Mitochondrial DNA variation and genetic structure in Old-World populations of Drosophila subobscura

To discover the relation between mitochondrial DNA (mtDNA) polymorphism and the geographic population structure of Drosophila subobscura previously established for other genetic traits, a wide Paleartic survey was carried out. A total of 24 nucleomorphs was observed among 261 isofemale lines assayed by 11 restriction endonucleases with 38 different sites in the mtDNA cleavage map. The differentiation of the Canary Islands populations (delta = 0.0119) compared with the mean among all the other continental and insular populations (delta = 0.0002) is striking. Both the great divergence among Canary Islands nucleomorphs (delta = 0.021) compared with the maximum nucleomorph distance in all other populations (delta = 0.017) and the abundance of endemic nucleomorphs (11) on the Canary Islands (50% of the total number of different nucleomorphs found in the entire distribution area) suggest that this molecular differentiation most probably results from the very old age of the Canary Islands populations rather than from drift and founder effects.     

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.     

New gene arrangements in natural populations of Drosophila subobscura

Four new gene arrangements were detected in natural populations of Drosophila subobscura near Zürich, Switzerland. Three of them are the result of inversions not recorded before, and one is a new combination of known gene arrangements. Another arrangement, I₃₊₄, was recorded for the first time in Switzerland. It may have been imported or possibly may have arisen de novo. It is suggested that inversion mutations are not so infrequent as so far supposed, if the presence of mobile elements is taken into account in D. subobscura.     

Studies of chromosome structure polymorphism in Asia Minor and Persian populations of Drosophila subobscura Coll

Summary By means of the male sample method samples of D. subobscura from the north and south coast of Asia Minor and from the south coast of the Caspian Sea were examined with regard to the gene arrangements contained in them. The samples were judged by the following three criteria: (1) the number of different gene arrangements present, (2) the frequencies in which these gene arrangements occur, and (3) the rate of zygotic combinations.The samples from Asia Minor were found to be highly polymorphic. In this respect, they are comparable with other samples from the northern border of the Mediterranean region.The Persian populations are poor as regards number of gene arrangements. Commonly, some gene arrangements are highly frequent and, accordingly, high rates of homozygous combinations dominate. Presumably, these are marginal populations.Comparisons between samples from the north and south coast of Asia Minor confirm the findings from other parts of the species' range. As a rule, the frequencies of various gene arrangements vary according to the latitude in which they occur. Since climate and vegetation are also largely correlated with latitude, it is assumed that this is due to adaptive reaction to ecological factors.Comparing populations from the Western and Eastern Mediterranean region, similar differences were recorded. Some gene arrangements seem to be confined to either the western or eastern part of the area, and other gene arrangements differ in frequency. Since the ecological conditions in these areas appear to be comparable, such differences might be due to the history of the formation and dispersal of gene arrangements.In general, the frequencies of the gene arrangements in the chromosomes A, U, E and O are most equal along the northern margin of the Mediterranean area and, correspondingly, the frequencies of the heterokaryotypes are highest. It may be assumed that many heterokaryotypes reach their highest degree of heterotic supremacy in this area, and/or that the different homokaryotypes are uniform with respect to adaptive values. The reasoning rests on the additional assumption that in this area the biological conditions are the least extreme, or, in other words, that D. subobscura is best adapted to this environment. As it is, the region appears to lie geographically near the center of the species' range. In the I-chromosome the most equal frequencies of gene arrangements along with the highest frequency of the heterokaryotypes are to be found in Central Europe.On the other hand, in various populations lying supposedly at the borders of the species' range, the frequencies of gene arrangements of at least some chromosomes are strongly unequal and, accordingly, the frequencies of the homokaryotypes are high.In populations where the polymorphism is high, the character of the polymorphism varies slightly from chromosome to chromosome. In the O-chromosome usually one gene arrangement is abundant, while a large number of others are rare; two gene arrangements tend to dominate in the I- and U-chromosome, whereas in the E-chromosome several gene arrangements are apt to share similar frequencies.     

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.     

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.     

Chromosomal inversion polymorphism leads to extensive genetic structure: a multilocus survey in Drosophila subobscura

The adaptive character of inversion polymorphism in Drosophila subobscura is well established. The O(ST) and O(3+4) chromosomal arrangements of this species differ by two overlapping inversions that arose independently on O(3) chromosomes. Nucleotide variation in eight gene regions distributed along inversion O(3) was analyzed in 14 O(ST) and 14 O(3+4) lines. Levels of variation within arrangements were quite similar along the inversion. In addition, we detected (i) extensive genetic differentiation between arrangements in all regions, regardless of their distance to the inversion breakpoints; (ii) strong association between nucleotide variants and chromosomal arrangements; and (iii) high levels of linkage disequilibrium in intralocus and also in interlocus comparisons, extending over distances as great as approximately 4 Mb. These results are not consistent with the higher genetic exchange between chromosomal arrangements expected in the central part of an inversion from double-crossover events. Hence, double crossovers were not produced or, alternatively, recombinant chromosomes were eliminated by natural selection to maintain coadapted gene complexes. If the strong genetic differentiation detected along O(3) extends to other inversions, nucleotide variation would be highly structured not only in D. subobscura, but also in the genome of other species with a rich chromosomal polymorphism.     

Adaptation of Drosophila subobscura chromosomal inversions to climatic variables: the Balkan natural population of Avala

Genetica - The adaptive value of chromosomal inversions continues raising relevant questions in evolutionary biology. In many species of the Drosophila genus, different inversions have been...     

