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First published online December 28, 2007
Journal of Experimental Biology 211, 224-233 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.007575
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When wings touch wakes: understanding locomotor force control by wake–wing interference in insect wings

Fritz-Olaf Lehmann

Biofuture Research Group, Institute of Neurobiology, University of Ulm, 89069 Ulm, Germany


Figure 1
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Fig. 1. Schematic reconstruction of wake pattern during wake–wing interaction in fruit fly and dragonfly model wings. (A–D) Wake capture mechanism at dorsal and ventral stroke reversal. At the beginning of each half stroke, the leading and trailing edge vortex system generates an inter-vortex stream towards the wing. (E–H) Schematic reconstruction of vortex generation and shedding during clap-and-fling maneuver using a generic Drosophila stroke kinematics (Lehmann et al., 2005Go). Chordwise wing segments at the end of the upstroke (F), during the clap phase (G), and during the fling phase (H) before the two wings separate for the downstroke. Leading and trailing edge vortices are shed into the wake at the end of each half stroke. The low-pressure region evolving between the wings during the fling pulls fluid around the leading and the trailing wing edge into the opening cleft. (I–K) Reconstruction of vortices and local flow conditions at maximum transient lift production in tandem model wings of a dragonfly. (I) Wing kinematics follows a generic pattern, as found in various species (Maybury and Lehmann, 2004Go). (J,K) Flow characteristics at 0.35 fraction of the stroke cycle at which either the fore- (upper wing; J) or the hindwing (lower wing; K) leads wing motion by a quarter stroke cycle. Local flow vector (black) in the vector diagrams is calculated from the velocity and angle of the combined fore–hindwing downwash (green vector) determined in a region below the hindwing's surface and the translational velocity of the hind wing section (gray vector). Effective angle of attack for the hind wing section (left value) and local flow velocity (right value) are shown, respectively, in parentheses below the vector diagram. Open arrows indicate the direction of wing motion. Vortical circulation in the hindwing's leading edge vortex (LEV) is shown in parentheses. The different strengths of starting and leading edge vortices are indicated approximately by the size of the plotted vortices. Blue and red arrows represent normalized vectors of total force attenuation and enhancement, respectively, compared to a wing flapping free from mirror- (F–H) and forewing (J,K) downwash. The exact inclination of the force vectors slightly differs from the orientation normal to the wing's surface, as shown in the schematics, because of shear forces in the fluid.

 

Figure 2
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Fig. 2. Wake vorticity during fling phase in model Drosophila wings using 2D-DPIV analysis. The images were captured using a commercial 2D-DPIV system and a mini-Nd:YAG laser (TSI, Shoreview, MN, USA). (A) Wing shape and relative distance of five layers from the wing base used for analysis. Scale bar indicates time for the data traces in B–F. (B,C) Fluid vorticity fields at 0.05 stroke cycle after dorsal wing clap in a layer near the wing base (B), 0.33 (C), 0.5 (D), 0.75 wing length (E) and in a layer close to the wing tip (F). Data above each image show circulation {Gamma} in the leading edge vortex (LEV), measured in a region of interest (ROI, white box). LEV circulation when flapping one and two wings are plotted in red and black, respectively. Gray bars in the graphs indicate wing clap and time at which the DPIV analysis was performed, respectively. White lines in panels indicate wing sections. The leading edge of the dorsal wing surface is indicated by a white triangle. The magnitude of vorticity is plotted in color code and arrows correspond to the magnitude of the velocity vector at each point in the fluid, longer arrows signify larger velocities. t=stroke cycle (0–1), R=wing length. See Lehmann et al. (Lehmann et al., 2005Go) for kinematics and further details on methods.

