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First published online February 12, 2007
Journal of Experimental Biology 210, 881-896 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.02704
A computational investigation of the three-dimensional unsteady aerodynamics of Drosophila hovering and maneuvering
Laboratory for Computational Physics and Fluid Dynamics, Naval Research Laboratory, Washington, DC 20375, USA
* Author for correspondence (e-mail: ravi{at}lcp.nrl.navy.mil)
Accepted 22 December 2006
| Summary |
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-like
structures that have not been previously observed in the wakes of flapping
fliers.
Key words: flapping wings, insect flight, insect hovering, insect maneuvering, Drosophila, incompressible flow, unstructured grid, unsteady aerodynamics
| Introduction |
|---|
|
|
|---|
Flapping foil propulsion has received considerable attention in the past
few years as an alternative to the propeller. Experimental and computational
investigations have been carried out on several fronts to gain insight into
insect flight performance and use those insights in vehicle design. Kellogg et
al. (Kellogg et al., 2001
;
Kellogg et al., 2003
) and Jones
and Platzer (Jones and Platzer,
2003
) have experimentally investigated the use of tandem flapping
foils for micro-air vehicle propulsion and were successful. Ramamurti et al.
(Ramamurti et al., 2005
)
computationally studied the unsteady thrust and lift generation of tandem
multiple flapping wing vehicles as well as rotary-wing vehicles to support
MicroAirVehicle (MAV) design efforts. For these types of vehicles, it is
important to accurately quantify the performance of flapping wings in order to
provide the controllability needed during a maneuver.
The effects of the wing rotation have been studied in the fruit fly,
Drosophila (Dickinson et al.,
1999
). 3-D unsteady flow computations over an insect wing
(Liu and Kawachi, 1998
) were
used to obtain qualitative agreement for the flow patterns over a hovering
hawkmoth with the visualizations previously obtained in a windtunnel
(Willmott and Ellington,
1997
). Force production in the single flapping wing of
Drosophila has also been studied
(Ramamurti and Sandberg, 2002
;
Sun and Tang, 2002
). Ramamurti
et al. also studied the force production about the flapping and deforming
pectoral fin and body of the swimming bird wrasse fish computationally
(Ramamurti et al., 2002
).
Their experimental results provided good agreement with measured force data
(Dickinson et al., 1999
) and
fish acceleration data (Walker and
Westneat, 1997
). Since one would like to know the importance of
wing deformation time history to lift and thrust production time history,
Ramamurti et al. investigated the fluid dynamics underlying the generation of
forces during pectoral fin oscillation as fin rigidity and resulting shape
deformation time history is varied
(Ramamurti et al., 2004
). All
of these investigations focused primarily on lift and thrust production
mechanisms for forward flight or straight ahead swimming.
Recently the question of force production during insect turning has been
addressed. Fry et al. (Fry et al.,
2003
) studied the wing and body kinematics of free flying
Drosophila performing rapid maneuvers. They measured insect wing and
body kinematics and used that data to drive a robotic model of the fruit fly.
They measured the total aerodynamic force and torque during the specified
saccade maneuver of the fly model. The results of Fry's work indicated that
inertia rather than viscous effects dominated the torque production. The
details of the unsteady aerodynamics of force and yaw moment production by
each of the two asymmetrically flapping wings interacting with the fly body
and how those were related to the overall insect torque production was still
an open question. We address that question with the computational
investigation reported here.
The primary objectives of this study were to (i) computationally simulate
the maneuver of the fruit fly using prescribed wing and body kinematics
observed from experiments, (ii) to validate the computed forces with the
experimentally measured forces, (iii) to compute the unsteady aerodynamics of
force production by each asymmetrically flapping wing, and (iv) to understand
the subtle changes in kinematics observed by Fry et al.
