First published online August 9, 2007
Journal of Experimental Biology 210, 2767-2780 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.000265
Fish biorobotics: kinematics and hydrodynamics of self-propulsion
George V. Lauder1,*,
Erik J. Anderson2,
James Tangorra3 and
Peter G. A. Madden1
1 Museum of Comparative Zoology, Harvard University, 26 Oxford Street,
Cambridge, MA 02138, USA
2 Department of Engineering, Grove City College, 100 Campus Drive, Grove
City, PA 16127, USA
3 Bioinstrumentation Laboratory, Massachusetts Institute of Technology, 77
Massachusetts Avenue, Cambridge, MA 02139, USA
*
Author for correspondence (e-mail:
glauder{at}oeb.harvard.edu)
Accepted 25 April 2007
 |
Summary
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|---|
As a result of years of research on the comparative biomechanics and
physiology of moving through water, biologists and engineers have made
considerable progress in understanding how animals moving underwater use their
muscles to power movement, in describing body and appendage motion during
propulsion, and in conducting experimental and computational analyses of fluid
movement and attendant forces. But it is clear that substantial future
progress in understanding aquatic propulsion will require new lines of attack.
Recent years have seen the advent of one such new avenue that promises to
greatly broaden the scope of intellectual opportunity available to
researchers: the use of biorobotic models. In this paper we discuss, using
aquatic propulsion in fishes as our focal example, how using robotic models
can lead to new insights in the study of aquatic propulsion. We use two
examples: (1) pectoral fin function, and (2) hydrodynamic interactions between
dorsal and caudal fins. Pectoral fin function is characterized by considerable
deformation of individual fin rays, as well as spanwise (along the length) and
chordwise (across the fin) deformation and area change. The pectoral fin can
generate thrust on both the outstroke and instroke. A robotic model of the
pectoral fin replicates this result, and demonstrates the effect of altering
stroke kinematics on the pattern of force production. The soft dorsal fin of
fishes sheds a distinct vortex wake that dramatically alters incoming flow to
the tail: the dorsal fin and caudal fin act as dual flapping foils in series.
This design can be replicated with a dual-foil flapping robotic device that
demonstrates this phenomenon and allows examination of regions of the flapping
performance space not available to fishes. We show how the robotic flapping
foil device can also be used to better understand the significance of flexible
propulsive surfaces for locomotor performance. Finally we emphasize the
utility of self-propelled robotic devices as a means of understanding how
locomotor forces are generated, and review different conceptual designs for
robotic models of aquatic propulsion.
Key words: fish, swimming, robotics, locomotion, flow visualization, digital particle image velocimetry, kinematics, fin, foil, propulsion
 |
Introduction
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The comparative biomechanics and physiology of moving through water has
long attracted the attention of both biologists and engineers, and recent
decades have witnessed considerable growth in the study of aquatic animal
locomotion. Major results of these efforts include a much more complete
understanding of how animals moving in the water use their muscles to power
movement, detailed descriptions of body and appendage motion during
propulsion, and experimental and computational analyses of fluid movement and
the attendant forces (for reviews, see
Biewener, 2003
;
Fish and Lauder, 2006
;
Lauder, 2006
;
Lauder and Drucker, 2004
;
Shadwick and Lauder, 2006
).
Although a number of areas remain in which currently dominant approaches can
still yield fruitful new insights (including, for example, analyses of
maneuvering locomotion, how animals effect control of multiple locomotor
surfaces to maintain stability, and examination of locomotor repertoires used
by animals in natural flow regimes), it is clear that substantial future
progress in understanding aquatic propulsion will require new lines of
attack.
Fortunately, recent years have seen the development of one such new avenue
that holds considerable promise for testing classical hypotheses, as a source
of new data on aquatic locomotion, and as a novel direction that greatly
broadens the scope of intellectual opportunity available to researchers: the
use of biorobotic models. Robotic models of body and appendage function, with
the attendant ability to program specified motions, the intrinsic abstraction
from detailed morphological features present in individual species, and the
ability to explore a broader parameter space of movement than exists in
nature, allows investigators to explore the biomechanics of aquatic propulsion
in wholly new ways. Recent investigations into aquatic biorobotics include
(Alvarado and Youcef-Toumi,
2005
; Anderson and Chhabra,
2002
; Bandyopadhyay,
2005
; Kato, 2000
;
Liu and Hu, 2006
;
Long et al., 2006a
;
Long et al., 2006b
;
Low, 2006
;
Tangorra et al., 2007b
;
Triantafyllou and Triantafyllou,
1995
). Comparative approaches that examine locomotor function in
different species, while invaluable, are limited by the investigator's
inability to control for the many non-locomotor differences among these
species. Furthermore, it is difficult to alter the natural motions of the body
and appendages in freely swimming animals to examine the effect of novel
movement patterns on locomotor performance. And, robotic models can have their
structure and material properties (such as flexibility) altered at will,
allowing a controlled investigation of the locomotor performance effects of
such changes.
