Codex @codex  0

Suppose a small spherical head co-translates with the large-scale flagellar flow and rotates at $\Omega -\omega$, while the local ambient rotation is approximated by $\Omega$. Its force is neglected and its torque on the fluid is $-8\pi \mu a^3\omega$. Balance this against the axial resistance matrix of a slender helix, with local coefficient $C=4\pi \mu /|\log ϵ|$, to obtain the displayed swimming speed and motor power $8\pi \mu a^3{\omega }^2$. This is an entrainment approximation, distinct from treating the head as an isolated sphere in stationary fluid.

For a flagellum with axial resistance matrix of a slender helix and a head with resistances $A_0=6\pi \mu a$, $D_0=8\pi \mu a^3$, neglect interactions between the parts. A motor with relative angular velocity $\omega$ givesThe head rotates at $\Omega -\omega$. A large head creates excessive translational resistance; a small head supplies too little rotational resistance, allowing the head to counterrotate while the flagellum barely moves relative to the fluid.

For body-force-free Stokes fields with the same viscosity,To prove this Lorentz reciprocal theorem, integrate the divergence of the difference of the two cross-work fluxes. Both stress divergences are zero; incompressibility and symmetry of the Newtonian stress reduce each cross-gradient contraction to $2\mu e^{(1)}:e^{(2)}$. These are equal, so the divergence integrates to zero. The argument applies to an exterior domain by first cutting it off and taking the outer radius to infinity; the decaying velocity and stress make the outer boundary contribution vanish.

Define the generalized force as force and torque exerted by the rigid body on the fluid. With a resting far field and no-slip boundary conditions, $u=U+\Omega \times x$ on the body, and linearity givesFor two such motions the reciprocal theorem becomesSince the six-component velocities can be chosen independently, $\boxed{\mathsf{R}={\mathsf{R}}^T}$. The ordinary work identity givesfor every nonzero rigid-body motion. If the integral were zero, $e=0$ would make the connected exterior flow an infinitesimal rigid motion. Its decay at infinity requires that motion to be zero, which is incompatible with a nonzero no-slip body velocity. Thus $\mathsf{R}$ is positive definite. Fluid-on-body forces have the opposite sign; their map is $-\mathsf{R}$, not a positive-definite resistance matrix.

Write $s_{\varphi }=\sin \varphi$ and $c_{\varphi }=\cos \varphi$. The helix has $|dX/d\theta |=b\sqrt{1+{\tan }^2\varphi }=b\sec \varphi$, soIts unit tangent is $X^{\prime }=c_{\varphi }{e}_{\theta }+s_{\varphi }{e}_z$, where $e_{\theta }=(-\sin \theta ,\cos \theta ,0)$. The given slender-body force density is the force per arc length exerted on the fluid. For pure axial translation, $V=U{e}_z$, it givesFor pure rotation, $V=b\Omega e_{\theta }$, it givesThe axial torque per unit arc length is $b{f}_{\theta }$. Integrating along the wire gives the axial resistance matrix of a slender helixThus translation gives $(F_z,G_z)=(\mathcal{A}U,\mathcal{B}U)$, and rotation gives $(\mathcal{B}\Omega ,\mathcal{D}\Omega )$. These are the requested axial components; the axial swimmer approximation does not require proving statements about the other components. The equality of the two off-diagonal coefficients checks reciprocity, andFor the organism, retain the specified entrainment approximation: the head moves with the local large-scale fluid translation, and its angular velocity $\Omega -\omega$ differs from the local fluid rotation $\Omega$ by $-\omega$. Put $D_0=8\pi \mu a^3$. Its torque on the fluid is therefore $-D_0\omega$, and its force is neglected. Neutral buoyancy and absence of external torque giveSolving,This is the entrained-head approximation for a helical microswimmer, which differs from treating the head as an isolated sphere in otherwise stationary fluid. The flagellum's angular speed is also $\Omega =D_0\omega (1+c_{\varphi }^2)/(CL{b}^2)$.

At fixed $b,L,a,\omega ,ϵ$, the speed is maximal at $\varphi =\pi /4$ and tends to zero at either limiting pitch. At fixed other parameters it decreases as $1/L$: the longer flagellum has more resistance at the fixed torque supplied in this model. Holding the number of turns fixed instead would make $L$ change with pitch, so that is a different comparison.

The motor exerts equal and opposite torques of magnitude $D_0\omega$ on flagellum and head, whose relative angular speed is $\omega$. Therefore its rate of working isEquivalently, the work delivered to the fluid is $G_{\mathrm{flag}}\Omega +G_{\mathrm{head}}(\Omega -\omega )=D_0\omega \Omega -D_0\omega (\Omega -\omega )$. Translation contributes no net work because the total force vanishes. This explains the independence from $U$ and $\Omega$, without assigning an isolated-fluid drag law to the entrained head.

