Data from: The relationship between morphology and flight in Drosophila: a first approach to its genetic basis
Data files
Oct 27, 2025 version files 5.14 GB
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Flight_analysis.R
4.76 KB
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LANDMARCKS53LINES.TPS
401.92 KB
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Morphology_analysis.R
19.20 KB
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Raw_Flight_data_for_individuals.csv
56.64 KB
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Raw_Flight_data_for_lines.csv
7.84 KB
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Raw_morpho_data.csv
130.75 KB
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Raw_morphogeo_data.csv
395 KB
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README.md
16.42 KB
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Video_Example.MOV
309.21 MB
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Wings_and_Thoraxes_photos.zip
4.83 GB
Abstract
Flight is a crucial activity for winged insects, involving diverse behaviors, and wing morphology has often been proposed as a key factor influencing flight capacity. Traits such as wing loading, wing:ratio, and wing aspect ratio have been suggested as targets of natural selection, exhibiting environmental and genetic variability. Here, we evaluate the relationship between morphological traits and two aspects of flight performance: flight duration (PTF) and its robustness (CVPTF) in Drosophila melanogaster. Additionally, we included wing conformation (i.e., wing shape without the component explained by size), using geometric morphometrics. We analyzed whether variation of flight capacity can be attributed to morphological variation, employing 53 lines of Drosophila melanogaster whose genome is fully sequenced.
Dataset DOI: 10.5061/dryad.h18931zzz
Description of the data and file structure
The 53 DGRP lines used in this study were obtained from the Bloomington Drosophila Stock Center and were randomly chosen. For each line, 100 sexually mature couples were released into an egg-collecting chamber (a plastic container of 15 x 10 x 4 cm) containing a Petri dish with egg-laying medium (2% agar in distilled water and baker’s yeast). Eggs were allowed to hatch, and batches of 30 first-instar larvae were transferred to culture vials containing 5 mL of lab medium. Larvae were raised at 25±1°C and 60–70% humidity with a 12:12 light:dark photoperiod until adult emergence. For each line, we generated ten replicates (vials), with five randomly assigned to morphological measurements and the other five to flight-capacity estimates.
Morphological measurements
For morphological characterization, we randomly chose 4 to 6 flies of each sex per vial. The wings and the thorax of each individual were removed and mounted on a flat surface, always following the same arrangement. Then, they were photographed using a binocular microscope (10x) with an attached digital camera connected to a computer. We used the images to measure different morphological traits using tpsDig. For the estimation of wing traits, 15 landmarks were digitized on the ventral face of the left wing of each fly (Figure S1). We estimated wing shape (a multivariate morphological trait) using the Morpho package (Schlager, 2017). This methodology allows the separation of shape into its components: size and conformation.
The images were used to obtain also the following lineal measurements: thorax length (TL, the distance between the anterior margin of the thorax and the tip of the scutellum), total wing length (TWL, the distance between landmarks 6 and 13; Figure S1), and wing width (WW, the distance between landmarks 12 and 15; Figure S1). Also, we calculated wing area (WAR) as TWL x WW. Then, we estimated the compound traits wing loading (WLD), wing:thorax ratio (WTR), and wing aspect ratio (WAS): WLD as (TL)3/(WAR) WTR as WAR/TL, and WAS as TWL/WW.
Files and variables
File: Flight_analysis.R
Description: Code of statistical analysis for flight performance.
File: Morphology_analysis.R
Description: Code of statistical analysis for morphological measurements.
File: Raw_morpho_data.csv
Description: This data set includes flies randomly chosen of each sex per vial (between 4 to 6). The wings and the thorax of each individual were removed and mounted on a flat surface, always following the same arrangement. Then, they were photographed using a binocular microscope (10x) with an attached digital camera connected to a computer.
The images were used to obtain also the following lineal measurements: thorax length (TL, the distance between the anterior margin of the thorax and the tip of the scutellum), total wing length (TWL, the distance between landmarks 6 and 13; Figure S1 in the manuscript), and wing width (WW, the distance between landmarks 12 and 15; Figure S1). Then, we estimated the compound traits wing loading (WLD), wing:thorax ratio (WTR), and WAS as TWL/WW.
Variables
- LINE:Name or code of the line from the Bloomington Drosophila Stock Center. (Qualitative random variable).