Interest in fish biorobotics is not new, and there is a long history,
dating back to early experimental work using models
(Houssay, 1912
;
Breder, 1926
;
Gray, 1953
). These
investigators constructed mechanical models that allowed them to investigate
power output, undulatory wave formation, and the function of the tail during
fish locomotion (for a review, see
Alexander, 1983
), which greatly
increased our early understanding of how fishes generate propulsive
forces.
Robotic models can be used to special advantage when coupled closely with
experimental studies of freely swimming animals. This allows direct
comparisons to be made between the function of models with various
configurations and the function of biological designs, and ensures reasonable
comparison of the performance spaces of the robotic models and natural
locomotion. In our view, the marriage of robotic models with experimental
analyses of biological locomotion promises to drive the next set of major
advances in our understanding of aquatic propulsion.
In this paper we discuss, using aquatic propulsion in fishes as our focal
example, how such a research program might lead to new insights in the study
of aquatic propulsion. Key questions that can be studied using robotic models
of fish propulsion include the following. How do fins generate thrust? How do
multiple fins arrayed along the body of fishes interact hydrodynamically? How
does flexiblity of the propulsive surface affect the speed and efficiency of
locomotion? And, do fishes use kinematic patterns that generate maximal
thrust, or would different motions, if biologically possible, improve
locomotor performance?
We use two specific examples: (1) pectoral fin function and (2)
hydrodynamic interactions between dorsal and caudal fins. We first present
kinematic and hydrodynamic data from experimental studies of fish pectoral,
dorsal and caudal fins to provide the biological context, and then present
data from a robotic model of the pectoral fin and from a flapping foil robotic
device that models dorsal–caudal fin interactions and allows
investigation of the propulsive properties of flexible foils. We emphasize the
utility of self-propelled robotic devices as a means of understanding
how locomotor forces are generated, and review different conceptual designs
for robotic models of aquatic propulsion.
 |
Fish locomotion: function of paired and median fins in vivo
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Fish generate propulsive forces and control body position through motions
of their body and median and paired fins
(Lauder, 2006
;
Webb, 2006
). Each of the fins
that act as control surfaces during locomotion possess distinct skeletal
supports and associated intrinsic musculature that provide active control of
fin surface movement (Fig. 1).
In addition, ray-finned fishes (but not sharks) have the ability to actively
control the surface conformation of their fins via a unique bilaminar
fin ray design, which allows musculature at the base of the fin to generate
curvature of the fin rays along their length, and thus resist hydrodynamic
loading (Alben et al., 2007
;
Lauder and Madden, 2006
;
Lauder et al., 2006
). The
pectoral fins of fishes are paired structures, while the dorsal, anal and
caudal fins are midline structures (Fig.
1). Although fluid dynamic interactions between the paired
pectoral fins and more posterior median fins are theoretically possible, no
such interactions have been demonstrated experimentally, and we will treat the
two sets of fins separately here. The dorsal and anal median fins generate
flows that interact with the caudal fin
(Drucker and Lauder, 2001
;
Drucker and Lauder, 2005
;
Standen and Lauder, 2007
), and
thus the median fins need to be considered as a group.

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Fig. 1. (A,B) Bluegill sunfish Lepomis macrochirus, hovering in still
water, and (C) snowy grouper Epinephelus niveatus skeleton, showing
the positions of the major fins and their internal skeletal supports. The
pectoral and pelvic fins are paired, while the dorsal, anal and caudal fins
are median (midline) fins. The dorsal and anal fins of ray-finned fishes have
internal skeletal supports (pterygiophores), which support musculature that
moves the fin rays. Fin rays are labeled in yellow for the dorsal and anal
fins. The caudal fin also has a complex series of intrinsic musculature that
allows fishes to actively control tail conformation
(Drucker and Lauder, 2001 ;
Lauder, 1982 ;
Lauder, 1989 ). Metal
supporting elements for the grouper skeleton have been digitally removed for
clarity.
|
|
We first consider the kinematics and hydrodynamics of pectoral fin
propulsion, before considering the function of the dorsal and caudal fins.
 |
Paired fin propulsion: pectoral fins
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The pectoral fins of fishes can undergo considerable deformation during the
fin beat cycle, and the recent availability of megapixel resolution high-speed
video has allowed detailed quantification of fin surface bending and the
motion of individual fin rays (Lauder et
al., 2006
; Standen and Lauder,
2005
). For example, in bluegill sunfish (Lepomis
macrochirus) swimming at a slow speed (0.5 L
s–1, where L is total length) involving only use of
the pectoral fins, the area of the fin changes by approximately 30% as the fin
rays separate, and there is considerable fin twisting as well as chordwise and
spanwise bending (Fig. 2).