(Fry et al., 2003
) that lead to
the net measured torque required for the maneuver. In this manner, a
computational investigation can complement physical observations and
measurements by providing the unsteady pressure time-histories on all moving
surfaces and the detailed velocity and vorticity map of the evolving flow
field associated with either measured or derived total force data. This can be
of value not only to biomimetic vehicle designers but also to the biological
community, since knowledge of surface pressure distribution time histories
during insect, bird or fish locomotion and maneuvering is essential to
understanding the bone, muscle and skin external loading experienced by these
creatures as they execute such motions. The surface pressure time-histories on
insect bodies and flapping wings are the crucial missing unsteady data that
cannot be measured in insect free flight, and only with great difficulty on
models. The biologist can now use this unsteady computed data to explore
questions such as why the wing hinge joint and wing root have the shape and
structure that they have. One possibility is that the shape and structure have
evolved to enable the object or prey avoidance maneuvers observed and hence
must be capable of supporting the wing hinge time-varying moments throughout a
maneuver. That is, the insect physiology and biomechanics enable it to operate
successfully in its environment.
There are many aspects to consider when investigating the insect's dynamic behavior in an unsteady environment and what is required for success. External stimuli can be optical, thermal, chemical and mechanical. Kinematics of insect responses to such stimuli can be measured by biologists and provided as input to 3-D unsteady aerodynamics computations. The resulting force and moment time histories and wake vorticity evolutions can then assist in answering functional morphology questions by providing otherwise unavailable information on what structural and dynamic characteristics the insect must possess to enable its dynamic responses. Computational investigations directed at answering morphology questions may then lead to the answers to other questions, such as whether the structure of the wing hinge joint, for example, has evolved to enable the saccade maneuver or possibly some other dynamic behavior such as stable flight in a wind gust, where an unsteady pressure field provides the external stimuli. Investigation of wind-borne chemical stimuli and subsequent dynamic responses is another area where computations can be of assistance. Examination of the geometry and location of chemical receptors in concert with the insect unsteady flight aerodynamics may provide insights into how the receptors enable successful tracking of pheromones or other chemicals of interest in a random wind-gust environment. Do the energy demands of flight in wind gusts restrict operations above a certain magnitude or is it a biomechanical limitation that dominates? There are numerous biological questions of this sort that are not answerable solely by laboratory or field measurements but on which progress can be made by complementary investigations that include the unsteady aerodynamic computations we describe below.
| Materials and methods |
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|
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The incompressible flow solver
The governing equations employed are the incompressible NavierStokes
equations in Arbitrary Lagrangian-Eulerian (ALE), formulation which are
written as:
![]() | (1) |
![]() | (2) |
and are discretized in time using an implicit time
stepping procedure. Thus the equations are Eulerian for zero mesh velocity and
Lagrangian if the mesh velocity is the same as the flow velocity. The present
time-accurate flow solver is discretized in space using a Galerkin procedure
with linear tetrahedral elements. The details of the flow solver have already
been discussed extensively elsewhere
(Ramamurti and Löhner,
1992
|
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| Results and discussion |
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The body of the fruit fly in Fig.
2 is rotated along the x-axis through a roll angle
,
followed by a yaw rotation of
about the y-axis, and the by a
pitch rotation of
about the z-axis. This set of rotations is
equivalent to the Euler angle systems commonly used in aeronautical
engineering. The coordinate transformation for this set of rotations is given
elsewhere (Greenwood, 1987
) and
is as follows.
![]() | (3) |
|
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|
The rotation angles for the wings are obtained from the vectors from the
wing hinges (RWH and LWH) to the wing tips (RWT and LWT) and the wing normal
vectors (RWN and LWN), e.g. RWHRWT and RWN vectors shown in
Fig. 1. For this purpose, the
body is pitched at an average pitch angle of 44.4° obtained over the first
6 hover cycles. The resulting stroke angle (
), the deviation from the
xz plane (
) and the angle of attack (
) of the wings
are shown in Fig. 5. The stroke
angles of the right and left wings are similar during hover,
t=0.00.03 s, and towards the end of the saccade maneuver,
t=0.07 s onwards, and differ considerably during the saccade, as seen
in Fig. 5A. A similar trend in
the deviation and angle of attack is shown in
Fig. 5B,C. In order to see the
differences in the angles between the two wings, the instantaneous differences
of these angles were plotted in Fig.
