AE 242 Biological Flows

Propulsion by Flagellum
Shashank Tomar

31 May 2022

1 Introduction
Unicellular micro-organisms have existed since time immemorial on Earth. They have existed, repro-
duced and survived in extreme conditions. In accordance with Darwin’s Theory of survival of fittest,
they evolved various characteristics and complex body parts for various applications, like finding food
and reproduction[Lig76]. This project studies one such whip-like structure, named flagella found on
some unicellular organisms.
The word flagellum is used both for the filamentous attachments that confer motility on some single
cell organisms and for the fundamentally different whip-like organelles that play an important role
in locomotion and feeding in a vast range of eukaryotic organism. The flagellum is simply a rather
gently curving helical filament of protein (Figure 1).

Figure 1: Flagellar organisms. Left: E Coli, Right: Spermatozoa

Due to the very small dimensions of micro-organisms, the motion is classified under low Reynolds
number low regime (≈ 10−4 ), the regime in which fluid dynamics is governed by Stokes equations.
ρva
Re = (1)
η
How a similar propeller would work at high Reynolds number is difficult to imagine. Imagine a ship
in a sea of molasses with its propeller turning at one revolution per month. This has a Re ≈ 10−3 .

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Due to linearity of stokes equations, a helix that translates along its axis under an external force but
without an external torque will necessarily rotate [Pur97]. Conversely, the same helix that is caused
to rotate due to an external torque will necessarily translate, which is the crux of dynamics of rotating
flagella.

2 Modelling Propulsion
On the heels of Purcell, we too go for a relatively simple model of propulsion via flagellum by assuming
that only two degrees of freedom of the isolated propeller exist, rotation at angular speed ω around
its axis and translation at velocity v along that axis (Figure 2). An external force F and some
external torque N is applied to the propeller by means not yet specified (by the motor in the head
of the organism). At the same time, a force -F and a torque -N are exerted on the propeller by the
surrounding fluid.
The governing equations in the low Reynolds number flow are the Stokes equations:

− ∇p + η∇2 v = 0 (2)

Figure 2: An isolated propeller, subjected to an external force F and an external torque N

The force and torque are assumed to be linearly related to v and ω for simplicity of the model.

F = Av + Bw (3)
N = Cv + Dw (4)

We call the 2 × 2 matrix " #

A B
C D
the propulsion matrix P of this propeller. It is also referred to as the resistance matrix in other
literature. The constants A,B,C, and D are proportional to the fluid viscosity η and depend otherwise

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only on the shape and size of the propeller.
From dimensional analysis, one can infer that they scale with propeller size in this way: if every
dimension of the propeller is increased by the factor l, the new propulsion matrix P’ has elements
A′ = lA, B ′ = l2 B, C ′ = l2 C, and D′ = l3 D. We also assume that the torque and force considered
above are as applied by a thin, stiff, untwistable axial wire, thus ignoring the elasticity/compliance of
a real flagellum. A benefit of defining the Propulsion matrix as above is that effect of adding multiple
propellors can be simply determined by adding the individual matrices, i.e:

P = P1 + P2 (5)

Another additional assumption that arises from the geometry of helical flagellum is B=C. The reason
for this assumption is noticing that forward motion v is caused by rotation ω of the flagellum and
vice-versa. Thus assuming B=C allows us to couple the Force and Torque terms, making the model
more aligned with observed physical phenomena.

We now need to connect the head to the tail, as it is the head (or bacterial cell) that provides
the rotational motion to the tail and also accounts for a majority of the drag. The cell is also mod-
elled by a diagonal Propulsion matrix P0 with A0 and D0 being the only non-zero entries. Here
A0 = 6πηa and D0 = 8πηa3 following Stokes’ relation for a sphere of radius a. The cell rotates

(a) Propulsion matrix of composite propellers (b) Head-tail interaction modelling

Figure 3: Images to illustrate the assumptions used in modelling flagellar motion

continuously in a direction opposite to the propeller rotation, there being no external torques on the
system. We denote Ω as the angular velocity of the cell. Both the flagellum and the cell translate
at the same speed v. The torque on the cell is equal and opposite to the torque on the propeller;
likewise, the force on the cell is equal and opposite to the force on the propeller. Figure 3 is helpful
to understand this configuration. With the chosen sign convention, for a right-handed screw B < 0.
The motor will have to drive the propeller in the sense ω < 0 to propel the system to the right, that
is with v > 0. The external force F acting on the propeller is negative, that is directed toward the