During abduction as the pectoral fin moves away from the body, the fin root
assumes a cupped shape (Fig.
2B, posterior view), resulting in two simultaneous leading edges,
while the area expansion and fin curvature is clearly seen on the return
stroke (Fig. 2C).

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Fig. 2. Motion of the pectoral fin in a bluegill sunfish (17 cm total length,
L) during steady locomotion at 0.5 L s–1.
Each row shows frames from simultaneous lateral, ventral and posterior digital
videos (taken at 250 Hz) at three time intervals, 97 ms (A), 142 ms (B) and
174 ms (C), during a single pectoral fin beat. Yellow arrows indicate the
major fin motions (smaller amplitude movements of the fin surface are not
labeled with yellow arrows), the small red and blue arrows show the position
of the upper (dorsal) and lower (ventral) pectoral fin edges respectively, and
the green arrow shows the location of the `dimple' on the dorsal fin margin
that forms as a wave of bending passes out along the fin from base to tip. The
large blue arrows and dot in A show the direction of water flow, which is
perpendicular to the page in the posterior view. Note the considerable
twisting and bending of the fin, and the cupped shape as the upper and lower
fin margins move away from the body at the same time (A: posterior view). The
fin beat begins at time 0 ms. Scale bars, 1.0 cm.
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The cupping of the pectoral fin during the outstroke is also clearly
visible in laser images, which permit particle image velocimetry analyses of
water flow induced by fin motion (Fig.
3) (Drucker and Lauder,
1999
; Drucker and Lauder,
2002
; Lauder and Drucker,
2002
; Lauder and Tytell,
2006
). Analyses of pectoral fin water flow patterns show that on
both the fin outstroke and return stroke, water is accelerated downstream at a
velocity greater than free-stream, indicating that thrust is produced by the
pectoral fin throughout the fin beat cycle
(Fig. 3E,F). This conclusion is
corroborated by computational analyses of pectoral fin function
(Mittal et al., 2006
), which
demonstrated two distinct thrust peaks, one each on the outstroke and the
return stroke of the fin.

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Fig. 3. Hydrodynamic function of the pectoral fin in bluegill sunfish swimming at
0.5 L s–1, as seen in posterior view looking
upstream. A laser-generated sheet of light illuminates a thin slice of water
flow as well as the pectoral fin and body, which casts a shadow to the right.
Laser light penetrates the translucent fin, allowing flow between the fin and
the body to be quantified. Water flow in this figure is out of the page,
toward the reader. Images were obtained from 500 Hz digital video. (A–C)
Particle image velocimetry images showing the movement of the fin illuminated
by the laser light sheet in relation to the body and position of the other
fins. Duration of movement shown=0.48 s from panels A–C. Yellow arrows
show the key fin movements: note the cupped fin shape in A and B. (D–F)
Water flow patterns as a result of pectoral fin movement. This column is from
a different sequence than the frames in the left column. Yellow arrows
indicate water velocities (every other vector is shown), and the background
color scheme is coded so that black color indicates free stream flow velocity
(7 cm s–1), red color flow accelerated by the fin to greater
than free stream velocity, and blue color showing flow slowed below free
stream. Note that the pectoral fin accelerates flow on both the outstroke and
return stroke (red color in E and F).
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 |
Median fin propulsion: the body, dorsal and anal fins
|
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The prominent median fins (dorsal, anal and caudal) of fishes
(Fig. 1) play an important and
active role in locomotor dynamics (Arreola
and Westneat, 1997
; Drucker
and Lauder, 2001
; Hove et al.,
2001
; Jayne et al.,
1996
; Lauder et al.,
2002
; Standen and Lauder,
2005
; Tytell,
2006
). In many spiny-finned fishes the dorsal and anal fins are
composed of an anterior spiny portion and a more flexible posterior region,
often termed the soft dorsal and anal fins
(Fig. 1). The spiny portions of
these fins can be erected and depressed but not moved laterally, while the
soft dorsal and anal fins possess inclinator musculature that powers lateral
movement (Geerlink and Videler,
1974
; Jayne et al.,
1996
). The inclinator muscles are active during both steady
swimming and maneuvering, and movement of the dorsal and anal fins generates a
vortex wake (Fig. 4). Together,
the dorsal and anal fins in bluegill sunfish contribute as much locomotor
force during slow swimming as the tail
(Tytell, 2006
). Perhaps the
most important consequence of the dorsal and anal fin vortex wake is that the
caudal fin moves through water that has a greatly altered flow structure
compared to undisturbed free-stream flow, far away from the swimming fish.