6A. From this figure it can be seen that the differences vary
during each wing beat cycle and no clear trend is observed. Hence, these
differences were filtered using an fft-filter and the results plotted in
Fig. 6B. It is clear that the
stroke angle undergoes the largest change during the maneuver. The left wing
lags the right wing by approximately 13° midway during the saccade, at
t=0.05 s approximately, and recovers at the end of the maneuver. The
difference in the angle of attack between the two wings shows that the left
wing is maintained at a smaller angle of attack by approximately 6°
throughout the maneuver. The difference in the deviation angle of the right
and left wings shows that the right wing is at a slightly higher elevation
angle during the maneuver.
Unsteady computations of forces and moments
Using the kinematics described above, the unsteady flow over a maneuvering
fruit fly was computed using the incompressible flow solver. The instantaneous
surface pressure distribution on the wings and the body of the fruit fly was
integrated to produce the time history of forces. These forces were resolved
in the y-direction to yield the lift force and the resultant of the
force in the x- and z-directions along the tailhead
vector to yield the forward horizontal force.
The results are shown in Fig.
7 and are compared with the experimentally obtained forces of Fry
et al. (Fry et al., 2003
). The
comparison shows that the computed results agree extremely well with the
experiments throughout the maneuver. From
Fig. 7A, it is clear from the
experimental measurements that the lift peaks during the upstroke are almost
twice as large as the peaks during the downstroke during hovering and in the
initial period of the saccade maneuver, from t=0.0 to 0.05 s. At the
beginning of the saccade, the computed lift during the downstroke is higher
than the measured lift while the computed lift on the upstroke is slightly
less than the measured lift. During the later part of the maneuver, the peak
in the lift force drops in the upstroke part of the cycle and the peak during
the downstroke remains similar to the pre-saccade value. The computed lift is
very close to the measured lift, on both the upstroke and the downstroke, as
the saccade maneuver progresses. The thrust force peaks to a value of nearly
25 µN during the upstroke during the hover and early stages of the saccade
and decreases during the later part of the maneuver. The computed thrust and
drag force time history is nearly identical to the measured thrust throughout
the entire maneuver. The peak drag force during the downstroke decreases
continuously during the maneuver.
The yaw moment experienced by the fruit fly during the maneuver is computed
about the mid point between the head and tail along an axis normal to the
headtail vector and the right wing hinge to the left wing hinge vector,
shown in Fig. 8. The
contributions of the yaw moment from the right and left wings are shown in
Fig. 9A. It is clear that most
of the yaw moment is created during the upstroke and the two wings produce
moments in the opposing direction, as would be expected. The mean moment
created by the right wing is slightly larger that that of the left wing
producing a net yaw moment. Fig.
9B shows the total instantaneous yaw moment created by the right
and the left wings. This plot does not show any continuous yaw moment
production during the saccade maneuver, t=0.030.07 s, although
there are instants where the net yaw moment is large, e.g. t=0.042 s,
0.052 s and 0.07 s. Also, it can be seen that the yaw moment is relatively
small during the downstroke compared to the large excursions during the
upstroke. This total yaw moment is then filtered through an fft-filter and the
result is shown in Fig. 9C. The
yaw moment continuously increases during the pre-saccade period and reaches a
maximum value of 2x109 Nm at t=0.03 s, and a
local maximum of 5x1010 Nm between t=0.05 s
and 0.07 s. This yaw moment behavior is very similar to the torque production
described by Fry et al. (Fry et al.,
2003
). The experimental torque shown in fig. 3C of Fry et al.
(Fry et al., 2003
) was been
obtained using a low pass filter and averaging over six species. The use of an
fft-filter can be justified by the fact that the fruit fly torque is dominated
by the inertia (Fry et al.,
2003
). Moreover, the body of the fruit fly may not be able to
respond to the instantaneous changes in forces and moments at the flapping
frequency of the wings. Hence, in this study the data was filtered with a
maximum frequency of 60 Hz, and both raw data and filtered data are presented.
