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Bending, twisting and beating trunk robot bioinspired from the 3+0 axoneme

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2013 Bioinspir. Biomim. 8 026006
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IOP PUBLISHING

BIOINSPIRATION & BIOMIMETICS

doi:10.1088/1748-3182/8/2/026006

Bioinspir. Biomim. 8 (2013) 026006 (12pp)

Bending, twisting and beating trunk robot

bioinspired from the 3 + 0 axoneme
Christian Cibert
Institut des Sciences et Techniques des Yvelines - ISTY, 10-12, Avenue de l Europe F-78140 Velizy,
France
E-mail: cibert.christian@wanadoo.fr

Received 19 October 2012

Accepted for publication 12 March 2013
Published 12 April 2013
Online at stacks.iop.org/BB/8/026006
Abstract
The axoneme is the skeleton and motor axis of flagella and cilia in eukaryotic organisms.
Basically it consists of a series of longitudinal fibers (outer doublets of microtubules) that
design a cylinder and whose sliding, due to the coordinated activities of dedicated molecular
motors (the dynein arms), is converted into a bending because outer doublets pairs are
stabilized by elastic links (the nexine molecules). In spite of these interesting mechanical
properties, mechanical and robotics engineers have never considered this amazing molecular
machinery as a model. The aim of this paper is to propose the robotic design and the kinematic
modeling of the 3 + 0 axoneme that makes motile the flagellum of Diplauxis hatti, the
simplest that exists. The model that we propose bends and twists and combines the two
movements. It is able to propagate wave trains that could be involved in the development of
biomimetic actuators of various mechanisms such as (sub)aquatic robotic propellers as well as
robotic trunks.
S Online supplementary data available from stacks.iop.org/BB/8/026006/mmedia
(Some figures may appear in colour only in the online journal)

1. Introduction

Lindemann 1994b, Nicastro et al 2011). It is a very long

structure (from 1 to 100 m) as compared to its diameter
(0.250 m)the ratio length/diameter ranges from 4 to 400
and is able to propel large cells (spermatozoa, protists, etc)
inside very viscous swimming mediums because of their
very low Reynolds number (Brokaw 1966, Gueron and LevitGurevich 1998, Gueron et al 1997, Holwill et al 1995, Yagi
et al 2005).
Different sorts of axonemes exist according to the outer
doublet numbers, their lengths, their accessory structures
(e.g. human spermatozoon (Fawcett 1975, Serres et al 1983),
trypanosoma (Gadelha et al 2006), eel (Woolley 1997, 1998),
and so on) and the shape of the wave train that they
generate.
Within the axoneme, the interval that exists between two
neighboring outer fibers is roughly constant because of the
presence of the two longitudinal and periodic series of dynein
arms, although it could be adjusted to allow the regulation of
the molecular motors (Cibert et al 2010, Lindemann 1994a).

Many nanomachines are bioinspired. The well-known example

is the spinning flagellum of bacteria (Erhardt et al 2010,
Ummat et al 2006, Yang et al 2009), but all the flagella (and
cilia) are not constituted by a passive spiral filament actuated
by only one engine inserted inside the cell membrane. The
axoneme is the axial actuator and the skeleton of the cilia and
flagella of eukaryotic cellsa very conserved machinery in
animals and plantsthat makes these organelles able to beat
with a frequency that ranges from few to hundred hertz.
The axoneme is constituted by a cylinder of parallel
longitudinal outer fibers (polarized microtubule doublets)
whose slidingdue to molecular motor series (dyneins)
(Roberts et al 2009, 2012)is converted into bending because
of the presence between them of the oriented molecular motors
themselves and elastic molecules (nexin links) (Barber et al
2011, Bui et al 2008, 2009, Cibert et al 2010, Dymek et al
2011, Gibbons 1981, Holwill and Satir 1994, Lin et al 2012b,
1748-3182/13/026006+12$33.00

2013 IOP Publishing Ltd

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Bioinspir. Biomim. 8 (2013) 026006

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(B)
(A)

(C)