Experimental studies of trout and bluegill sunfish
(Drucker and Lauder, 2001
;
Drucker and Lauder, 2005
;
Standen and Lauder, 2007
) show
that the dorsal and anal fins produce vortices that pass downstream and are
encountered by the caudal fin as it sweeps from side to side
(Fig. 4). These vortices can
potentially enhance thrust produced by the tail if they encounter the tail at
an appropriate phase of movement. Akhtar et al.
(Akhtar et al., 2007
) showed
through a computational fluid dynamic analysis of the sunfish dorsal fin and
tail that the phase relationships reported by Drucker and Lauder
(Drucker and Lauder, 2001
) did
indeed produce enhanced thrust by enhancing the leading edge vortex as the
tail surface is inclined forward.

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Fig. 4. Hydrodynamic analysis of the dorsal fin and caudal fin in swimming bluegill
sunfish, to show that these two fins can act as dual flapping foils in series,
and that flow leaving the dorsal fin can affect caudal fin function. The
caudal fin of ray-finned fishes does not move through undisturbed free stream
flow, but rather has its flow environment highly modified by upstream fins.
The left panels show the laser-imaged dorsal fin and tail of a bluegill
sunfish (16.5 cm L) swimming at 17 cm s–1; laser
light illuminates from top to bottom in these images, and the dorsal fin and
tail cast shadows toward the bottom. Yellow arrows in B show the
left–right oscillatory motion of the dorsal fin and tail as seen from
above. In the right panels these images are analyzed to show water flow
velocities around the fins (vectors were not calculated in the fin shadows)
and vorticity. The views shown in this figure are from above, looking down on
the upper surface of the fish with the dorsal fin and tail (also see
Fig. 1). In A, the dorsal fin
has shed a clockwise vortex that is moving toward the tail. This vortex passes
above the tail (B) while the dorsal fin sheds a new vortex of opposite sign on
the return stroke. This pattern repeats as a clockwise vortex is just leaving
the dorsal fin again (C). Note that flow in the gap between the dorsal fin and
tail is nearly orthogonal to free-stream flow. Free-stream flow has been
subtracted from the right panel images to reveal flow structure; images on the
left have been contrast-enhanced.
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Another significant finding of experimental studies of median fin function
is the strong side force component of the wake. This result can be seen in
Fig. 4, where the dorsal fin
wake generates flows that are nearly orthogonal to the free-stream. The caudal
fin is also capable of generating a thrust wake signature, with a momentum jet
formed at nearly a 45° angle to the direction of travel
(Fig. 4C). However, a thrust
wake is not present in steadily swimming anguilliform fishes, and the
difference among fish wake flow patterns may be related to the
three-dimensional body shape and the presence of a discrete propeller-like
caudal fin (Lauder and Tytell,
2006
).
 |
Fish locomotion: studied with robotic models
|
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The study of fish locomotor physiology and biomechanics is greatly enhanced
by the ability to manipulate movement patterns experimentally in ways not
possible to achieve through comparative analyses of living fishes, and to
directly measure forces produced by fin-like flapping foils.
A variety of different fish robotic designs have been produced, and among
the most well known are the autonomous robotic fish-like devices
(Anderson and Chhabra, 2002
;
Kato, 2000
;
Liu et al., 2005
;
Long et al., 2006b
). But much
of the progress in fish biorobotics has occurred through the use of laboratory
models in still water or flow tanks that allow controlled study of specific
movement patterns and simultaneous force measurement.
Fig. 5 summarizes several
alternative concepts for robotic platforms useful for the study of fish
locomotor mechanics. One common approach is to attach a robotic fish-like
device to a carriage and tow this carriage through a water tank
(Fig. 5A). Under these
conditions, force and torque sensors can be used to quantify the lift and
thrust forces produced by the model when it is towed through the water at
known speeds (e.g. Barrett et al.,
1999
; Triantafyllou and
Triantafyllou, 1995
). This approach has been used in many studies
of flapping foil-based propulsion and to evaluate the effect of locomotion in
the wake of upstream bluff bodies such as a cylinder or rock placed in the
flow (e.g. Beal et al., 2006
;
Hover et al., 2004
;
Techet et al., 1998
).

