An adaptive chromosomal polymorphism affecting size-related traits,and longevity selection in a natural population ofDrosophila buzzatii

Size-related phenotypic variation among second-chromosome karyotypes inDrosophila buzzatii was examined in an Argentinian natural population. For all measured traits (thorax and wing length; wing, head and face width), this inversion polymorphism exhibited a significant and (additive) linear contribution to the phenotypic variance in newly emerged wild flies. The results suggest that only overall body size, and not body shape, is affected. as no karyotypic variation was found for any trait when the effects of differences in within-karyotype size were removed with Burnaby's method. Likewise, in an experiment of longevity selection in the wild, variation in chromosomal frequencies was verified in the direction predicted on the basis of: (i) previous studies on longevity selection for body size in the wild and (ii) the pattern of chromosomal effects we observed on size. The direction of such selection is consistent with a pattern of antagonistic selection detected in previous studies on the inversion polymorphism.     

INVERSION LENGTH AND BREAKPOINT DISTRIBUTION IN THE DROSOPHILA BUZZATII SPECIES COMPLEX: IS INVERSION LENGTH A SELECTED TRAIT?

Length and position of breakpoints are characteristics of inversions that can be precisely determined on the polytene chromosomes of Drosophila species, and they provide crucial information about the processes that govern the origin and evolution of inversions. Eighty-six paracentric inversions described in the Drosophila buzzatii species complex and 18 inversions induced by introgressive hybridization in D. buzzatii were analyzed. In contrast to previous studies, inversion length and breakpoint distribution have been considered simultaneously. We conclude that: (1) inversion length is a selected trait; rare inversions are predominantly small while evolutionarily successful inversions, polymorphic and fixed, are predominantly intermediate in length; a nearly continuous variation in length, from small to medium sized, is found between less and more successful inversions; (2) there exists a significant negative correlation between length and number of polymorphic inversions per species which explains 39% of the inversion length variance; (3) natural selection on inversion length seems the main factor determining the relative position of breakpoints along the chromosomes; (4) the distribution of breakpoints according to their band location is non-random, with chromosomal segments that accumulate up to eight breakpoints.     

Increased body size confers greater fitness at lower experimental temperature in male Drosophila melanogaster

Genetic variation of body size along latitudinal clines is found globally in Drosophila melanogaster, with larger individuals encountered at higher latitudes. Temperature has been implicated as a selective agent for these clines, because the body size of laboratory populations allowed to evolve in culture at lower temperatures is larger. In this study, we investigated the hypothesis that larger size is favoured at lower temperature through natural selection on adult males. We measured life‐span and age‐specific fertility of males from lines of flies artificially selected for body size at two different experimental temperatures. There was an interaction between experimental temperature and body size selection for male fitness; large‐line males were fitter than controls at both temperatures, but the difference in fitness was greater at the lower experimental temperature. Smaller males did not perform significantly differently from control males at either experimental temperature. The results imply that thermal selection for larger adult males is at least in part responsible for the evolution of larger body size at lower temperatures in this species. The responsible mechanisms require further investigation.     

Excess of nonsynonymous polymorphism at Acph-1 in different gene arrangements of Drosophila subobscura

Nucleotide variation in the Acph-1 gene region was analyzed in a natural population of Drosophila subobscura from Bizerte (Tunisia). The lines studied differed in their gene arrangement for segment I of the O chromosome: 21 lines were O3+4+8, 21 were O3+4+23, and 3 were O3+4. According to chromosomal phylogenies, O3+4 is a central arrangement from which O3+4+8 and O3+4+23 originated. Strong genetic differentiation at Acph-1 was detected among the different arrangements, which is reflected in strong linkage disequilibrium between the variants at informative polymorphic sites and the type of arrangement. Estimates of silent nucleotide diversity are slightly lower within O3+4+23 (pisilent = 0.0166) than within O3+4+8 (pisilent = 0.0228) or O3+4 (pisilent = 0.0234). In contrast, nonsynonymous nucleotide diversity estimates (around 0.1%) are similar in the three arrangements. Most nonsynonymous rare variants are singletons, which results in highly significant Tajima's neutrality tests within the young O3+4+8 and O3+4+23 arrangements. This test is not significant for nonsynonymous mutations within a large Spanish O3+4 sample. In addition, a significant and marginally significant excess of nonsynonymous polymorphism was detected by the McDonald and Kreitman test within O3+4+23 and O3+4+8, respectively. This excess results in a rather high neutrality index (NI = 5.25) when both arrangements are jointly analyzed, in contrast to its value within the old O3+4 arrangement (NI = 1.74). The pattern of variation at Acph-1 within the young arrangements is unusual for nuclear genes and has the same characteristics previously detected in most genes of the mitochondrial genome. Assuming that most nonsynonymous mutations at Acph-1 are under weak negative selection, a smaller effective size of the young arrangements relative to O3+4 might explain the observed results. The relatively low frequency of O3+4+8 and O3+4+23 in the distribution area of D. subobscura, the more recent origin of these arrangements relative to O3+4 and the suppression of recombination in heterokaryotypes might contribute to the relatively small effective size of the young arrangements. Therefore, present results indicate that the differences in effective size and recombination caused by chromosomal arrangements are modulating nonsynonymous variation at Acph-1.     

Inbreeding reveals interpopulation differences in inversion polymorphism of Drosophila subobscura

The adaptive significance of inversion polymorphism of Drosophila subobscura is well established. However, little is known about gene combinations within inversions which are coadapted because of population-specific effects. We studied this aspect of Dobzhansky's coadaptation hypothesis, using the systematic inbreeding method. Differences in magnitude and quality of inversion polymorphism in two ecologically and topologically distinct habitats were compared after several generations of continuous full-sib inbreeding. Populations from the two habitats differ in frequency of homokaryotypes after third and fifth generation of inbreeding and in the levels of homozygosity of different gene arrangements. The effect of homozygosity appears population and chromosome specific.     