(D )

Figure 1. (A) The 9 + 2 axoneme, viewed base to tip, is constituted by a polarized cylinder of nine outer doublets of microtubules that
carry: the dynein arms (molecular motors), nexin links (elastic junctions) and the radial spokes (Cibert and Heck 2004, Lin et al 2012a). All
these appendages are periodically distributed along each of the outer doublets (B). The central apparatus is organized around two
microtubules and carries different periodic structures involved in the regulation of the axonemal machinery. (C) Organization of the 3 + 0
axoneme. Symbols are the same as in (A). Dashed arrows define the polarity of the axonemal triangle that is defined as a function of the
orientation of the dynein arms and the architecture of the outer doublets. (D) The sliding amplitude is defined as the interval that exists
between the ends of the two outer doublets of microtubules (whose lengths are equal) which constitute a bent pair.

Schematically, one of the two neighboring fibers is the cargo

carried by the two parallel dynein arm series and each of
the dynein arms walks on the surface of the other fiber in a
coordinated manner with the other arms. This mechanism has
been observed in real time (Brokaw 1989).
The local regulation of the activities of the molecular
motors distributed along the outer doublet pairs could be either
geometric (Cibert et al 2010, Lindemann 1994a) or conceive
in a more theoretical context; using bundle dynamics it was
demonstrated that solitary waves propagate along the axoneme
which is able to generate self-organized phenomenon (Camalet
et al 1999, Hilfinger and Julicher 2008, Hilfinger et al 2009).
However, this question remains open.
Whereas snake and fish models are often used to generate
crawling or swimming robots (Khan et al 2010, Kim et al
2012, Sato et al 2012), the axoneme remains unexploited by
roboticists, because the basic biological model of actuators
is the skeletal muscle, and many models try to mimic these
organs that pulls on two opposite ligaments (e.g. McKellins
model (Daerden and Fefeber 2002), electron muscles (Odhner
et al 2006), and so on).
Whatever the sophistication of these bioinspired
mechanical systems they are the intuitive translation of the

apparent macroscopic muscle properties, in spite of the

questions raised by the actual biochemical and physiologic
micro-control of the sarcomere, the active unit of the muscle
cells (Gordon et al 2000, Herzog et al 2010, Jarosch
2008, Smith and Stephenson 2009). These macroscopic
apparent properties correspond to the sum of the elementary
passive extension and active contraction of a few millions
of sarcomeres that depend on the sliding of the contractile
filaments: actin and myosin. However, it is not necessary to
fully know the biochemical/physiological functioning of the
sarcomeres to correctly mimic the apparent properties of the
ensemble that they constitute and to construct a humanoid
machine (Gravato Marques et al 2010).
The mechanical transposition of the axoneme in
macroscopic terms constitutes a more interesting challenge.
Actually (i) no macroscopic axoneme exists because this
organelle is only defined at the cellular scale; (ii) there is only
one axoneme inside a given cilium (or flagellum) and (iii) it
is made of macromolecules that do not have any macroscopic
counterparts.
The aim of the present study is to propose a mechanical
modeling of the axoneme into a dynamics software (ADAMS
software), taking into consideration the basic biochemical
mechanisms of the simplest 3+0 axoneme.
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(A)

(B)

Figure 2. Sliding and bending of segment of a 9 + 2 axoneme. We

have modeled (under ImageJ (rsbweb.nih.gov/ij/download.html)) a
canonic axonemal cylinder constituted by nine longitudinal fibers
linked by transverse series of molecular motors and elastic links, the
transverse links located between two neighboring fibers. The
orientation of the axoneme is the same as that described in
figure 1(A).
Figure 3. Sliding and bending of a segment of a 3 + 0 planar
axoneme. (A) The top view of the model. The inset shows the
orientation of the bending plane when the axoneme is viewed from
the tip. The orientation of the model is the same as that shown in
figure 1(C). The red arrows are the oriented transverse links that
correspond to the dynein arms and the nexin links. Because the
transverse links are always perpendicular to the longitudinal fiber on
which they are permanently bound, they are involved in either
pinching or divergence. (B). Perspective view of the same bent
model. The sliding amplitude corresponds to the extrusion of the
fiber #1 from the axonemal triangle. Because the longitudinal fibers
#2 and #3 are included in a plane that is perpendicular to the
bending plane, the pair [2,3] does not slide; the two pairs [1,2] and
[3,1] are active.