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Fig. 5. Schematic figure to illustrate two different categories of fish-like
aquatic robot design and the measurements that might be made from each design.
(A) Robot is attached to a sting (a rod holding the robotic model vertically
from the carriage above) and either fixed in place while forces are measured
on the sting, or towed at a fixed velocity on a moving carriage. In either
case, the robot is not self-propelled, but rather moves at externally imposed
speed. In this case, there need be no equality between thrust and drag forces,
as it is not known if the robot is generating sufficient thrust to overcome
drag. (B) Robot swims at a self-propelled time-averaged constant speed as a
result of thrust generated by heave and pitch motions, and mean thrust force
per cycle must equal the mean drag force. The flow speed in the tank is
adjusted to a value, Ueq, where the robot propels itself
at a constant equilibrium X position, termed Xeq.
The robot is free to move itself upstream and downstream on a low friction air
bearing system. Once Xeq is determined for a particular
heave and pitch motion pattern during self-propulsion, the robot can be fixed
in position at Xeq to measure forces and torques while the
same motion pattern and flow speed used for self-propulsion are imposed. This
allows force measurement under conditions identical to self-propulsion, when
thrust and drag forces must be equal.
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Another approach, illustrated in Fig.
5B, is to allow the robotic models to self-propel. Under the
condition of self-propulsion at a steady speed, the thrust generated by the
model must equal the drag force experienced by the robotic device. In the
robotic devices described in this paper, self-propulsion is allowed by
mounting the carriage holding the device motors on extremely low friction air
bearings. These air bearings permit motion in the upstream–downstream
direction but not side-to-side movement of the whole carriage (although heave
motors do move the fin-like foils from side to side). The flapping surface is
immersed into a flow tank, and the speed of the external flow adjusted so that
the flapping robotic device generates sufficient thrust to precisely hold
position at a mean fixed position. When the robotic device holds at an
equilibrium position in the flow tank (termed Xeq), the
flow speed is noted as Ueq
(Fig. 5Bi). Different phase
relationships between heave and pitch movement will give rise to different
Ueq values for which the robotic device holds position at
Xeq.
Once Ueq has been determined for a given set of
movement parameters, the robotic device can be attached to a force transducer
at position Xeq to allow quantification of thrust, lift,
and side forces at this equilibrium position under imposed flow of
Ueq (Fig.
5Bii). By measuring forces at known self-propelled speeds, forces
are measured under conditions at which mean thrust must equal mean
stroke-averaged drag. One issue of practical concern is the force generated by
the cable bundle that is necessary to provide power to the motors and to read
data from the force, torque and position sensors. The effect of the cable
bundle can be minimized by using an Xeq value that
corresponds to the neutral position of the carriage on the air bearings: if
all measurements are made at this neutral Xeq measured
under conditions of no flow, then the cables have no effect on the final
measured Ueq and force values.
 |
A robotic fish pectoral fin
|
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We have constructed a self-propelled robotic pectoral fin that replicates
many of the anatomical features of a real fish pectoral fin
(Fig. 6). This biorobotic
pectoral fin is attached to a carriage that is itself mounted on air bearings
that have very low friction for motion in the upstream–downstream
direction (Fig. 6A). The
pectoral fin robot uses motor-driven nylon tendons to actuate bilaminar fin
rays (each ray is composed of two halves, or hemitrichs)
(Lauder, 2006
) that are of
similar design to real fish fin rays: displacement of one half of the ray
induces a curvature in the ray, and curvature of the whole fin surface can be
controlled by producing curvature in all rays. Like the biological fin ray,
small displacements at the base cause a large displacement at the tip of the
fin ray. The whole fin can be reoriented by moving a compliant base
(Fig. 6B). In the design of the
biorobotic fin we used five bilaminar fin rays embedded in a flexible urethane
webbing with a modulus of elasticity of approximately 0.10 MPa, similar to
that of the (relatively extensible) fin membrane in bluegill sunfish
(Lauder et al., 2006
). The
webbing is pleated so that it can be expanded easily. The complex geometry of
the fin rays and fin ray base plate (Fig.
6C) was manipulated to adjust the fin ray's passive stiffness and
ability to curl when the bases of the hemitrichs were displaced, and they were
constructed using stereo-lithography (which allowed rapid polymerization of
different prototype ray structures). Nylon tendons are attached through small
holes in the base of the hemitrichs.