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left. The counter rotation of the cell itself is negative: Ω < 0. With due regard to these signs, Force
and moment balance on the propeller shaft implies:

A0 v = −Av − Bω (6)
D0 ω = −Bv − Dω (7)

The above equations on simplification reveal the following relation between the constants:

B2
D0 + D = (8)
A0 + A
The rotation speed of the motor itself, that is, the speed of the ”rotor” attached to the flagellum
relative to the ”stator” attached to the cell wall, is ω − Ω. Denoting the motor speed Ωm , the
following relation is derived between v and Ωm :
BD0
v=− Ωm (9)
(A0 + A)(D0 + D) − B 2

The Torque N exerted by the motor on the propeller is:

B 2 − D(A0 + A)
N= v (10)
B
The force F exerted by the propeller on the fluid is:

A(D0 + D) − B 2
F = v (11)
D0 + D
Therefore we see that this model can be used to evaluate both the linear propagation
speed v, rotation of the flagellum, rotor and the Torque and Thrust exerted by the
Flagellum.

To compute the propulsive efficiency we compare the power output of the motor, which is N Ωm
with the least power required to move the cell at speed v, i.e. A0 v 2 (as the head accounts for the
majority of drag). Thus the propulsive efficiency ϵp is given by:

A0 v 2 A0 D 0 B 2
ϵp = = (12)
N Ωm (A0 + A)(D0 + D) − B 2

From practical measurements in existing literature, it turns out that B 2 << AD. Similarly D0 /(D0 +
D) is just the ratio ω/|Ωm | which is fairly close to unity for cases of interest. The above equations
can be then simplified to:
B
v=− Ωm (13)
A0 + A
A0 B 2
ϵp = (14)
(A0 + A)2 D

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3 Optimizing the propeller size
Consider a flageller propeller of a known/specified shape. Suppose we are free to scale it up or down
in size by a factor k. Then the resulting propulsion matrix will have elements kAp , k 2 Bp and k 3 Dp
where Ap , Bp and Dp refer to the initial propeller parameters. In this case, the propulsive efficiency
is given by:
A0 Bp2 k
ϵp = (15)
Dp (A0 + kAp )2
Differentiating wrt k, we can find the size of propeller that maximizes ϵp for a cell of given A0 . The
maximum efficiency ϵp,max is attained when k=A0 /Ap and has the value
Bp2
ϵp,max = (16)
4Ap Dp
We see that the maximum propulsive efficiency depends only on the shape of the propeller. The
swimming speed of a cell driven by a maximally efficient flagellum is
−kBp
vmax = Ωm (17)
2Ap
For any given shape of a propeller the elements of the propulsion matrix A,B and D can be determined
by very simple experiments with a model[Pur97]. Thus this model allows us have a simple estimate
of the following quantities:

• Propulsive speed

• Rotation speed of flagellum and bacterium cell

• Propulsive Thrust

• Torque acting on the flagellum

• Propulsive efficiency (optimized)

4 Simulation
In order to verify the accuracy of the model, we compare it against numerical simulations on Ansys
Fluent. We consider an experiment where helix shaped screws are dropped in a highly viscous fluid and
their sinking speed v1 and angular velocity Ω1 is measured. Then the helix is connected to another
helix with the same shape, but opposite orientation (viz right/left handed), creating a ”racemic”
element that doesn’t rotate when dropped in fluid. The propulsion matrix of this ”racemix” composite
will be diagonal with elements 2A and 2D. It’s sinking speed is also measured and denoted by v2 .
This experiment was proposed first by Purcell to measure coefficients A, B and D. The expressions
are:
WD
v1 = (18)
AD − B 2

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 WB
Ω1 = − (19)
AD − B 2
W
v2 = (20)
A
where W is the weight of the screw. We model the first propeller of length 52 mm (as seen in Table
4 ) to check the accuracy against numerical simulation.

Figure 4: Elements of Propulsion matrices and propulsion efficiencies for flagellar models dropped in
silicon oil

4.1 Propeller Specification

• A helical spring-like left-handed test propeller was modelled in CATIA V5 using the specifica-
tions for propeller one in the Table above.

• The propeller has a pitch of 10.4 mm and outer diameter of 9.4554 mm as calculated from the
above data (Figure 5).

• After assuming a cross sectional diameter of 1 mm, and steel as the material, the mass of the
propeller was found to be 8.402e-4 kg.

4.2 ANSYS Geometry

• A Boolean Fluent Geometry was used with fluid domain in an enclosure of 50 × 50 × 100 mm3
volume.