2. Mechanical properties of the axoneme

The basic properties of the axoneme were mainly studied
using two biological models: the flagellum of the sea urchin
spermatozoon and the flagella of Chlamydomonas. The 9 + 2
axoneme is a cylinder made of nine longitudinal outer fibers
(nine outer doublets of microtubules) that surround two
microtubules that organize the central apparatus, hence its
name (figure 1(A) and (B)).
The nine outer fibers are embedded inside the peripheral
cytoplasm of the cells and are free at the distal end of the
organelle. The outer doublets carry three main longitudinal and
periodic series of appendages: the molecular motors (dynein
arms), elastic links (nexin) and radial spokes that link them
to the central apparatus. From a mechanical point of view,
the sliding of the outer doublet pairs (figure 1(D)), due to the
molecular motors series, is converted into bending because of
the action of the elastic links series (Brokaw 1989, Gibbons
1981, Lindemann 1994a, 1994b, 1996).
The three main mechanical characteristics of the axoneme
are: (i) the coupling between the sliding of the outer doublet
pairs and bending, (ii) summation of the elementary sliding
along each of the outer doublet pairs and (iii) the polarity of
the molecular motors functioning; the motors associated with
a given outer doublet walk mainly baseward on the surface of
the neighboring outer doublet.

Because curvature is either positive or negative and

because of the polarity of the dynein arms (Gao 2006), the
bending plane divides the axoneme into two hemi-axonemes
whose cyclic activities are necessarily opposite during a
beating cycle (Cibert 2002). From a mechanical point of
view, the axonemal architecture appears as hyper-redundant
(Wingert et al 2002), it means: robust, lightweight and easy to
control.
The sliding mechanism of the outer doublets that occurs
within the 9+2 axoneme is described in figures 1(D) and 2.
The deviated bending of the outer doublets of microtubules,
which is due to the adhesion of two microtubules (Cibert and
Heck 2004, Cibert et al 2010), was not taken into consideration
in this calculation.
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Figure 4. Wave train and sliding. The calculation of the curves was
done using a C-program written under the C-interpreter of ImageJ.
Definitions of the curves are indicated on the plot directly. Arbitrary
units were used on both axes. See movie 1 available at
stacks.iop.org/BB/8/026006/mmedia.

(A)

Figure 6. Mechanical architecture of the 2D model is capable of

planar bending. The upper and lower diagrams are the ADAMS
model and the kinematic scheme of this construction, respectively.
The bodies are labeled with plain Arabic numbers; the numbers of
the bodies they connect designs the links; they are written in italic
and placed between parentheses. Prime is relative to the rank of the
upper planar 2D module. One of the two joints (22 -2) and (2 -21 )
must be a universal joint instead of a ball joint as designed, to endow
the system with the convenient number of degree of freedom;
ball-joints were used in this scheme because they easier to design.
The transverse bars link the middles of the two segments of the
longitudinal fibers; they are embedded in the left fiber and slide
freely along the right one, when the left and right sides are
defined by the polarity of the scheme only. The actuated link is the
(21 -22 ) translational joint.

(B)

Figure 5. Planar bending of a 2D model (Cibert 2001, Cibert and

Ludu 2010). (A) Configuration of a rigid transverse link that joins
two longitudinal fibers. Gliding is the displacement of the active end
of the transverse link along the element of the convenient segment
of the longitudinal fiber. (B) Configuration of a transverse link
constituted by two articulated segments.