The compliant base mechanism supports the fin rays and serves as a
deformable joint about which the fin rays are moved. Fin rays are held in
place by tension in the nylon tendons. A stiff base plate
(Fig. 6B,C) is mounted between
the compliant base mechanism and the actuators and contains channels through
which the tendons pass. The compliant base has multiple segments designed to
allow it to bend and generate a range of biologically realistic motions that
would be difficult to achieve with a rigid hinged mechanism
(Fig. 6D–G): expansion
and hence area increase, curling of the surface, and cupping (similar in
character to motions of the sunfish pectoral fin during locomotion, Figs
2,
3).
When the carriage supporting the fin is mounted above the flow tank
(Fig. 6A), the biorobotic fin
in the tank can be imagined to represent the pectoral fin of a fish swimming
on its side just below the water's surface. The design allows the submerged
fin to be reoriented through ±60° pitch and 180° yaw. To
measure thrust force, the fin carriage was mounted on the air bearings in an
equilibrium position and attached to a force transducer (as in
Fig. 5Bii). The force produced
by the biorobotic fin along the X axis (fore and aft) was measured
during fin beats as the fin was cycled between approximately 0°
(horizontal or `against the fish body') and 90° (vertical or `maximum
outstroke'). The fin's movements were created by superimposing combinations of
curl, expansion and cupping onto a basic sweep motion. Simple sinusoids were
used to drive the sweep, curl and expansion motions at about 0.60 Hz. Cupping
was activated using a square wave with the fin cupped during out-stroke and
flat during in-stroke. Since the commanded velocity was the same on the
out-stroke and in-stroke, changes in force were related directly to changes in
the fin's shape and its effect on the water. All tests on the robotic fin
shown in Fig. 6 were conducted
in still water.
Robotic pectoral fin forces produced under three different movement
patterns are shown in Fig. 7.
The most biologically realistic movement pattern is the `cup and sweep', which
is the closest to the pattern observed in sunfish
(Fig. 2). The pectoral fin is
cupped as it sweeps out from the body and flattens as it sweeps back on the
return stroke (Fig. 2A). The
cup and sweep motion generates a biphasic thrust trace, with one thrust peak
during abduction (out-stroke) and a second peak during adduction (in-stroke).
This closely resembles the force pattern calculated using computational fluid
dynamics based on the actual movement of sunfish pectoral fins
(Lauder et al., 2006
;
Mittal et al., 2006
), and the
experimental observation that thrust is produced throughout the fin beat
(Fig. 3)
(Lauder and Madden, 2006
;
Lauder et al., 2006
). There is
a small period of drag for the cup and sweep motion near mid-stroke that
corresponds to the time when the pectoral fin is changing from abduction to
adduction (Fig. 7).
Interestingly, two (smaller) peaks in thrust force are obtained with a cupping
only movement and the small period of net drag force is absent. This suggests
that the observed cupping motion of the bluegill pectoral fin is an important
component of thrust generation. Thrust production during initial cupping may
reflect the dual leading edge vortices observed during cupping in bluegill
sunfish fins (Lauder et al.,
2006
). In contrast, for a sweep motion without cupping, large drag
forces are generated on the outstroke, while thrust is generated on the
return. Little net thrust is produced by the sweep-only motion
(Fig. 7).

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Fig. 7. Force in the X-direction (see
Fig. 5) reflecting thrust and
drag, measured from the robotic pectoral fin during a single fin beat under
three different imposed movement patterns (shown in different colors). Robotic
models of the pectoral fin allow analysis of the effects of different movement
patterns in a way not possible with studies of live animals alone. When the
fin executes a cupping motion only (black trace), a force curve with two
distinct peaks is produced with no drag force at the transition from outstroke
(abduction) to instroke (adduction). A cupping and sweeping motion (blue
trace) generates considerably higher thrust forces as well as a small drag
force during the transition. Moving the fin in sweep only (red trace),
produces large drag forces on the outstroke and roughly equivalent thrust on
the return stroke.
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There are several noteworthy differences between the function of the
robotic pectoral fin and that of sunfish. Most importantly, we observed no
wave of spanwise bending along the robotic fin (e.g.
