     

Open ground individuals and population structure in Drosophila subobscura Collin

The distribution of Drosophila subobscura adults within, and their movement through, open ground habitats has been investigated, along with the status of such individuals within populations as a whole. Movement occurs between open ground and woodland, to some extent as a result of passive transport by air currents, but flies can remain in open ground for at least several days at a time. There are no detectable differences in either the reproductive age-structure or the body-size distributions of flies collected from woodland and open ground habitats, although a significantly higher proportion of females are found in open ground than in woodland. The large number of gravid females in open ground provide the potential for breeding there, but circumstantial evidence suggests that this is unlikely to occur. Rather, their presence appears to reflect the foraging strategy adopted by females in response to their high nutritional requirements. The flux of flies through open ground between woodlands suggests that previous estimates of the genetically-important parameter, effective population size, have been seriously underestimated for natural D. subobscura populations.     

O-chromosome lethal frequencies in Serbian and Montenegrin Drosophila subobscura populations

Lethal chromosomal frequencies were obtained from three Drosophila subobscura samples from the Mt. Avala (Serbia) population in September 2003 (0.218), June 2004 (0.204) and September 2004 (0.250). These values and those from other Balkan populations studied previously (Petnica, Kamariste, Zanjic and Djerdap) were used to analyze the possible effect of population, year, month and altitude above sea level on lethal chromosomal frequencies. According to ANOVAS no effect were observed. Furthermore, the lethal frequencies of the Balkan populations did not vary according to latitude. This is probably due to the relative proximity and high gene flow between these populations. From a joint study of all the Palearctic D. subobscura populations so far analyzed, it can be deduced that the Balkan populations are located in the central area of the species distribution. Finally, it seems that lethal chromosomal frequencies are a consequence of the genetic structure of the populations.     

Microsatellite variation in colonizing and palearctic populations of Drosophila subobscura

The recent colonization of North America by Drosophila subobscura has provided a great opportunity to analyze a colonization process from the beginning. A comparative study using 10 microsatellite loci was conducted for five European and two North American populations. No genetic differentiation between European populations was detected, indicating that gene flow is high among them and that the microsatellites used in the present work represent neutral markers not subject to differentiation due to selection. Extensive reduction in the number of alleles and a significant decrease in heterozygosity in colonizing populations were detected that could be explained by the founder effect and a subsequent quick but not infinite expansion. Assuming that all alleles present in the colonized area were carried by the sample of colonizers, we estimated that most probably 4-11 individuals expanded in the new area. F(ST) and the chord distance measures reflect the colonization process more accurately, since drift has been the major force in differentiating the Old and New World populations, and thus other measures considering allele size differences, such as Rho(ST) and deltamu2, are less reliable for studying nonequilibrium populations. Finally, our results were consistent with the two-phase microsatellite mutational model, indicating that most alleles are generated by gain or loss of a repeat unit, while some alleles originate by more complex mutations.     

Rapid evolution of wing size clines in Drosophila subobscura

Parallel latitudinal clines across species and continents provide dramatic evidence of the efficacy of natural selection, however little is known about the dynamics involved in cline formation. For example, several drosophilids and other ectotherms increase in body and wing size at higher latitudes. Here we compare evolution in an ancestral European and a recently introduced (North America) cline in wing size and shape in Drosophila subobscura. We show that clinal variation in wing size, spanning more than 15 degrees of latitude, has evolved in less than two decades. In females from Europe and North America, the clines are statistically indistinguishable however the cline for North American males is significantly shallower than that for European males. We document that while overall patterns of wing size are similar on two continents, the European cline is obtained largely through changing the proximal portion of the wing, whereas the North American cline is largely in the distal portion. We use data from sites collected in 1986/1988 (Pegueroles et al. 1995) and our 1997 collections to compare synchronic (divergence between contemporary populations that share a common ancestor) and allochronic (changes over time within a population) estimates of the rates of evolution. We find that, for these populations, allochronically estimated evolutionary rates within a single population are over 0.02 haldanes (2800 darwins), a value similar in magnitude to the synchronic estimates from the extremes of the cline. This paper represents an expanded analysis of data partially presented in Huey et al. (2000).     

Sperm dysfunction in sex ratio males of Drosophila subobscura

Drosophila subobscura has 128 spermatids per cyst, enclosed by two cyst cells. At beginning of elongation in control males the spermatid nuclei surround the head cyst cell nucleus, in sex ratio males nuclei are found throughout the cyst. Spermatid nuclei can elongate in any position in the cyst. Nuclei can be eliminated during individualization or degenerate after individualization. The number of sperm in any wrong position in the cyst varies in control males from 0 to about ten, in sex ratio males from 0 to more than 50. Two cyst sizes are distinguishable. At beginning of elongation small cysts have homogeneously stained spherical nuclei which later on are rod like. Large cysts have granulated nuclei which at first become spindle shaped and then slender. The length of the DNA containing part of elongated sperm heads of the long class is about 33 m in sex ratio and control males. The small sperm heads are 15 m in sex ratio but 20 m in control males. The complete DNA-containing-sperm-length is about 10% less in short sperm and 5% less in long sperm of sex ratio males than in those of control. Sex ratio males have more cysts per testis than control males. In sex ratio we counted 53.8%, in control males 49.4% short cysts.The work was supported by the grants N.S.F. GB-43209 and NIH GM 21732 and the Schweizerischer Nationalfonds zur Foerderung der wissenschaftlichen Forschung grant Nr. 3.815.72.