In this scheme, the ensemble constituted by the dynein

arms and nexin links correspond to the periodic transverse
beams that link two neighboring longitudinal fibers, the outer
doublets. When the molecular motors walking on the surface of
the neighboring outer doublet is clearly demonstrated, that of
the nexin links was proposed from experimental observations
and modeling (Cibert 2001, Yagi and Kamiya 1995). Radial
spokes and central apparatus are omitted in spite of their crucial

importance of their regulative functions (Barber et al 2011,

Cibert 2003, Dymek et al 2011, Huang et al 1982, Lin et al
2012b, Mitchell and Nakatsugawa 2004).
Because the outer doublets of microtubules are
continuous, inextensible and incompressibleas compared
4

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according to Gray (Gray 1955, 1958, Gray and Hancock 1955)

as the products of an exponential envelope and a periodic
function (Cibert 2008) (figure 4).
The fish-shaped envelop of the curves that represent
the cumulative sliding along the axoneme characterizes the
curvilinear abscissas where the local sliding of the outer
doublets pairs tends to extrema. The abscissas where the
sliding magnitude is the lowest are the proximal ends of the
axonemewhere sliding and bending are nil by construction,
even if this point is controversial at least in mammal systems
(Vernon and Woolley 2002)and a point whose longitudinal
abscissa depends on the shape of the wave train that propagates
along the axoneme. At this abscissa, in spite of the low sliding
amplitude of the nine outer doublets, the axoneme is able to
bend normally as it does along the other segments where the
sliding magnitude is the largest (Cibert 2008).
Because the bending of the 9+2 axoneme generates
inner forces which are difficult to take into consideration
in a mechanical modeling (Cibert et al 2010, Lindemann
1994a, 1994b, Mitchell 2003), we have modeled the simplest
biological 3+0 axoneme whose mechanism is homologous
to that of the 9+2 axoneme. This simplest axoneme is
constituted by three longitudinal outer fibers and contains
neither central apparatus nor radial spokes (Prensier et al
1980) (figure 1(C); each of the three sides of this triangular
axoneme is homologous to that of the nonagon. Figure 3 shows
the sliding to bending conversion of this simple model.
In this figure, pinching and divergence of the transverse
links (red arrows) are very uniform along each side of in
such way that their spatial frequencies are constant along
each pair of fibers. This is due to the orientations of the
links which are perpendicular to the fiber on which they are
linked. In spite of these small variations, the sliding amplitude
is very large; it corresponds to the extrusion of the fiber #1
outside the axonemal triangle. The same event is observed
in figure 2. This is the consequence of the summation of
the elementary displacement of each of the transverse links
that induces a velocity gradient in the gliding of the active
ends the transverse links. In the biological axoneme, the
consequence of these processes is the clustering of the dyneins
arms (Cibert 2008, Cibert and Ludu 2010). These figures show
that to translate in mechanical terms the sliding to bending
conversion, it is enough to adjust conveniently the intervals
that exist between the transverse links whose active ends glide
along the longitudinal fiber.
Using ADAMS software, we have constructed a 2D
model to test the main mechanical characteristics of one fiber
pair before constructing a 3D model capable of bending and
twisting, and to mix bending and twisting as the biological
model does.

Figure 7. Planar bending of the 2D model.

Figure 8. Modification of the 2D model.

with the sliding amplitude (Kasas et al 2004a, 2004b, Kis

et al 2002)sliding is cumulative (Brokaw 1993, 1996, Cibert
2001, 2002, 2003, Goldstein 1976). Consequently, along a
given outer doublet pair, at a given curvilinear abscissa, the
sliding equals the sum of the activities of the molecular motors
located upstream. This mechanism induces the extrusion of
the outer fibers from the axonemal cylinder at the end of the
curved models shown in figures 2 and 3. Sliding amplitude
(s) between two neighboring outer fibers and curvature (c) are
linked (Cibert 2008):
s = c h,

(1)

where h is the projection on the bending plane of the interval

between the two outer doublets.
Since sliding is cumulative, there is no direct relation
between the local activity of the molecular motors along a
given segment and the curvature of this segment. To illustrate
this important point, we have calculated the wave train that
propagates along the flagellum of a sea urchin spermatozoon