Fig. 2B), and this may be due
in part to the relatively stiffer fins rays in the robotic model. Bluegill
sunfish fin rays and membrane have an elastic modulus of about 1 GPa and
0.3–1 MPa, respectively (Lauder and
Madden, 2006
), which makes sunfish fins more flexible than the
robotic pectoral fin we constructed. In sunfish, relatively high fin
flexibility allows generation of a chordwise wave, which may contribute to
thrust production during the transition from abduction to adduction, and the
lack of such motion in the robotic model may result in the small period of
negative thrust at midstroke.
 |
A flapping foil robot
|
|---|
The dorsal and anal fins and the caudal fin of fishes can be considered as
representing dual flapping foils arranged in series (Figs
1,
4). The dorsal and anal fins
constitute the upstream foil, leaving a wake encountered by the downstream
caudal fin foil (Fig. 4). The
pattern of median fin function in fishes observed experimentally to date
involves a relatively narrow range of fin movement amplitudes and phasings
(Drucker and Lauder, 2001
;
Drucker and Lauder, 2005
;
Hove et al., 2001
;
Standen and Lauder, 2005
;
Standen and Lauder, 2007
), and
in order to control for interspecific differences and to explore a large
parameter space of heave and pitch motions, it is useful to have a robotic
device that can execute programmed motions. Because the median fins of fishes
are attached to the undulating fish body, only relatively small differences in
phase and amplitude are observed between the dorsal, anal and caudal fins. A
robotic device allows the uncoupling of the motion of the dorsal and anal fins
from that of the caudal fin, and new (biologically impossible) movement
patterns can be executed to examine their effect on foil thrust and on the
wake patterns produced by different foil motions. A robotic flapping foil
device is also useful for testing the effect of foil flexibility on wake flow
patterns, for quantifying hydrodynamic foil–foil interactions, and for
understanding why some movement patterns produce an anguilliform wake, while
others generate the classic carangiform fish wake flow pattern
(Lauder and Tytell, 2006
).
We have constructed a dual-foil flapping robotic device in which two foils
(we used the NACA 0012 airfoil cross-sectional geometry, which approximates
the shape of many aquatic propulsive surfaces)
(Fish and Lauder, 2006
) are
separately mounted on carriages attached to an air bearing system to allow
independent self-propulsion of each foil
(Fig. 8A) with little
frictional loss. Linear encoders attached to each foil carriage allow
measurement of foil position. During self propulsion, we measure the flow tank
speed (Ueq) required for the foil to hold position at the
equilibrium location (Xeq;
Fig. 5B) and determine the
locomotor performance of different heave and pitch amplitudes and frequencies
while simultaneously quantifying the wake flow pattern generated by each foil
using particle image velocimetry. Once Ueq is determined,
we attach the foils to a force transducer at position Xeq
and measure the force generated by each set of movement parameters. These
experiments are repeated for a range of interfoil spacings, ranging from 0.5
to 2.0 chord lengths, where the chord is the width of the foil.

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Fig. 8. Design of the self-propelled dual flapping foil robot to study fish fin
function. (A) Carriage that holds the dual foils, with the heave and pitch
motors for each foil mounted above the flow tank on air bearings that allow
horizontal translation in the X-direction with little friction. This
design feature is critical to allowing self-propulsion. In this image, the two
foils are suspended above the flow tank. (B) Close view of the two foils (NACA
0012 in cross-sectional shape); the foils are 6.85 cm in chord length (width)
and 19 cm high. (C) Close view of the pitch and heave motors for one foil
mounted on the carriage and air bearing system.
|
|
This flapping robotic device also allows us to investigate the locomotor
properties of biomimetic foils with varying degrees of flexiblity. The role of
flexibility in biopropulsion is still not well understood. Both rigid foils
and flexible foils can generate thrust, but fin and body propulsion in most
aquatic systems involves flexible hydrofoils. What do biological systems gain,
if anything, by utilizing flexible propulsive surfaces? By mounting foils with
varying degrees of flexibility on the robotic flapping device, we can
correlate self-propelled speed with flexibility while controlling all other
parameters.
Here we focus first on the presentation of foil wake flow patterns during
movement with 0.5 chord length spacing between the two foils under conditions
of self-propulsion (Fig. 9),
for comparison with the experimental data from bluegill sunfish dorsal and
caudal fins (Fig. 4) and
computational fluid dynamic results (Akhtar
et al., 2007
). The influence of the upstream foil on the flow
encountered by the downstream foil is clearly evident as a stream of trailing
vorticity encounters the downstream foil
(Fig. 9B) as it crosses the
wake of the upstream foil. Furthermore, the reduced flow between the two foils
is evident, as is the enhancement of the leading edge vortex on the downstream
foil (Fig. 9C). This is
precisely the mechanism identified by Akhtar et al. in their computational
study for thrust enhancement when the wake of the upstream foil encounters the
trailing foil and causes increased leading edge suction
(Akhtar et al., 2007
).
The strong thrust wake of the downstream foil is also evident as the foil
reaches the extremes of side-to-side motion
(Fig. 9A,C) with two
counter-rotating regions of vorticity and a large region of water with
relatively high momentum moving back and to the side in a manner similar to
the carangiform wake of swimming fishes (e.g.
