Drosophila Wing Integration and Modularity: A Multi-Level Approach to Understand the History of Morphological Structures.

Abstract

Simple SummaryThe diverse components of any morphological structure are integrated with respect to each other since they have developed, functioned, and evolved together, a phenomenon known as integration. However, this integration is not absolute but organized in units (i.e., modules) that are relatively independent while participating to generate a structure that acts as a functional whole. Even though most of the studies on modularity and integration have focused on variation among individuals within populations, there are more levels of variation that exhibit modularity and integration, deriving from distinct sources such as genetic variation, phenotypic plasticity, fluctuating asymmetry, evolutionary change, among others. Consequently, the present study focused on analysing the integration and modularity of the wing shape of some of the best-known model organisms, i.e., the genus Drosophila, at the static, developmental, and evolutionary levels to acquire a better insight about how modularity and integration act at different analytical levels. The strong integration and overall similarities observed in the variation pattern at multiple levels suggest a shared mechanism underlying the observed variation in Drosophila’s wing shape and added a new piece of evidence of stasis in the evolutionary history of Drosophila wing.Static, developmental, and evolutionary variation are different sources of morphological variation which can be quantified using morphometrics tools. In the present study we have carried out a comparative multiple level study of integration (i.e., static, developmental, and evolutionary) to acquire insight about the relationships that exist between different integration levels, as well as to better understand their involvement in the evolutionary processes related to the diversification of Drosophila’s wing shape. This approach was applied to analyse wing evolution in 59 species across the whole genus in a large dataset (~10,000 wings were studied). Static integration was analysed using principal component analysis, thus providing an integration measurement for overall wing shape. Developmental integration was studied between wing parts by using a partial least squares method between the anterior and posterior compartments of the wing. Evolutionary integration was analysed using independent contrasts. The present results show that all Drosophila species exhibit strong morphological integration at different levels. The strong integration and overall similarities observed at multiple integration levels suggest a shared mechanism underlying this variation, which could result as consequence of genetic drift acting on the wing shape of Drosophila.