3. The 2D model
To be as close as possible to the biological 3 + 0 axoneme,
the first goal was to construct the 3 + 0 axonemal model
using three flexible fibers (not shown). It is difficult, however,
to adjust the size of compliant fibers as a function of their load
and the controlled movement they must support. Therefore, we
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(A)

(B)

Figure 9. Mechanical adjustments that make enable the module described in figure 8 to generate 3D movements. These modifications are:
(i) longitudinal segments are linked by ball joints; (ii) transverse segments split into three elements in place of two, the joints (1i2i) being
cylindrical and (iii) the joints (32i+13i) and (31i32i) are a universal joint and a ball joint (or vice versa), respectively. (A) The proximal
module anchored on a solid basis; (B) one of the modules along the series.

have decided to model these fibers as series of solid segments

joined by ball joints. Transverse links (molecular motors and
elastic links (nexin)) that stabilize a pair of longitudinal fibers
were rigid too.
In figure 5(A), the system is constructed by two
longitudinal fibers #1, 4 and #3, 6 linked by a transverse beam
#2, 5 that mimics the ensemble constituted by the outer and
inner dynein arms and a nexin link.
When the model bends, the moving end of the transverse
link #5 (arrow head) glides baseward or tipward along the
segment #6, inducing the sliding (figure 1(C)) of the facing
longitudinal segments #4 and 6. This principle is that we have
used to model the bending of the 9 + 2 model shown in
figure 2. This construction is mechanically complex because
the active head of the transverse beam must glide and pivot at
the same time. In figure 5(B), pivoting and gliding functions
are separated. This configuration is easier to build and does
not change significantly the bending angle. Figure 6 shows the
model that we have constructed under the ADAMS software
according to these principles and whose bending is shown in
figure 7.
The three panels in figure 7 demonstrate that the
same increment (or decrement) of the interval between
two successive transverse leads to a cumulative the gliding
amplitude of the head of the transverse links along the
longitudinal fibers, and induces the expected bending of the
model.
Within the 2D model, when the gliding is directed
tipward, the end of the very distal transverse link overlaps
the distal end of the segment along which it moves (figure 7,
left panel). However, when the gliding is directed baseward
the gliding end of the transverse segment might overlap the
proximal ball joint that links two successive segments of the

Figure 10. Base of the 3D MODEL (ADAMS).

longitudinal fiber; in this case, the distal longitudinal segment

of the longitudinal fiber is at odds (figure 7, right panel).
To address this problem, we moved the transverse link at
the distal end of each of the segments of the longitudinal outer
fibers and constructed cantilevers along which the gliding ends
of the transverse links move freely (figure 8).
It is noteworthy that the length of this cantilever defines
the amplitude of the cumulative sliding and consequently the
bending angle of the model.

4. The 3+0 3D model

The main modifications that convert the 2D model into the
3D one are described in figure 9. They consist of adding
degrees of freedom to enable the 2D module to bend in a
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(A)

(B)

Figure 11. (A) ADAMS view of a series of three 3D modules (figure 9) and of the mechanism (tang) that allows the recovery of the
displacement of the free end of the model. (B) ADAMS view of the tang.

direction perpendicular to the plane it forms and to twist about

its axis.
Actually, along the transverse links, cylindrical joints
(1i2i) are added, and the revolute joints that connect two
successive segments of the longitudinal outer fibers in the
2D model are converted into ball joints.
Figure 10 shows the ADAMS models of the base of the
3D model. The three sides of a given stage define a module;
the 3 + 0 model involves three modules (figure 11).
Figure 12(A) and table 1 describe the degrees of freedom
of this model; the nine motors, it involved, control completely
its 9 degrees of freedom.
As stated above and shown in figures 1(C) and 2, when the
model bends, the free distal ends of the longitudinal fibers are
included in a plane whose orientation is not that of the bending.
This occurs whatever the number of outer longitudinal fibers.
To bypass this problem, we have constructed a tang that can
transfer bending and twisting observed at the very end of the
model to any object (figure 11).