Nauen and Lauder, 2002a
;
Nauen and Lauder, 2002b
).
The robotic flapping apparatus also demonstrates that a passively flexible
foil can generate thrust and a wake pattern similar to that seen in swimming
fishes. Fig. 10 shows
self-propulsion at 24 cm s–1 by a flexible plastic foil
actuated only in heave, a motion orthogonal to free stream flow. Thrust is
generated by momentum transfer to the surrounding water as a wave of bending
passes down the deforming plastic foil, and also by attached leading vortices
on the forwardly inclined foil surface that remain attached for most of the
movement cycle (Fig. 10A,C).
The wake generated by the flexible plastic foil resembles that of anguilliform
swimming in eels (Tytell and Lauder,
2004
), with large laterally directed momentum jets and elongate
shear layers that break up into multiple centers of vorticity
(Fig. 10C).

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Fig. 10. Hydrodynamics of a single, flexible, flapping foil self-propelling at a
speed of 24 cm s–1. The white arrow shows the heave motion
(3.5 cm heave amplitude) of the rod that actuates the flexible foil, composed
of a plastic sheet of the same dimensions as the foils in
Fig. 9. Foil thickness is 0.32
mm, foil length=19 cm, foil height=6.8 cm, and the video sample rate is 250
Hz. The left panels show the flexible foil and water illuminated by a laser
light sheet from top to bottom; the flexible foil casts a shadow toward the
bottom of each image; these images have been contrast-enhanced. Large yellow
arrows in the left-hand panels show the direction of foil surface motion from
one panel to the next. The actuating rod to which the foil is attached and the
thin black foil itself have been enhanced by a white dot and line,
respectively, for clarity. In the right panels these images are analyzed to
quantify water flow velocities and vorticity around the flexible foil (vectors
could not be calculated in the fin shadows), as in the previous analysis of
two foil self-propulsion (Fig.
9). Note that an attached leading edge vortex (LEV) is visible at
0 ms as the foil leading edge nears the end of its downward motion and begins
to move up. This attached LEV persists throughout the duration of the
downstroke, until almost 930 ms (not shown). A distinct thrust wake is evident
behind the flexible foil, with a strong side component.
|
|
Even using simple flexible foils and actuation only in heave, complex wake
flows can be generated that strongly resemble fish locomotor wake patterns.
Furthermore, the comparison of rigid and flexible foils (Figs
9 and
10) suggests that even simple
models of flexible deforming surfaces can generate complex wake patterns with
biological relevance.
 |
Prospectus
|
|---|
The construction of robotic models that capture key features of the
functional design of fishes promises to provide a significant new avenue for
the exploration of longstanding questions in aquatic locomotion. Fish
biorobotics, coupled with the ability to study the hydrodynamics of locomotion
in vivo, provides a powerful intellectual combination for pursuing
key questions in aquatic propulsion.
The generality of the approach discussed here also indicates that many
broader issues relating not just to fish propulsion but also to the design of
aquatic organisms in general and underwater vehicle design using biologically
inspired features, can be examined using biorobotic models
(Bandyopadhyay, 2004
;
Fish, 2004
;
Long et al., 2006b
;
Miklosovic et al., 2004
;
Tangorra et al., 2007a
;
Tangorra et al., 2007b
). The
question of how flapping foils in series interact is a general one, with
implications for flapping propulsion in both water and air, and the propulsive
significance of flexibility, a hallmark of biological systems, is still not
well understood. What do organisms gain in performance, if anything, by having
flexible propulsors? New directions and questions require new approaches, and
integrating the study of biorobotic models with experimental analyses of
animals moving in vivo promises to be one such new avenue.
 |
Acknowledgments
|
|---|
This work was supported by an ONR-MURI Grant N00014-03-1-0897 on fish
pectoral fin function, monitored by Dr Thomas McKenna and initiated by Dr
Promode Bandyopadhyay, and by NSF grant IBN0316675 to G.V.L. We thank Drs
Rajat Mittal and Promode Bandyopadhyay for many helpful discussions on
bio-inspired propulsion, Eric Tytell for his help in designing the dual
flapping foil robot, Naomi Davidson for her many contributions to the pectoral
fin robot, and Eliot Drucker for collaborative work on sunfish fin function.
Two referees provided helpful suggestions on the manuscript. Karsten Hartel
kindly provided the photograph in Fig.
1C. Tony Julius, Mary Hong and Julie Idlet provided invaluable
assistance in the lab.
 |
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