An interspecific QTL study of Drosophila wing size and shape variation to investigate the genetic basis of morphological differences.

The Drosophila wing has been used as a model in studies of morphogenesis and evolution; the use of such models can contribute to our understanding of mechanisms that promote morphological divergence among populations and species. We mapped quantitative trait loci (QTL) affecting wing size and shape traits using highly inbred introgression lines between D. simulans and D. sechellia, two sibling species of the melanogaster subgroup. Eighteen QTL peaks that are associated with 12 wing traits were identified, including two principal components. The wings of D. simulans and D. sechellia significantly diverged in size; two of the QTL peaks could account for part of this interspecific divergence. Both of these putative QTLs were mapped at the same cytological regions as other QTLs for intraspecific wing size variation identified in D. melanogaster studies. In these regions, one or more loci could account for intra- and interspecific variation in the size of Drosophila wings. Three other QTL peaks were related to a pattern of interspecific variation in wing size and shape traits that is summarized by one principal component. In addition, we observed that female wings are significantly larger and longer than male wings and the second, fourth and fifth longitudinal veins are closer together at the distal wing area. This pattern was summarized by another principal component, for which one QTL was mapped.

Quantitative trait analysis and geographic variability of natural populations of Zaprionus indianus, a recent invader in Brazil.

Five natural samples of a recent South America invader, the drosophilid Zaprionus indianus, were investigated with the isofemale line technique. These samples were compared to five African mainland populations, investigated with the same method. The results were also compared to data obtained on mass cultures of other populations from Africa and India. Three quantitative traits were measured on both sexes, wing and thorax length and sternopleural bristle number. We did not find any latitudinal trend among the American samples, while a significant increase in body size with latitude was observed in the Indian and, to a lesser degree, in the African populations. American populations were also characterized by their bigger size. Genetic variability, estimated by the intraclass correlation among isofemale lines, was similar in American and African populations. The intraline, nongenetic variability was significantly less in the American samples, suggesting a better developmental stability, the origin of which is unclear. A positive relationship was evident between intraline variability of size traits and the wing-thorax length correlation. Altogether, our data suggest that the colonizing propagule introduced to Brazil had a fairly large size, preventing any bottleneck effect being detected. The big body size of American flies suggests that they came from a high-latitude African country. The lack of a latitudinal dine in America seems to be related to the short time elapsed since introduction. The very rapid spread of Z. indianus all over South America suggests that it might rapidly invade North America.

Genetic architecture of wing morphology in Drosophila simulans and an analysis of temperature effects on genetic parameter estimates.

The Drosophila wing has been used as a model to investigate the mechanisms responsible for size and shape changes in nature, since such changes might underlie morphological evolution. To improve the understanding of wing morphological variation and the interpretation of genetic parameters estimates, we have established 59 lines from a Drosophila simulans laboratory population through single pair random matings. The offspring of each line were reared at three different temperatures, and the wing morphology of 12 individuals was analyzed by adjusting an ellipse to the wings' contour. Temperature, sex and line significantly affected wing trait variation, which was mainly characterized by longer wings having the second, fourth and fifth longitudinal veins closer together at the wing tip. As for the genetic parameter estimates, while the cross-environment heritability of some traits, such as wing size (SI), decreased with an increasing difference between the temperatures at which parents and offspring were reared, wing shape (SH) heritability did not seem to change. Since we found indications that neither an increase in the phenotypic variation nor the occurrence of genotype-environment interactions could fully explain the low heritabilities of SI estimated by cross-environment regressions, we discuss the importance of other effects for explaining this discrepancy between the SI and SH heritability estimates. In addition, although the genetic matrix was not entirely represented in the phenotypic matrix, several correspondences were identified, suggesting that the observed patterns of wing morphology variation are genetically controlled.

Plasticity of Drosophila melanogaster wing morphology: effects of sex, temperature and density.

In this paper we use an adjusted ellipse to the contour of the wings of Drosophila as an experimental model to study phenotypic plasticity. The geometric properties of the ellipse describe the wing morphology. Size is the geometric mean of its two radii; shape is the ratio between them; and, the positions of the apexes of the longitudinal veins are determined by their angular distances to the major axis of the ellipse. Flies of an inbred laboratory strain of Drosophila melanogaster raised at two temperatures (16.5 degrees C and 25 degrees C) and two densities (10 and 100 larvae per vial) were used. One wing of at least 40 animals of each sex and environmental condition were analyzed (total = 380), a measurement of thorax length was also taken. Wing size variation could be approximately divided into two components: one related to shape variation and the other shape independent. The latter was influenced primarily by temperature, while the former was related to sex and density. A general pattern could be identified for the shape dependent variation: when wings become larger they become longer and the second, fourth and fifth longitudinal veins get closer to the tip of the wing.