The mean yaw moment during each cycle is also shown in
Fig. 9C and is very similar in
both magnitude and trend compared to the filtered moment. The filtering
process used here seems to be a method to obtain the mean cycle values without
the knowledge of beginning and end of each flapping cycle.
|
Origin of the yaw moment
In order to find the origin of this yaw moment, the force vector acting on
each wing is decomposed into its forward, lift and sideways components. For
this purpose, the forward direction is taken to be the vector from tail to
head of the insect and parallel to the xz plane, as shown in
Fig. 8. The lift is in the
direction normal to the flight and is aligned along the y-axis, i.e.
the vertical component of force. The sideways direction is then defined
orthogonal to the lift and forward directions.
Fig. 10A shows the forward component of the computed forces produced by the wings during the entire maneuver. During hovering, the thrust produced during the upstroke is nearly anti-symmetric to the force produce during the downstroke and the contributions from the right and left wings are nearly the same. During the saccade, t=0.03 s0.07 s, there is a considerable increase in the thrust produced during the upstroke compared to the drag produced in the downstroke, and the right wing produces slightly larger thrust. Fig. 10B shows the difference in the thrust produced by the right and the left wings and exhibits both positive and negative values. Hence, this difference in the forward force is further filtered and is shown in Fig. 10C. From this figure it is clear that the right wing produces additional forward force compared to the left wing during the saccade maneuver. The differences in the sideways and lift components of the forces between the right and left wings were filtered in a similar manner and are shown in Figs 11 and 12. The differential side force component remains positive during the maneuver with a peak value of 1.78 µN at the beginning of the maneuver, t=0.045 s in Fig. 11B, compared with a peak value of 0.8 µN at t=0.035 s in Fig. 12B, and therefore could contribute to the yaw moment. The differential lift force is positive in the initial period of the maneuver and becomes negative in the later half of the maneuver.
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Fig. 14A,B show the length of the moment arms for the forward and side forces, respectively. These moment arms were obtained from the center of pressure (CP) for each wing and computing the distances of these CP to the center of rotation along the side and forward directions, shown in Fig. 8. From Fig. 14A, it can be seen that the center of pressure for the right wing is located away from the center along the side ways direction throughout the maneuver.
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During the later phases of the saccade, the moment arm of the right wing is larger in magnitude thus producing the restoring negative torque. Fig. 15A shows the distance of the center of pressure from the center of rotation for each wing. The center of pressure for the right wing is located at nearly 2.5 mm from the center of rotation midway between the head and tail of the insect and that of the left wing is at 2.0 mm throughout the saccade. Fig. 15B shows the location of the center of pressure from the wing hinges along the spanwise direction. This distance is nearly constant at 1.4 mm throughout the maneuver, which is 60% of the span of the wing.
Force production and surface pressures during 3 typical cycles
Fig. 16 shows the wing
motion and the force vectors generated by the wings during a typical hover
cycle. The wing motion shown by the leading edge dot and the orientation of
the chord shows a symmetric or U-shaped profile both during the downstroke and
upstroke. The motion is similar for the right and left wings. The force
production is also similar during this cycle as expected. The force vector
shown in here is the resultant of the lift and the thrust components and does
not include the side force component.