 

Figure 3
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Fig. 3. Mean lift production in dragonfly model wings depends on both the phase-shift between fore- and hindwing stroke cycle and the vertical separation between the two wing hinges. (A) Vortical structures produced by a dragonfly hindwing at 0.35 hindwing stroke cycle. The wing moves from left to right (open arrow). The wing blade element that runs through the center of force at 0.65 wing length is indicated by a white line and the leading edge is indicated by a triangle, as shown in G. Inclination and length of the black vectors indicate direction and relative magnitude of fluid velocity. (B,C) Schematic explanation of how vertical spacing ({Delta}s) between fore- and hindwing stroke plane changes kinematic phase relationship between both wings ({Delta}t) due to vortex travel time at 0.35 stroke cycle. When the forewing leads wing motion, destruction of hindwing LEV by the forewing's start vortex depends on the phase-shift between both stroke cycles and the vertical distance between both ipsilateral wing hinges. (D) DPIV image showing the interference between vortical structures produced by fore- and hindwing at 0.15 hindwing stroke cycle. The hindwing leads wing motion by 0.25 stroke cycle. While the forewing (upper blade element) starts wing rotation at the end of the downstroke, its trailing edge vortex interferes with the LEV produced by the hindwing (lower blade element). The hindwing's start vortex still appears to be attached to the wing's trailing edge. (E,F,H,I) Vertical position (E,F) and vertical travel velocity of start vortices (H,I) shed from the fore- (black) and hindwing (blue). The vortex positions were only reconstructed from flow images in which the vortical structures were clearly visible. The data are recorded while the forewing (E,H) and the hindwing (F,I) was leading wing motion by a quarter stroke cycle, respectively. Shaded areas in H and I indicate the period during which the forewing start vortex interferes with the hindwing wake, and the dotted lines indicate maximum downwash velocity measured in the hindwing wake. R=wing length, LEV=leading edge vortex, HW=hindwing, FW=forewing.

 

Figure 4
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Fig. 4. Modulation of lift with changing phase-shift between the stroke cycles of dragonfly tandem wings. (A,B) Data show modulation of forewing (A) and hindwing (B) lift averaged over the entire flapping cycles of four subsequent strokes, while phase lag systematically varied between –50% (forewing lead by a half stroke cycle) and 50% (hindwing leads by a half stroke cycle). Colors show measurements at different vertical distances between the stroke planes (1.25 to 5 mean wing chord c). (C) Phase-shift between the stroke cycles of both wings during mean peak lift changes with changing vertical distance between the two wing hinges. Black, forewing; red, hindwing; aspect ratio of the wings=2.7; flapping frequency=666 mHz; wing rotation symmetrical, starting 10% prior to and ending 10% after stroke reversal; Reynolds number=137; c=4.0 cm. Data in A and B were smoothed using a numerical 3 data point adjacent averaging filter.

 

Figure 5
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Fig. 5. Phase-shift induced modulation of mean lift production depends on wing length in tandem model wings. The length of the model wings was 6.9 (light gray), 7.7 (blue), 8.5 (purple), 9.3 (orange), 10.1 (black), 10.9 (red), 11.7 (green) and 12.5 cm (dark gray). (A) Lift modulation of the lower hindwing (12.5 cm wing length), while length of the upper forewing varied between 6.9 and 12.5 cm. (B) Lift modulation in length-changing forewing and length constant hindwing of 12.5 cm. (C) Combined fore- and hindwing lift as shown in A and B, respectively. (D) Hindwing lift modulation during length changes of the hindwing. Forewing length is constant at 12.5 cm. (E) Peak phase at which fore- (black) and hindwing (red) produce maximum mean lift. Data are plotted against the ratio between hind- and forewing length. A value of 1.0 means that both wings have equal length (12.5 cm). (F) Strength of peak-to-peak modulation of fore- (black) and hindwing (red) lift production. Modulations were derived from sinusoidal fits to the data set (forewing lift: fit on data between ±15% phase-shift, hindwing lift: fit on all data) as shown by the colors in A–D. Insets in E show the shape of the model wings used in the experiments. The rounded wing tip was similar in all wings while total wing length varied between minimum and maximum values. Aspect ratio varied between approximately 1.7 (6.9 cm wing length) and 3.1 (12.5 cm wing length), respectively. Wing chord=4.0 cm. Vertical distance between both wings was 1.25 mean wing chord. For kinematic pattern, see legend of Fig. 4. More information on the methods is given elsewhere (Maybury and Lehmann, 2004Go).

 





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