Size and shape heritability in natural populations of Drosophila mediopunctata: temporal and microgeographical variation.

'Traditional morphometrics' allows us to decompose morphological variation into its major independent sources, identifying them usually as size and shape. To compare and investigate the properties of size and shape in natural populations of Drosophila mediopunctata, estimating their heritabilities and analysing their temporal and microgeographic changes, we carried out collections on seven occasions in Parque Nacional do Itatiaia, Brazil. In one of these collections, we took samples from five different altitudes. Measurements were taken from wild caught inseminated females and up to three of their laboratory-reared daughters. Through a principal component analysis, three major sources of variation were identified as due to size (the first one) and shape (the remaining two). The overall amount of variation among laboratory flies was about half of that observed among wild flies and this reduction was primarily due to size. Shape variation was about the same under natural and artificial conditions. A genetic altitudinal cline was detected for size and shape, although altitude explained only a small part of their variation. Differences among collections were detected both for size and shape in wild and laboratory flies, but no simple pattern emerged. Shape variation had high heritability in nature, close to or above 40% and did not vary significantly temporally. Although on the overall size heritability (18 +/- 6%) was significant its estimates were not consistent along months--they were non-significant in all but one month, when it reached a value of 51 +/- 11%. Overall, this suggests that size and shape have different genetic properties.

Heritability, phenotypic and genetic correlations of size and shape of Drosophila mediopunctata wings.

We have studied the morphology of wings of Drosophila mediopunctata employing the ellipse method, a procedure that allows precise descriptions of wing size (SI), wing shape outline (SH), and placement of longitudinal wing veins. We have found that the SH and the points which determine the position of the apices of the third, fourth and fifth longitudinal wing veins show high heritability in nature (the lower bound for the natural heritability is above 0.25). The values found are similar to those obtained for the broad-sense heritabilities (H2) in the laboratory. However, SI and the point which determines the apex of the second longitudinal wing vein showed small lower bounds for heritability in nature, 0.05 and 0.07, respectively, in spite of the high estimates of H2 in the laboratory. These results suggest that size and shape have different genetic properties. We observed a high positive phenotypic correlation between the SH, the fourth and the fifth longitudinal wing veins, which contrasts with a negative correlation between these traits and the second longitudinal vein. That is, as the SH gets longer, the apices of the second and fifth veins become closer to each other. Positive genetic correlations in the field were detected between SH, the fourth and the fifth longitudinal veins and also between the third and the fourth veins.

Variation and heritability of aristal morphology in a natural population of Drosophila mediopunctata.

We studied the major sources influencing the variation of the number of aristal branches in a natural population of Drosophila mediopunctata. Flies were collected on six occasions at different altitudes in Parque Nacional do Itatiaia (Brazil). The progenies of these flies were reared in the laboratory at 16.5 degrees C. The number of aristal branches ranges from 11 to 15 and is influenced by sex. Estimates of the natural heritability showed that at least 20% of the total phenotypic variation is due to additive genetic variation. Although the heritability of this trait estimate in the laboratory was larger (42%), the difference between the two estimates is not statistically significant. Thus, for the number of aristal branches, laboratory estimates of heritability provide reasonable estimations of both the magnitude and significance of heritabilities in nature. The mean numbers of aristal branches in the wild-caught flies from different altitudes or months are homogeneous. The same was observed for the means of its progeny kept in the laboratory under controlled conditions. On the other hand, wild-caught females have significantly fewer aristal branches than their laboratory-raised daughters, which suggests that an environmental factor or factors may have an important influence on this trait.

Morphological variation in a natural population of Drosophila mediopunctata: altitudinal cline, temporal changes and influence of chromosome inversions.

To characterize the morphological variation in a natural population of Drosophila mediopunctata, males were collected on three occasions at a single locality. From each wild-caught male 14 body measures were taken and the karyotype for inversions on chromosomes X and II was determined. Through a principal components analysis, two sources of variation, identified as size and shape, accounted for approximately 80 and 6 per cent of the total morphological variability, respectively. The shape component was determined primarily by variations in the position of the wing second longitudinal vein. Differences between collections were detected both for size and shape. An altitudinal cline was observed in respect of wing shape, although altitude explained only a small part of the shape variation. Size and shape were affected by chromosome II inversions. However, in respect of size, no direct differences were detected between karyotypes but a significant interaction between collecting date and karyotype was found. This suggests that karyotypes might differ in their norms of reaction in the field.