At the beginning of the saccade maneuver, the wing motion changes from the symmetric shape to a `banana-shape' during the downstroke and the upstroke remains relatively symmetric, as shown in Fig. 17. The forces on the two wings also show differences, the right wing producing larger force during the downstroke and the later half of the upstroke.
|
Fig. 19 shows the pressure distribution on the surface of the wings during a hover cycle and is almost symmetric between the right and left wings. Hence, the forces produced by the wings are also symmetric. At an instant midway during the downstroke, t=0.0169 s shown in Fig. 19A, the lift achieves a maximum. At this instant, a low-pressure region is visible on the top of the wings and a high-pressure region is present on the bottom of the wings. At t=0.0176 s, just before stroke reversal shown in Fig. 19B, a small low pressure region is visible on the top of the wings in the leading edge region and at mid span. The pressure on the bottom of the wings is nearly constant at this instant. Fig. 19C shows the pressure distribution midway during the upstroke with a high-pressure region on the lower side of the wing. The extent of the high-pressure region is larger and the suction pressure is lower compared to the middle of the downstroke, leading to a higher lift on the upstroke. Fig. 19D shows the pressure distribution just after stroke reversal and is nearly symmetric between the right and left wings leading to a force vector opposing each other.
Fig. 20 shows the pressure distribution during the first cycle of the saccade maneuver. At t=0.0299 s, the force vectors oppose each other and with the wings aligned nearly in the vertical direction producing zero lift. The pressure distributions on the right and left wings do not exhibit any symmetry during this cycle with the bottom left wing showing higher pressure extending over a larger region. At midway through the downstroke, t=0.0310 s, a maximum lift is produced. At stroke reversal, the pressure is nearly constant on the bottom of the wing at t=0.0324 s. At t=0.0334 s, the lift produced reaches a peak. The instantaneous yaw moment also reaches a peak at this time. The lift and the thrust produced reduce toward the end of the cycle, close to stroke reversal at t=0.0341 s. The yaw moment produced during the downstroke is nearly zero for each of the wings whereas during the middle of the upstroke the yaw moment reaches an appreciable value, albeit the left wing producing a negative moment opposing the right wing.
|
Vorticity evolution during hover and saccade cycles
Fig. 22 shows the sequence
of evolution of the vorticity generated by the leading and trailing edges of
the wings during a hover cycle. At t=0.2309 s, at stroke reversal we
see vortices generated from the leading and trailing edges. At
t=0.0233 s the vortices from the trailing edges are pulled up due to
the wing motion and continue to get stretched at t=0.02358 s where
they form an arch-like structure below the head. At t=0.02394 s, the
wings are in the middle of the upstroke with the vortex from the leading edge
forming a loop as the wing sweeps back to a position close to stroke reversal
at t=0.02453 s. At stroke reversal, t=0.02493 s, the
arch-like vortices that were shed at the end of the previous downstroke are
pronounced and soon after reversal, this structure pinches off of the loop,
t=0.02527 s, forming a
-like structure, which follows
downstream until t=0.02729 s.
Fig. 23 shows this evolution
viewed from the back of the hovering fruit fly. At t=0.02233 s, the
-like structure connected to the loop can be seen. At
t=0.02309 s, the arch pinches off from the loop and at
t=0.02333 s, this arch joins the
-structure from the front
and pinches off from it at t=0.02376 s. Between t=0.02413 s
and 0.02493 s, the wing is executing the upstroke producing the loop structure
and at t=0.02552 s, just after stroke reversal, we can see the
-like structure emerging.
|
|
-like structure generated from the previous upstroke. This structure,
in contrast to the hover cycle exhibits asymmetry in the lengths of the legs
mainly since the leg on the left side is formed due to an earlier pinching off
compared to the right side. At the stroke reversal, t=0.05662 s, we
see the leading edge vortex being shed and drawn into a loop throughout the
upstroke. | Summary and conclusions |
|---|
|
|
|---|
The yaw moment reached a peak value at the beginning of the maneuver,
remained positive throughout the remainder of the maneuver, and exhibited a
restoring moment after the maneuver. The origin of the yaw moment was
investigated by computing the center of pressures on each wing and the
individual moment arms. This showed that the forward force and a component of
the lift force produced the turning moment while the side force produced the
restoring torque during the maneuver. The vorticity shed from the wing leading
edge and the tips shows a loop-like structure, which during stroke reversals
pinched off into
-like structures that were advected downstream. The
role of these vortex structures on the force production mechanism is being
investigated.
| Acknowledgments |
|---|
| References |
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