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
Ravi Ramamurti* and
William C. Sandberg
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
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Summary
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Three-dimensional unsteady computations of the flow past a fruit fly
Drosophila under hovering and free flight conditions are computed.
The kinematics of the wings and the body of the fruit fly are prescribed from
experimental observations. The computed unsteady lift and thrust forces are
validated with experimental results and are in excellent agreement. The
unsteady aerodynamic origin of the time-varying yaw moment is identified. The
differences in the kinematics between the right and the left wings show that
subtle change in the stroke angle and deviation angle can result in the yaw
moment for the turning maneuver. The computed yaw moment reaches a peak value
at the beginning of the maneuver and remains positive throughout the remainder
of the maneuver. The origin of the yaw moment is investigated by computing the
center of pressures on each wing and the individual moment arms. This
investigation leads to the conclusion that it is the forward force and a
component of the lift force that combine to produce the turning moment while
the side force alone produces the restoring torque during the maneuver. The
vorticity shed from the wing's leading edge and the tips show a loop like
structure that during stroke reversals pinches off into
-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
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Introduction
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Flapping wing locomotion and maneuvering mechanisms are of interest to
behavioral biologists, biomechanics researchers, and engineers attempting to
develop systems that can match the performance of living creatures. The flight
dynamics of insects are quite complex. For example, the motion of the fruit
fly Drosophila melanogaster involves several successive sharp right
angle turns. These turns, called saccades, are characteristic of many insects.
Understanding the three-dimensional (3-D) unsteady aerodynamics of the two
flapping wings and the insect body that results in the saccade maneuver would
be extremely valuable to designers in their attempts to create vehicles
capable of emulating such obstacle-avoidance performance. Unlike rotary wing
flight that produces a steady torque on the body, flapping flight should
generate an oscillatory torque. There are numerous questions one would wish to
answer before embarking on a vehicle design for saccade-like maneuvering
performance. A particularly important question is `How does the unsteady flow
field about multiple flapping wings and an insect body interact and evolve to
produce lift and thrust?' and secondly, `If the flapping is asymmetric what is
the instantaneous flow field and how does it evolve to produce the desired
maneuvering moment?' Or to put it another way, `How much asymmetry between
multiple flapping wings is needed to achieve a specific maneuvering moment and
what are the underlying aerodynamics of the moment generation?' Also, insect
flight is largely in the laminar regime, with vortices being shed persistently
from the leading edge and the wing tips. Understanding the role of these shed
vortices on the lift and thrust development is also important.
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.
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Materials and methods
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To study the aerodynamics of the maneuvering flight, Fry et al.
(Fry et al., 2003
) captured the
saccade of the fruit fly, Drosphila melanogaster, using a three
high-speed video camera system. The flies were untethered and were enticed
towards a target with a drop of vinegar. Some of the flies performed collision
avoiding maneuvers, resulting in the measurement of the body and wing
kinematics throughout the maneuver that were used in the present study.
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) |
In Eqn 1, p denotes the pressure,
va=vw is the advective velocity vector,
where v is the flow velocity and w is the mesh velocity and the
material derivative is with respect to the mesh velocity w. Both the
pressure p and the viscous stress tensor have been normalized by the
(constant) density
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
; Ramamurti et al.,
1994
; Ramamurti et al.,
1995
; Ramamurti et al.,
1999
) in connection with successfully validated solutions for
numerous 2-D and 3-D, laminar and turbulent, steady and unsteady flow
problems.

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Fig. 1. Schematic of the hovering Drosophila. LWT, RWT, left/right
wingtip; LWH, RWH, left/right wing hinge; LWN, RWN, left/right wing normal
vector.
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Results and discussion
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The unsteady flow past an isolated Drosophila wing has been
computed (Ramamurti and Sandberg,
2002
), and the effect of phasing between the translational and
rotational motions on the thrust production of a hovering fruit fly was
studied. In the present study, we extend this work and compute the unsteady
forces and moments produced by both flapping wings and including the body of
the fruit fly as it hovers and then executes a saccade maneuver. The
feflo incompressible flow solver described briefly above is used to
compute the unsteady 3-D flow. The kinematics of the body and the wings
obtained from the experiments of Fry et al.
(Fry et al., 2003
) are
prescribed in this study. The first step in the simulation is to convert the
experimentally observed kinematics to rigid body translations and rotations in
the computational frame.

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Fig. 5. Angular positions of the wings during the maneuver (from
Fry et al., 2003 ). (A) Stroke
angle, (B) deviation from the stroke plane and (C) the angle of attack.
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Kinematics of the maneuver
Fig. 1 shows the hovering
Drosophila in the experimental coordinate system (XYZ). The
prescribed data consist of the coordinates of the head, the tail, the wing
hinges and tips and the unit vector normal to the wings. In order to convert
this data to rigid body translation and rotation, first the fruit fly body is
placed in the computation coordinate system (xyz) with the head to
tail aligned along the x-axis, the two wings placed flat on the
(y-z) plane, as shown in Fig.
2.
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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Fig. 6. Difference in angular positions between the wings during the maneuver. (A)
Experimental raw data and (B) filtered data.
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Fig. 7. Comparison of time history of (A) lift (L) and (B) thrust (T) forces. Gray
and white bars indicate downstroke and upstroke, respectively.
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The rotation angles for the body are obtained from the bodytail
vector, the right wing hinge to the left wing hinge (RWHLWH) vector and
the vector orthogonal to these two vectors, as shown in
Fig. 3. Examination of the
computed Euler angles show that the yaw angle reaches a maximum of 53°,
while the pitch and roll angles reach a value close to 90° at the end of
the maneuver. Although these are a consistent set of angles, the actual yaw,
pitch and roll angles of the fly body can be obtained by pitch and yaw angles
from the bodytail vector and then obtaining the roll angle from the
RWHLWH vector. This set of angles is shown in
Fig. 4, where the fly is seen
to yaw 90° and the pitch remains between 40° and 50° and the roll
is nearly zero.
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.