Temporal changes of chromosomal polymorphism in natural populations of Drosophila subobscura

The behaviour of the chromosomal polymorphism of D. subobscura was analyzed in relation to temporal changes, daily, seasonal and annual. Firstly, chromosome analyses were carried out over a year in a natural population. Samples were taken at monthly intervals in Tibidabo, a locality close to Barcelona. In all the months except January, the number of individuals captured was enough to carry out a chromosome analysis of that population. The A, E and O chromosomes showed a great uniformity in the frequencies of gene arrangement over the year. However, significant changes occur in the frequencies of J and U chromosomes. The J₁ and U₁₊₂₊₈ arrangements showed a similar tendency, with two maxima, in June and February, and a minimum in September. These changes seem to be unrelated to the cyclical climatic changes. A chromosome analysis of the subsamples captured during the day, in the May sample, was done. In no case was the change in chromosome frequencies statistically significant. The behaviour of the U_st arrangement must be noted, the frequency of which decreased through the day. Finally, the data for the Tibidabo population were compared with samples captured in nearby localities over a period of 21 years. Significant differences were found in the frequencies of all the chromosomes, with the exception of J. The most differentiated population was the last sampled, from 1982. In this population the st arrangements tended to decrease in the A, E and O chromosomes and A₂, E_1+2+9+12 and O₃₊₄₊₇ increase, respectively.     

Temperature-related genetic changes in laboratory populations of Drosophila subobscura: evidence against simple climatic-based explanations for latitudinal clines

Parallel latitudinal clines to the long-standing ones in the original Palearctic populations have independently evolved at different rates for chromosomal polymorphism and body size in South and North American populations of Drosophila subobscura since colonization around 25 years ago. This strongly suggests that (micro) evolutionary changes are largely predictable, but the underlying mechanisms are unknown. The putative role of temperature per se was investigated by using three sets of populations at each of three temperatures (13 degrees , 18 degrees , and 22 degrees C) spanning much of the tolerable range for this species. We found a lower chromosomal diversity at the warmest temperature; a quick and consistent shift in gene arrangement frequencies in response to temperature; an evolutionary decrease in wing size, mediated by both cell area and cell number, at 18 degrees C; no relationship between wing size and those inversions involved in latitudinal clines; and a shortening of the basal length of longitudinal vein IV relative to its total length with increasing standard dose. The trends for chromosomal polymorphism and body size were generally inconsistent from simple climatic-based explanations of worldwide latitudinal patterns. The findings are discussed in the light of available information on D. subobscura and results from earlier thermal selection experiments with various Drosophila species.     

The genetic structure of Drosophila ananassae populations from Asia, Australia and Samoa

Information about genetic structure and historical demography of natural populations is central to understanding how natural selection changes genomes. Drosophila ananassae is a widespread species occurring in geographically isolated or partially isolated populations and provides a unique opportunity to investigate population structure and molecular variation. We assayed microsatellite repeat-length variation among 13 populations of D. ananassae to assess the level of structure among the populations and to make inferences about their ancestry and historic biogeography. High levels of genetic structure are apparent among all populations, particularly in Australasia and the South Pacific, and patterns are consistent with the hypothesis that the ancestral populations are from Southeast Asia. Analysis of population structure and use of F-statistics and Bayesian analysis suggest that the range expansion of the species into the Pacific is complex, with multiple colonization events evident in some populations represented by lineages that show no evidence of recent admixture. The demographic patterns show isolation by distance among populations and population expansion within all populations. A morphologically distinct sister species, D. pallidosa, collected in Malololelei, Samoa, appears to be more closely related to some of the D. ananassae populations than many of the D. ananassae populations are to one another. The patterns of genotypic diversity suggest that many of the individuals that we sampled may be morphologically indistinguishable nascent species.     

The genetics of Drosophila subobscura populations. 3. Inversion polymorphism and climatic factors

Summary The natural population of Drosophila subobscura of Mt. Parnes has been followed for two successive years and the frequency of the gene arrangements determined. Some changes occured in the first year but do not explain the existence of the geographical clines for these frequencies. Within the second year no changes were observed. For chromosomes J and U there is evidence for a greater fitness of the heterozygotes.Experiments on temperature effect on viability of adult male flies, on their temporal sterility, on the viability of flies during all their biological cycle and experiments on the effect of dryness on the viability of adult male flies did not show an important change of gene arrangement frequencies in the resistant individuals. Nor did diapausing females have very different gene arrangements, except for chromosome O. Cage experiments with temperature as a selective agent did not elucidate the action of this factor. It is concluded that temperature or dryness do not affect very much the polymorphic system, nor explain the existence of these clines.This polymorphic system is much more stable in D. subobscura than in D. pseudoobscura, its nearctic relative. It is also richer. It is concluded that it is historically older and helps the populations to cope with many environmental changes.     

Mitochondrial DNA variation and genetic structure in populations of Drosophila melanogaster

The understanding of the genetic structure of a species can be improved by considering together data from different types of genetic markers. In the past, a number of worldwide populations of Drosophila melanogaster have been extensively studied for several such markers, including allozymes, chromosomal inversions, and quantitative characters. Here we present results from a study of restriction- fragment-length polymorphisms of mitochondrial DNA (mtDNA) in 92 isofemale lines from many of the same geographic populations of D. melanogaster. Eleven restriction enzymes were used, of which four revealed restriction-site polymorphism. A total of 24 different haplotypes were observed, of which 18 were unique to single populations. In many populations, the unique haplotypes have reached high frequency without being observed in neighboring populations. A Wagner parsimony tree reveals that mutationally close variants show geographical clumping, suggesting local differentiation of mtDNA in populations. The Old-World and the New-World populations are differentiated, with the predominant Old-World haplotype being virtually absent from the New World. These results contrast with those for the nuclear genes, in which many loci show parallel clines in different continents, and suggest a common origin of D. melanogaster populations in North America.