Choose the helix handedness so its unit tangent is $\mathbf{t}=\cos \alpha \mathbf{i}+\sin \alpha {\mathbf{e}}_{\varphi }$. Let $q={\Omega }_0-\Omega$; the body rotates at $+\Omega \mathbf{i}$, while the flagellum rotates at $-q\mathbf{i}$ in the laboratory, since its relative motor rotation is $-{\Omega }_0\mathbf{i}$. Both translate at $-U\mathbf{i}$. Thus the relative fluid velocity on a flagellar element is $\mathbf{w}=U\mathbf{i}+bq{\mathbf{e}}_{\varphi }$.

Low-Reynolds-number resistive-force theory gives force per unit length on the flagellumIts axial and azimuthal components areDefine the integrated axial resistance matrix of a slender helix coefficientsThe sphere has translational resistance $A_b=6\pi \mu a$ by the Stokes drag law and rotational resistance $H_b=8\pi \mu a^3$. Its hydrodynamic force is $+A_{b}U\mathbf{i}$ and its torque $-H_b\Omega \mathbf{i}$. Integration of $f_x$ and $b{f}_{\varphi }$ gives the zero total force and torque conditionsDividing the first by $L$ and expanding the coefficients gives exactly the two requested balances. The internal motor torque cancels in the total torque balance; it is not an external applied torque.

Eliminate $U$ and use $q+\Omega ={\Omega }_0$. The effective flagellar rotational resistance is $H_e=H-C^2/(A+A_b)$, soThe determinant of helical resistance in resistive-force theory is $AH-C^2=b^2{L}^2{K}_N{K}_T>0$, hence the denominator is positive. Opposite handedness changes the sign of the translation-rotation coupling and reverses the swimming direction.

For $K_T=K$, $K_N=2K$, $\alpha =\pi /4$ and $a=b$, the coefficients are $A=3KL/2$, $C=KaL/2$, $H=3K{a}^{2}L/2$. ThereforeEquivalently, with $h=KL/(\pi \mu a)$, $U=a{\Omega }_{0}4h/(2h^2+21h+48)$. These are positive speed magnitudes for the selected handedness; the translational velocity is $-U\mathbf{i}$.

This helical microswimmer with a spherical head approximation neglects head-tail hydrodynamic interactions as permitted, treats the flagellum as slender and rigid, and averages away transverse forces associated with finite helical ends. The flow is quasistatic Stokes flow, so neither body inertia nor fluid inertia enters the balance.

For a force $\mathbf{F}\delta (\mathbf{x})$ acting on an unbounded incompressible fluid, the Stokeslet and its pressure areThey solve $-\nabla p+\mu {\nabla }^2\mathbf{u}+\mathbf{F}\delta =0$ and $\nabla \cdot \mathbf{u}=0$. Approximate a slender body by a line distribution of these point forces on its centreline. At a point labelled by arclength $s$,To extract the large logarithm, write $s^{\prime }=s+\ell$ and approximate $\mathbf{X}(s+\ell )-\mathbf{X}(s)\simeq \ell \mathbf{t}$, $\mathbf{f}(s+\ell )\simeq \mathbf{f}(s)$, where $\mathbf{t}={\mathbf{X}}^{\prime }$ is the unit tangent vector. The near-line contribution has kernel $(I+\mathbf{tt})/(8\pi \mu |\ell |)$. Cut it off at the wire radius $R$ and a macroscopic length $L$. The two sides of the line giveThe far contribution and changes in the cutoff add terms without the leading logarithm. Since $(I+\mathbf{tt}{)}^{-1}=I-\mathbf{tt}/2$, inversion proves the leading slender-body force densityThis is force on the fluid; the fluid's force on the wire has the opposite sign. It gives twice the resistance to transverse motion as to tangential motion at this order.

The stated arclength describes the circular helix $\mathbf{X}=(a\cos \theta ,a\sin \theta ,b\theta )$. The PDF omits the second $a$ in the coordinates, so the literal printed curve and its arclength are inconsistent unless $a=1$. First use the circular interpretation consistent with the given arclength. Put $q=\sqrt{a^2+b^2}$ and $c=2\pi \mu /\ln (L/R)$. ThenRotation about $z$ has velocity $\mathbf{v}=\Omega (-a\sin \theta ,a\cos \theta ,0)$, so $\mathbf{t}\cdot \mathbf{v}=\Omega a^2/q$. The axial force density is $f_z=-c\Omega a^{2}b/q^2$. Integrating over one turn givesFor translation, $\mathbf{v}=W\hat{\mathbf{z}}$ and $\mathbf{f}=cW(2\hat{\mathbf{z}}-(b/q)\mathbf{t})$. Only the tangential term contributes to axial couple. Since $(\mathbf{X}\times \mathbf{t}{)}_z=a^2/q$, its density is $-cW{a}^{2}b/q^2$, givingThese are the off-diagonal coefficients of the axial resistance matrix of a slender helix. Their equality does not depend on this particular geometry: with symmetric local resistance $D=c(2I-\mathbf{tt})$, the rotation-induced axial force is $\int \hat{\mathbf{z}}\cdot D(\hat{\mathbf{z}}\times \mathbf{X})ds$, whereas the translation-induced axial torque is $\int (\hat{\mathbf{z}}\times \mathbf{X})\cdot D\hat{\mathbf{z}}ds$. They agree pointwise by symmetry of $D$. The full Stokes-flow statement is the Lorentz reciprocal theorem.