Continuing studies on chromosomal polymorphism in a Korean natural population of Drosophila melanogaster

Investigations on the chromosomal inversion polymorphism were conducted on a Korean (Taenung) natural population of D. melanogaster during the period 1978 to 1992. A total of 66 different endemic and cosmopolitan inversions were found on both major chromosome pairs II and III. Some of them proved to be rare cosmopolitan types (2LKA, 2LNS, 2LF, 2RCy, 3LM, 3RKI, and 3RK), while others were endemics. The distribution of breakpoints for endemic and rare cosmopolitan inversions are not random along the two autosome arms.With respect to frequency changes, the 15-year survey revealed that five of the cosmopolitan types (2Lt, 2RNS, 3LP, 3RC, and 3RMo) exhibit cyclical frequency changes, whereas gene arrangement 3RP shows relatively stable frequencies. Tests for correlations between gene arrangement frequencies and several climatic variables gave no clear evidence for such relationships. Only one correlation coefficient out of 64 was statistically significant. This revised version was published online in July 2006 with corrections to the Cover Date.     

EVOLUTION AND DEVELOPMENT OF BODY SIZE AND CELL SIZE IN DROSOPHILA MELANOGASTER IN RESPONSE TO TEMPERATURE

We examined the evolutionary and developmental responses of body size to temperature in Drosophila melanogaster, using replicated lines of flies that had been allowed to evolve for 5 yr at 25°C or at 16.5°C. Development and evolution at the lower temperature both resulted in higher thorax length and wing area. The evolutionary effect of temperature on wing area was entirely a consequence of an increase in cell area. The developmental response was mainly attributable to an increase in cell area, with a small effect on cell number in males. Given its similarity to the evolutionary response, the increase in body size and cell size resulting from development at low temperature may be a case of adaptive phenotypic plasticity. The pattern of plasticity did not evolve in response to temperature for any of the traits. The selective advantage of the evolutionary and developmental responses to temperature is obscure and remains a major challenge for future work.     

RESPONSES AND CORRELATED RESPONSES TO ARTIFICIAL SELECTION ON THORAX LENGTH IN DROSOPHILA MELANOGASTER

Two sets of four replicate lines of Drosophila melanogaster were selected for large and small thorax with controls. F, progeny of crosses between the selected lines within each size category showed (a) a reduction in preadult viability in large lines relative to control and small lines when they were cultured at medium or high density in competition with a standard mutant marked competitor stock, and (b) an increase in larval development time in large lines relative to control and small lines. Natural selection for increased body size in adults may therefore be opposed by adverse effects on larval viability. The results are discussed in terms of the developmental mechanisms probably responsible for the change in body size. The preadult survival of the large and control lines was measured at three different temperatures, and there was no evidence for a significant interaction between size and temperature. The observed evolutionary increase in body size in response to reduced temperature in Drosophila must therefore involve either different genes from those subject to selection for size at a single temperature, or a fitness component other than preadult survival. There was no significant asymmetry in response to selection, and thorax length showed heterosis in crosses between the selected lines.     

Colour-associated mating success in a polymorphic Ladybird Beetle, Harmonia axyridis

1. Body size is often an important character in mating success, but has been only infrequently mentioned in regard to colour polymorphism. In this study, mating success was investigated in a colour polymorphic Ladybird Beetle, Harmonia axyridis , with reference both to colour morph and to body size.
2. In the non-melanic males the mating individuals were significantly larger than solitary individuals, while in melanic males there was no significant difference.
3. The mating pattern was close to random mating with respect to colour morph and there was no significant deviation.
4. The results suggest both body size and colour morph affect the male mating success and males of different body size obtain mating advantage according to the colour morph. Colour polymorphism in this species is controlled by alleles on a single locus. Thus, the alleles on that locus significantly influence the effect of selection on the quantitative character.     

The evolutionary history of Drosophila buzzatii. XXXV. Inversion polymorphism and nucleotide variability in different regions of the second chromosome

Inversions are portions of a chromosome where the gene order is reversed relative to a standard reference orientation. Because of reduced levels of recombination in heterokaryotypes, inversions have a potentially important effect on patterns of nucleotide variability in those genomic regions close to, or included in, the inverted fragments. Here we report sequence variation at three anonymous regions (STSs) located at different positions in relation to second-chromosome inversion breakpoints in 29 isochromosomal lines derived from an Argentinean population of Drosophila buzzatii. In agreement with previous findings in Drosophila, gene flux (crossing over and/or gene conversion) between arrangements seems to appreciably increase as we approach the middle sections of inversion 2j, and patterns of nucleotide variability within, as well as genetic differentiation between chromosome arrangements, are comparable to those observed at the molecular marker outside the inverted fragments. On the other hand, nucleotide diversity near the proximal breakpoint of inversion 2j is reduced when contrasted with that found at the other regions, particularly in the case of derived inverted chromosomes. Using the data from the breakpoint, we estimate that the inversion polymorphism is approximately 1.63 N generations old, where N is the effective population size. An excess of low-frequency segregating polymorphisms is detected; mostly in the ancestral 2st arrangement and probably indicating a population expansion that predates the coalescent time of inversion 2j. Heterogeneity in mutation rates between the markers linked to the inversions may be sufficient to explain the different levels of nucleotide diversity observed. When considered in the context of other studies on patterns of variation relative to physical distance to inversion breakpoints, our data appear to be consistent with the conclusion that inversions are unlikely to be "long-lived" balanced polymorphisms.