• Here an inlet and outlet is provided (for flow towards negative z), as the spring geometry will
not be translating, but will be modelled to rotate about z - axis as the fluid flows across it
(Figure 6).

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 Figure 5: Propeller geometry modelled in CATIA V5

4.3 Meshing
• A 3D Mesh with 510006 Elements (Limit in the student version : 512000) was employed with
element size of 3.2e-02 m.

• A Face sizing of 3e-04 m and an inflation with first layer thickness 1e-05 m for the propeller
surface was also given (Figure 7).

4.4 Fluent Setup and Input Boundary conditions

• The model has no gravity, is steady and pressure based with a viscous laminar model

• The fluid domain is given density and viscosity 971 kg/m3 and 100 kg/ms of Silicon oil, which
is the same as used by Purcell in his experiment

• Coupled scheme used for solution, with reference values from the inlet conditions

• Hybrid initialization was employed

• For the input boundary conditions v1 , v2 and Ω1 were calculated using eqns 18-20 to be
v1 =6.66007e-4 m/s, v2 =6.5261 e-4 m/s, Ω1 =-0.02804 rad/s

4.5 Results and inference

The velocity contour for the converged Fluent simulations can be seen in Figure 8.

• The expected net Force and Moment in the axial direction on the propeller was calculated and
compared with corresponding ANSYS results.

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 Figure 6: Geometry of the setup modelled in ANSYS

Figure 7: Left: Meshing of the entire domain. Right: Finer meshing near the propeller

• The results differ by more than 60 % (Table 1), which is due to the small size of the fluid
domain (due to the restriction of Ansys Student version) and improper inclusion of the head-
tail interactions in the modelling.

4.6 Validation
Similar to the above simulation, a sphere of 60 mm diameter was simulated for validation. The
expected drag force from Stokes’ law is:

F = 6πη(D/2)v = 0.04853N

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 (a) Velocity magnitude contours (b) Velocity vector contours

Figure 8: Results from the converged Ansys Fluent simulations

Variable Theoretical Model ANSYS

F (N) -0.01437 -0.03499
N (N-m) 0 6.83858e-06

Table 1: Comparison of results obtained in Ansys and from the developed theoretical model

By conducting a Fluent simulation with similar settings as above (Figure 9), the calculated drag is
0.1164 N. Thus the inaccuracy in results above could also be due to inaccuracy in the Numerical
simulation.

Figure 9: Fluent simulation of flow over a sphere at low Reynolds number

5 Conclusion
Propulsion by a helical flagellum was modelled, simulated and then compared. The current theoretical
model can predict the linear and angular velocities of propagation (i.e. the motion of center of mass)
as well as the generated Thrust and Moment. Propulsive efficiency was also derived and the size of
the flagellum for optimal propulsive efficiency was also derived.

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Numerical simulations show that the results match within the same order of magnitude. The match
could be closer with better implementation of the numerical scheme and further work can investigate
this. The model however can model the motion only in a straight line and additional degree of freedom
needs to be added into future models to simulate turning and tumbling motion of such organisms.

References
[Lig76] James Lighthill. Flagellar hydrodynamics. SIAM review, 18(2):161–230, 1976.

[Pur97] Edward M Purcell. The efficiency of propulsion by a rotating flagellum. Proceedings of the
National Academy of Sciences, 94(21):11307–11311, 1997.

6 Appendix - Validation of the Force and Moment models used

In all the calculations above, we assumed that the Force (F) and the Torque (N) varies linearly with
velocity (v) and angular velocity (w). In order to check the validity of these assumptions, we nu-
merically model two flows (Figure 10). One with only translational motion of the helix (ω=0) and
other with only rotation of the helix (v = 0). Then in each case the Force (F) and Moment (N) were
measured as a function of v/ω and plotted.

From Figures 11 and 12 we see that F and N vary linearly with ω and v over the tested ranges.
This implies that the supposition of linear variation of the variables is indeed the right one.

(a) Streamlines for ω=0 case (b) Streamlines for v=0 case

Figure 10: Comparison of streamlines over the Helical flagellum

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 (b) Variation of Moment (M) with velocity (v) when
(a) Variation of Force (F) with velocity (v) when ω=0 ω=0

Figure 11: Comparison of Forces and Moments for the case with ω=0

(a) Variation of Force (F) with angular velocity ω when (b) Variation of Moment (M) with angular velocity (ω)
v=0 when v=0

Figure 12: Comparison of Forces and Moments for the case with v=0

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