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Fig. 9. Time history of yaw moment Myaw. (A) Contributions from right
and left wings, (B) the total moment and (C) the filtered moment.
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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. 10. Time history of forward force produced by the wings. (A) Contributions from
right and left wings, (B) the difference in force and (C) the filtered
difference in force.
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In order to find out what the contribution of each of these force
components is to the yaw moment, the corresponding moment arms have to be
determined from the center of pressure.
Fig. 13 shows that the forward
force has a positive contribution to the yaw moment during the entire maneuver
and the contribution from the side force is positive during the initial phase
of the turning and remains negative during the rest of the maneuver, thus
providing a restoring torque. The remainder of the yaw moment arises from the
lift force acting in the y-direction, resulting in a non-zero
component of moment along the yaw axis.
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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Fig. 15. Distance (D) of the center of pressure from (A) the center of
rotation and (B) the wing hinge.
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Fig. 16. Force vectors (red arrows) on the left and right wings during a hover
cycle, t=0.0156 s to 0.0204 s.
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Fig. 17. Force vectors (red arrows) on the left and right wings at the beginning of
the saccade, t=0.0298 s to 0.0346 s.
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Fig. 18. Force vectors (red arrows) on the left and right wings in the middle of the
saccade, t=0.0538 s to 0.0586 s.
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The moment arm for the sideways force component,
Fig. 14B, shows that the
center of pressure is located between the center of rotation and the tail of
the fruit fly, and is indicated by the negative values of the moment arm.
Further, during hovering the location of the center of pressure nearly
coincides between the right and left wings, hence reducing the contribution
due to the side force.
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.
During the middle of the saccade maneuver, the wing motion shows an
asymmetric banana-shaped profile both during up and downstrokes,
Fig. 18, with the right wing
showing a more pronounced turning at stroke reversal. Also, the stroke angle
for the right wing is considerably larger compared to the left wing. The
magnitudes of the force vector on the left wing for this cycle is slightly
larger than that of the right wing.
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.

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Fig. 22. Vorticity generated by the leading and trailing edges of the wings shown by
iso-vorticity surface during a hover cycle.
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During a cycle in the middle of the maneuver, the asymmetries in the wing
positions and the pressure distributions can be seen in
Fig. 21. At t=0.0591
s and at t=0.0619 s, the lift produced attains a peak during the
downstroke and the upstroke, respectively. At t=0.621 s, a maximum
thrust is produced during the upstroke.
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.
Fig. 24 shows the evolution
of vorticity shed from the wings through one cycle in the middle of the
saccade maneuver. The vortex loops shed from the leading edge and the wing tip
clearly show asymmetry between the right and left wings. The loop ahead of the
body on the right wing is still connected at t=0.05328 s while on the
left side it has already pinched off. We can see the vortex shedding off the
leading edge at stroke reversal, t=0.05403 s, and being made into a
loop through the downstroke. At t=0.550.57 s, we can see the
-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
|
|---|
3-D unsteady computations of a maneuvering fruit fly Drosophila
have been carried out. The kinematics of the wings and the body were obtained
from experimental observations and converted to 6-DOF motion for the body and
the wings. The unsteady force and moments via direct integration of
the pressure on the surfaces were obtained and compared to the experimental
results and show excellent agreement for the entire hover and saccade
maneuver, consisting of over 20 wing beat cycles. The kinematics of the wings
show that subtle changes can result in the yaw moment required to perform the
turning maneuver. During the maneuver, the largest change between the right
and the left wings occurred in the stroke angle. The left wing lagged the
right wing by approximately 13° midway during the saccade, at
approximately t=0.05 s and recovered at the end of the maneuver. The
difference in the angle of attack between the two wings shows that the left
wing was 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 show that the right wing was at a slightly higher elevation
angle during the maneuver. These subtle changes in the kinematics can be
incorporated for controlling unconventional vehicles such as the Biplane
Insectoid Travel Engine (BITE) being developed at the Naval Research
Laboratory.
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
|
|---|
This work was supported by ONR through an NRL 6.1/6.2 project: `Flapping
Propulsion for Unconventional MAVs', with Dr James Kellogg as the project
monitor. The valuable discussions and the information on the experimental
kinematics with Dr Steven Fry of the Institute of Neuroinformatics,
Zürich, Switzerland are greatly appreciated. Also, regular discussions
with Dr Rainald Löhner of George Mason University is greatly appreciated.
This work was supported in part by a grant of HPC time from the DoD HPC
centers, ARL MSRC SGI-O2K and NRL SGI-O2K.
 |
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