- REPLICATE:Vial from which the fly was bred. (Qualitative random variable).
- SEX: Sex of the flies (male or female). (Qualitative fixed variable).
- Wlenght_mm:Total wing length, (TWL) the distance between landmarks 6 and 13; Figure S1, expressed in mm.
- Wwidth_mm:Wing width (WW), the distance between landmarks 12 and 15; Figure S1, expressed in mm.
- Tlenght_mm:Thorax length (TL), the distance between the anterior margin of the thorax and the tip of the scutellum, expressed in mm.
- Warea_mm2:we calculated wing area (WAR) as TWL x WW, expressed in mm2.
- WLD_mm:Wing loading was calculated as (TL x TL x TL)/(WAR), expressed in mm
- WTR_mm:Wing:thorax Ratio was calculated as WAR/TL, expressed in mm
- WAS:Wing aspect Ratio was calculated as TWL/WW.
File: Raw_Flight_data_for_individuals.csv
Description: This file contains the value of time in flight in two minutes of experiment for 1905 flies belonging to 53 lines of Drosophila melanogaster of the Bloomington Drosophila Stock Center. These lines were were randomly chosen from all stock of avalaible lines (aproximately 200).
Variables
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BLOCK: Flight time measurement date. (Qualitative random variable).
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LINE:Name or code of the line from the Bloomington Drosophila Stock Center. (Qualitative random variable).
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SEX: Sex of the flies (male or female). (Qualitative fixed variable).
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TIME_FLIGHT:Flight time in seconds during 2 minutes of experiment. (Quantitative variable)
PTF:Mean proportion of flight time for each sexual line. (Quantitative variable).
File: Raw_Flight_data_for_lines.csv
Description: This file contains the value of time in flight in two minutes of experiment for 1905 flies belonging to 53 lines of Drosophila melanogaster of the Bloomington Drosophila Stock Center. These lines were were randomly chosen from all stock of avalaible lines (aproximately 200).
Variables
- LINE:Name or code of the line from the Bloomington Drosophila Stock Center. (Qualitative random variable).
- SEX:Sex of the flies (male or female). (Qualitative fixed variable).
- PTF:Mean proportion of flight time for each sexual line. (Quantitative variable).
- SD:Standard deviation of the proportion of flight time for each sexual line. (Quantitative variable).
- CV:Coefficient of variation (SD/TFP) of the proportion of flight time for each combination of line and sex. (Quantitative variable).
- BLOCK:Flight time measurement date. (Qualitative random variable).
- WLD_mm:Wing loading was calculated as (TL x TL x TL)/(WAR), expressed in mm
- WTR_mm:Wing:thorax Ratio was calculated as WAR/TL, expressed in mm
- WAS:Wing aspect Ratio was calculated as TWL/WW.
File: Raw_morphogeo_data.csv
Description: For the estimation of wing traits, 15 landmarks were digitized on the ventral face of the left wing of each fly (Figure S1 in the manuscript). We estimated wing shape (a multivariate morphological trait) using the Morpho package. This methodology allows the separation of shape into its components: size and conformation. We employed 1077 wing photos, corresponding between 4-6 individuals per sex and line combination.
All wings were superimposed to minimise differences relative to size, position and orientation from Procrustes Analysis. This procedure generated 26 new Procrustes coordinates and eliminated four degrees of freedom, resulting in 26 shape-space dimensions. Shape variables generated afterwards, known as partial warps, indicate the partial contributions of hierarchically scaled vectors spanning a linear shape-space. Subsequently, principal components analysis of the partial warps scores matrix was conducted to obtain 26 new shape variables called relative warps (RWs). Hence, the dataset includes 26 relative warps for both sexes of lines of D. melanogaster.
Relative deformations are represented by the principal components. In this case, PC1 to PC26, and they dont have units.
On the other hand, the variables (here represented by X.1, Y.1 to X.15 and Y.15 correspond to coordinates of each landmarck in the wing.
Variables
- LINE:Name or code of the line from the Bloomington Drosophila Stock Center. (Qualitative random variable).
- REPLICATE:Vial from which the fly was bred. (Qualitative random variable).
- SEX:Sex of the flies (male or female). (Qualitative fixed variable).
- LINE_SEX: qualitative variable combination of LINE and SEX.(Qualitative random variable).