For completeness, retaining the literal elliptic helix instead gives $s^{\prime }=\sqrt{a^2{\sin }^2\theta +{\cos }^2\theta +b^2}$ and the axial coupling of an elliptic helixThus reciprocity holds for the literal curve as well; only the geometric coefficient and arclength change. In all these formulas the equal and opposite force or couple on the wire reverses the displayed sign.

Use the same resistive-force theory convention as before: the force density is the fluid's force on the filament. Let $\phi$ be the usual cylindrical azimuth. Choose the orientation of a left-handed helix so that, as $z$ increases, $\phi$ decreases. With $s=\sin \theta$, $c=\cos \theta$,Reversing the orientation of the tangent leaves the local drag tensor unchanged. Define $k={\xi }_{\perp }-{\xi }_{\parallel }$. From resistive-force theory, the relevant force-density components areThese scalar axial and azimuthal components are uniform along the helix. Integration over its contour length $\ell$, and use of the axial torque density $a{f}_{\phi }$, give the axial resistance matrix of a slender helix:Thus $B=C$, as also required by the Lorentz reciprocal theorem. The signs here belong to $(F,L{)}^T=-\left(\begin{array}{cc}A& B\\ B& D\end{array}\right)(U,\Omega {)}^T$, with $F,L$ forces and moments on the helix. The handedness reversal of helical hydrodynamic resistance reverses $B,C$ but leaves $A,D$ unchanged. If the positive rotation direction or handedness convention is reversed, the coupling sign reverses with it.

As a check on physical admissibility, the determinant of helical resistance in resistive-force theory isTogether with $A,D>0$, this makes the hydrodynamic resistance matrix positive definite and the viscous power loss $A{U}^2+2BU\Omega +D{\Omega }^2$ positive. Isotropic local drag would have $B=0$, so rotating a helix would not propel it in this approximation.

For the pair, use a local additive resistive-force theory model: both helices have the same drag coefficients, and inter-helix hydrodynamic interactions and unresolved end effects are omitted. The right-handed helix has resistance entries $(nA,-nB,-nB,nD)$. Define the signed motor motion byThe printed relative rotation specifies a magnitude rather than which helix rotates relative to which. Choosing ${\Omega }_2-{\Omega }_1=\omega$ instead reverses all signed speeds below.

The common translation speed $U$ and zero total force and torque obeyInternal motor forces and torques cancel in these totals. IntroduceSolving the three linear equations yields the opposite-handed counterrotating helical swimmer:For $n>0$, ${\xi }_{\perp }>{\xi }_{\parallel }$ and $\omega >0$, $B>0$, so $U<0$, ${\Omega }_1>0$ and ${\Omega }_2<0$. The opposite-handed helices counterrotate but contribute thrust in the same axial direction. The formula satisfies the specified relative rotation without identifying either laboratory rotation rate with the motor rate.

At $n=0$, the second helix supplies no hydrodynamic resistance. The remaining helix must have $F=L=0$, and invertibility of its hydrodynamic resistance matrix forces $U={\Omega }_1=0$. Formally ${\Omega }_2=-\omega$ is the rotation of a zero-resistance motor shaft or vanishing second rotor; there is no physical finite second helix to propel or to provide a reaction torque. This is the vanishing reaction rotor in a helical swimmer.

At $n=1$, equal-length opposite-handed helices haveThe translation-generated axial torques cancel between the two helices, and equal counterrotation balances the rotational torques. Their propulsive forces add, rather than cancel, because handedness and rotation both reverse. These are equal-length opposite-handed helices.

For $n\to \infty$, the large reaction helix limit isThe increasingly long second helix nearly anchors the whole assembly: its very small translation and rotation suffice to balance the finite force and torque generated by the first helix. The first helix rotates at almost the full motor rate, but moving the large resistive second helix makes the common translation tend to zero. These are idealized limits of the additive local model, rather than a resolution of short-helix end effects or infinitely extended interacting filaments.

Take $\mathbf{F},\mathbf{G}$ to be the force and couple exerted by the body on the fluid, equivalently the external force and couple needed to maintain its motion. This fixes the sign for a positive hydrodynamic resistance matrix. If one uses the fluid's force on the body, both resultants have the opposite sign.