Table 1. Mobility (m) of the model described in (figure 12(A). When

N is the number of independent cycles of the system, Ec = N 6
because of the 3D movements of the model. In niki, ni and ki are
the number of links and their degree(s) of freedom, respectively.
Mobility equals 9 and equals the number of motors involved in the
three stacked modules. Calculation was done according to Grubler
(Gogu 2005, Grubler 1917).
Kinematic
equations (Ec)
Bodies = 46
Links = 63
N = 63 46 + 1 = 18
Ec = 18 6 = 108

Independent
unknowns (Ic)

Ic = ni ki
Ic = 117

Mobility (m)
m = IcEc
m = 117108 = 9

Figure 12 and table 2 show that the addition of this

mechanism, which is encaged within the last two modules,
does not change the mobility of the system. To document this
conclusion, we simplify the kinematic diagram of the end of
the model (figure 13).
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Table 2. Mobility (m) of the model described in figure 12(A) and

(B). Mobility is the same as that of the 3D modules without the
tang. See table 1.
Kinematic
equations (Ec)

Independent
unknowns (Ic)
Mobility (m)

Bodies = 46 + 10 = 56 Ic = ni ki
m = IcEc
Links = 63 + 15 = 78 Ic = 117 + 30 = 147 m = 147 138 = 9
N = 78 56 + 1 = 23
Ec = 23 6 = 138
Table 3. Mobility of the distal end of the 3D module series, it
means that of the last two modules that include the tang (figures 11
and 15). The calculation is done according to figure 13(C). The
model includes ten real bodies and one theoretical one. The
number of links equals 13the degree of freedom associated with
each of theoretical links equals 3, the highest degree of freedom
(rotations) allowed by the set of gray bodies defined in figure 13(A).
The motility of the distal end of the module series equals 9.
Kinematic
equations (Ec)
Bodies = 11
Links = 13
N = 13 11 + 1 = 3
Ec = 3 6 = 18

Independent
unknowns (Ic)

Ic = ni ki
Ic = 27

Mobility (m)
m = IcEc
m = 27 18 = 9

(A)

As shown in table 3 the mobility of this mechanism equals

9, and the nine actuators involved in the three modules series
fully control the system.
The 3 + 0 model that we have constructed
is able to bend (figure 14, movie 2 available at
stacks.iop.org/BB/8/026006/mmedia), twist (figure 15,
movie 3) and to mix bend and twist (movie 4). The model is
able to generate wave train (movie 5).
Using ADAMS we have tested the effects of forces and
torques applied at the end of the tang, and observed that they
do not have any effect on the shape of the model (not shown).

(B)

5. Discussion
This work demonstrates that the 3 + 0 axoneme is one
of the biological systems that could be used to construct a
bending, twisting and beating trunk robot or propelling system.
The model we propose here is intended for use as a spine
in a humanoid robot (ROMEO, ALDEBARAN-ROBOTIC
(France)).
Because of the triangular architecture of the model, the
active changes of the positions of the gliding ends of the
transverse links along one side of the module series move
necessarily the gliding ends of the other transverse links along
the two other sides; the actuators movements are converted
into rotations while they are basically parallel to the model
axis. To generate the same movement, the six actuators of
the Stewart platform (parallel mechanism with 6 degrees of
freedom) form necessarily an angle with the axis of the
platform (Liang et al 2010, Stewart 1965). As compared to
the pioneer Stewart platform, the actuators of the axonemal
model do not support the constraints applied on the tang,
because these forces are manly transferred on the longitudinal