- Centroide_Size:Centroid size was calculated as the square root of the sum of the squared distances from each landmark to the centroid (geometric center) of the configuration of landmarks. The units are pixels.
- PC1:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC2:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC3:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC4:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC5:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC6:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC7:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC8:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC9:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC10:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC11:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC12:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC13:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC14:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC15:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC16:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC17:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC18:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC19:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC20:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC21:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC22:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC23:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC24:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC25:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- PC26:This column correspond to Relative warps. The response variables are the relative warps, and them do not have unities.
- 1.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 1.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 2.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 2.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 3.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 3.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 4.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 4.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 5.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 5.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 6.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 6.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 7.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 7.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 8.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 8.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 9.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 9.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 10.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 10.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 11.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 11.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 12.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 12.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 13.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 13.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 14.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 14.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 15.X:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
- 15.Y:This column correspond to coordinates of each landmarck in the wing respect the X or Y axis.
Raw Data
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LANDMARCKS53LINES.TPSThis file consists of the position of each reference point used to study the conformation and size of the wing. -
Video_Example.MOVThis is a video recorded associated with a flight of one fly included in this study. -
Wings_and_Thoraxes_photos.zipThis compressed file contains all the photos of the wings and thoraces used in this study.
This file contains all the wing and thorax photos used in this study in two different folders. The name of each photo follows this code, where the underscore separates the other parts of the code. For example, 38_A_H_THORAX_01_02The first number corresponds to the name of the isogenic line (in this case, 38).
The first letter corresponds to the name of the replicate, the vial where the flies were stored (in this case, A).
The next part of the code is a dichotomous coding that refers to the sex of the fly, where H corresponds to female and M to male.
The next part of the code is a dichotomous coding that refers to the morphological trait of the fly (THORAX or WING). Here, these terms are written in Spanish, so ALA corresponds to WING and THORAX to THORAX.
The next number, in this case 01, corresponds to the fly's identifier. It is possible that, in the case of a particular fly, the wings or thorax were damaged during the mounting of the specimen under the dissection magnifying glass, so it is possible that only one photo of the fly was available.
Finally, the last number (in this case, 02) corresponds to the photo number of this specimen. This last part of the code is irrelevant.
In addition, this file includes two photos associated with the scale used. All photos were taken at the same magnification. Each square in the photo corresponds to 1 millimeter.
Code/software
All statistical analyses were performed in R version 4.3.3 (R Core Team, 2024).
Here we provide the script for the analysis associated with flight and morphological data.
For the morphological analysis, 10-20 flies of each sex per line were randomly selected. Wings and thoraxes were dissected and mounted in a standardized way, then photographed using a binocular microscope. Morphological traits were measured from these images using tpsDig. For wing traits, 15 landmarks were placed on the ventral side of the left wing. Wing shape was estimated using the Morpho package, allowing shape to be decomposed into size and conformation components.
The images were used to obtain also the following lineal measurements: thorax length (TL, the distance between the anterior margin of the thorax and the tip of the scutellum), total wing length (TWL, the distance between landmarks 6 and 13; Figure S1), and wing width (WW, the distance between landmarks 12 and 15; Figure S1). Then, we estimated the compound traits wing loading (WLD), wing:thorax ratio (WTR), and wing aspect ratio (WAS): WLD as (TL)3/(WAR), WTR as WAR/TL , and WAS as TWL/WW. Additionally, for each morphological trait, we estimated its coefficient of variation at the line level.
We estimated flight capacity using two different traits: the proportion of time in flight (PTF) over two minutes, and its coefficient of environmental variation, which represents a measure of its robustness (CVPTF). To evaluate PTF, we used a previously tested flight chamber associated with a camera set at a 1080 x 720 resolution and a frame rate of 30 frames per second.
All flight capacity measurements were carried out in controlled temperature conditions (25±1°C), employing virgin flies of 3–7 days old. We defined flight time as the time spent by flies flying above a height of five cm inside the chamber. Finally, each recorded video was analyzed using specially designed motion-tracking software (https://github.com/galimba/drosophila). Flies that were damaged by the rod or those that could not fly were removed from the analysis. CVPTF was estimated for each sex separately as σ/μ, where μ stands for the PTF line’s mean and σ represents the standard deviation for the corresponding line.