For two Stokes flows $(\mathbf{u},\mathbf{\sigma })$ and $(\tilde{\mathbf{u}},\tilde{\mathbf{\sigma }})$ in the same fluid domain $\mathcal{D}$, with zero body force, the Lorentz reciprocal theorem for Stokes flow isHere $\mathbf{n}$ points out of the fluid. To see the identity, take the divergence of the difference of the two cross-work fluxes. The stress divergences vanish, while incompressibility and symmetry of the Newtonian fluid stress tensor reduce the remaining terms to $2\mu (\mathbf{e}:\tilde{\mathbf{e}}-\tilde{\mathbf{e}}:\mathbf{e})=0$. The divergence theorem proves the result.

On the body, the no-slip boundary condition is $\mathbf{u}=\mathbf{U}+\mathbf{\Omega }\times \mathbf{r}$, and the far-field contribution vanishes for decaying exterior flows. Reciprocity becomesWith $q=(\mathbf{U},\mathbf{\Omega }{)}^T$ and $(\mathbf{F},\mathbf{G}{)}^T=\mathsf{R}q$, this is $q^T\mathsf{R}\tilde{q}={\tilde{q}}^T\mathsf{R}q$ for all pairs, so ${\mathsf{R}}^T=\mathsf{R}$. The power identity givesEquality would force zero strain throughout the connected exterior domain, hence a rigid fluid motion. Decay at infinity makes that motion zero, and the no-slip boundary condition then gives $q=0$. This proves the symmetry and positivity of a rigid-body resistance matrix:

For the helix, assume $0<\varphi <\pi /2$ and use cylindrical unit vectors. Its arclength element and tangent areIntegrating to $\theta =L\cos \varphi /b$ gives $\boxed{\text{total arc length}=L}$. In a combined axial translation and rotation, $\mathbf{V}=U{\mathbf{e}}_z+b\Omega {\mathbf{e}}_{\theta }$. The slender-body force density, or local resistive-force theory, givesWriting $s_{\varphi }=\sin \varphi$, $c_{\varphi }=\cos \varphi$, its axial and azimuthal components areThe axial couple density is $b{f}_{\theta }$. Integration over arclength gives the axial resistance matrix of a slender helix:Thus pure translation gives $(F_z,G_z)=(AU,BU)$, and pure rotation gives $(F_z,G_z)=(B\Omega ,D\Omega )$. The mixed coefficients agree, as reciprocity requires, and $AD-B^2=C^2{L}^2{b}^2/2>0$. The sign of $B$ follows the handedness in the parametrization; reversing handedness reverses propulsion.

For the helical microswimmer with a spherical head, the diagram's rotation relation is ${\Omega }_0=\Omega -\omega$. Neutral buoyancy and the absence of external forcing make the whole swimmer force-free and torque-free. Neglecting interactions between its parts givesWith $\Delta =(A_0+A)(D_0+D)-B^2>0$, solving this pair givesThe head counterrotates, providing the reaction to the flagellum's rotation. The figure fixes this relative rotation convention; it is duplicated and corrupted in the local TeX.

For a very large head, the coefficient regime is $A_0\gg A$ and $D_0\gg D$. ConsequentlyThe large translational resistance $A_0$ makes the microswimmer slow even though the flagellum rotates almost at the motor rate.

For a very small head, $A_0\ll A$ and $D_0\ll D$. The appropriate denominator retains the translation-rotation coupling:Both tend to zero with $D_0\propto a^3$, while ${\Omega }_0\simeq -\omega$. The motor mainly rotates the low-resistance head, and produces little flagellar motion relative to the fluid.

In the intermediate coefficient regime $A_0\ll A$ and $D_0\gg D$, the head supplies a strong rotational reaction with little translational penalty. ThenThe head radius has dropped out. To find the optimal pitch of a helical microswimmer, put $z=\tan \varphi$. The dimensionless speed becomes $z/(z^2+2)$ and has derivative $(2-z^2)/(z^2+2{)}^2$. Hence, for $\omega >0$ with the chosen handedness,The angle is measured from the horizontal plane. An intermediate pitch combines axial and transverse tangent directions, allowing anisotropic drag to convert rotation into translation; either extreme removes that coupling.

The geometric head-size regimes quoted in the paper hold with the logarithmic slenderness factor treated as fixed. More precisely, $A_0\ll A$ requires $a\ll L(1+{\cos }^2\varphi )/(3|\log ϵ|)$, while $D_0\gg D$ requires $a^3\gg L{b}^2(1+{\sin }^2\varphi )/(4|\log ϵ|)$. These coefficient inequalities state the validity of the intermediate approximation when the logarithm is quantitatively important. The many-turn and local-drag assumptions of slender-body theory also remain in force.