Figure 12. (A) Kinematic diagram of a series of three 3D modules

as shown in figure 11(A). This circular scheme describes the
mobility of a model constituted by the superposition of three 3D
modules. Bodies are represented as discs; links are the lines. Bodies
of the second and third modules are tagged with a ( ) and ( ),
respectively. 0 represent the solid base on which the model is
anchored. The numbers associated with the links are their degrees of
freedom. The first 3D module (the central crown) is described as
followsthe elements of the second and third modules (crowns) are
simply incremented. The longitudinal fibers of this 3D module are
numbered: 1, 4 and 7. The three elements of the three transverse
segments are numbered: [1,2,3], [4,5,6] and [7,8,9]. As shown in
figure 9, the first element of each transverse segment is embedded at
the end of the corresponding segment of the longitudinal fiber. The
third elements of these three triplets of bodies glide freely on the
fibers (bodies) #4, #7 and #1, respectively. The motors of the three
sides of the 3Dmodel are the links (3132), (6162) and (9192).
(B) Kinematic diagram of the tang encaged in the last two 3D
modules of the 3D model (figure 11(A) and (B). The dashed line
represents the last two segments of the longitudinal fibers. The
central circle is the tang.

Bioinspir. Biomim. 8 (2013) 026006

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(A)

(B)

(C )

Figure 13. Simplified kinematic diagram of the tang. (A) Description of the architecture of the system dedicated to gather the deformations
of the model at the distal end of the 3D model (figure 11), which is encaged in the distal end of the 3D model. Transverse links and the
motors are omitted from this scheme, because they induce the sliding of the fiber which is compensated by the architecture of the system
that we propose. The gray beams constitute an ensemble, which is linked to the tang by a ball joint. (B) These gray beams should be
removed from the scheme and replaced by a theoretical body and three theoretical links (double dashed arrows) as shown in (C). DOF:
degree(s) of freedom. In (A) and (B), numbers included in the discs are the degrees of freedom allowed by the joints, and bodies (beams) are
represented as lines. In the resulting kinematic diagram (C) bodies are the discs (figure 12). (a)(c) are the links involved in the construction
of the mechanism. The black disc is the tang.

fibers and consequently along the axes of the transverse links.

Nothing excludes however, the possibility to make motile the
tang axis to allow axial translation, although the goal of the
axonemal machinery modeling is not to mimic the clever
platform.
Key words such as fish, jellyfish, robot, drone, swimming,
provide access through different servers to few millions of
images and videos and demonstrate the great interest of new
propulsion systems able to move these nano-, micro-, miniand macro-swimming machines whose application fields are
very different in both industry and research. Nevertheless,
on the side of their sizes, their main differences lie in their
intelligence more than their mechanics, which are founded
on classic fish (or snake) model associated to deflection
mechanisms (swimming appendages), which are analogous

to the fishs fins. It is noteworthy that a jellyfish model1 moves

because of the use of a crown of eight elastic undulating
appendages whose extremities are in the form of fish
caudal fin.
Propeller propulsion involves a rapid engine and generates
necessarily cavitation noise because of the velocity of the
fluid flow on the fan surfaces (Rhee et al 2005). By contrast,
waving systems can be slow being efficient and cannot generate
consequently large cavitation noise. Their only sound signature
is due to their mechanical defaults and their engines. In the
case of the axonemal system that we propose, the amplitudes
of the engines displacements are very low because of their
summation along the model. It may be proposed that the
activities of the engine series could be preset as a function of
1

www.kk.org/thetechnium/archives/2008/05/loving_robotic.php.

Bioinspir. Biomim. 8 (2013) 026006

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(A)

(A)

(B )

(B )

Figure 15. Left-hand (A) and right-hand (B) helices due to the twist
of the 3D AIM. Twisting is obtained when all the motors promote
movements of the same polarity and same amplitude.

Acknowledgments
I thank Vincent Hugel and Pierre Blazevic for helpful
discussions. Grants were supplied by the ROMEO FUI
program (ALDEBARAN, Paris (France)) and personal funds
of the author, specifically for bibliography that is not available
via UVSQ.

Figure 14. Left-handed (A) and right-handed (B) planar bending of

the 3D model. The two bending directions are arbitrarily defined.
Bending is calculated as follows: the orientation of the bending
plane is defined, and the magnitude of extension (or compression) of
the motors, which are distributed along a given side of the 3D
model, is calculated as a function of the algebraic module of the
projections (figure 1(B)) of the homologous transverse links on the
bending plane.

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