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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.

A force-free, torque-free microswimmer in a stationary Newtonian fluid, with negligible fluid and body inertia and steady Stokes flow at each instant, gains no net displacement from a reciprocal stroke: one whose sequence of shapes is exactly retraced in reverse. The environment and boundary conditions must also be unchanged during that retracing. Opening a two-shell swimmer and then closing it follows the same sequence backwards; doing the closure faster does not help, because Linearity of Stokes flow changes the instantaneous velocity by the same factor as the stroke rate.

A single real shape coordinate that travels out and back along an interval gives this cancellation, as computed in the force-free two-sphere stroke. Two independently controlled shape coordinates can instead execute a loop that is not retraced, avoiding this obstruction. Continuous rotation also need not retrace an interval: the helical microswimmer with a spherical head provides another route to propulsion. The scallop theorem is a restriction on a stroke under stated conditions, not a claim that small organisms cannot swim. Purcell presented the opening-and-closing scallop and a swimmer with two hinges in Life at low Reynolds number (1977), figures 6–7.

A squirmer is an idealized spherical microswimmer propelled by a prescribed tangential surface slip velocity. The Lorentz reciprocal theorem for Stokes flow determines its rigid translation and rotation without solving its complete exterior flow.

Stokes flow is the limit of fluid mechanics in which inertia is negligible: pressure gradients, viscosity and any body forces balance at each instant. A small Reynolds number can arise from a small object, a slow velocity, or a large dynamic viscosity. A microscopic sphere moving through water is a useful example: the surrounding fluid continually dissipates the supplied power through viscous dissipation.

The picture is a meridional slice of the three-dimensional axisymmetric flow around a sphere, with the sphere held fixed and the distant fluid moving rightward at speed $U$. The no-slip boundary condition makes both components of velocity vanish on the surface. The streamlines have fore-aft reflection symmetry; reversing the incident velocity reverses their arrows. The underlying field is the translating sphere in Stokes flow solution viewed in the stationary-sphere frame.

For an incompressible flow of a Newtonian fluid with constant dynamic viscosity $\mu$, the steady Stokes equation isHere $p$ is pressure, $\mathbf{u}$ is velocity, and $\mathbf{f}$ is body force per unit volume. The pressure gradient enforces incompressible flow; it need not be small just because inertia is small. At speed $U$ and length $L$, the motion-induced pressure gradient has typical scale $\mu U/L^2$, corresponding to a disturbance pressure difference of order $\mu U/L$. See Shreyas Mandre's fluid dynamics notes, sections 9.1.1–9.1.2.

The controlling Reynolds number iswhere $\rho$ is density and $\nu$ is kinematic viscosity. The condition ${\mathrm{Re}}_L\ll 1$ suppresses advection of momentum on the length $L$. Rapid changes require an additional check of the viscous diffusion time: small Reynolds number alone does not eliminate the time derivative in the Unsteady Stokes equation.

Two rewards of this limit are the exact Stokes's law for the drag on a sphere and kinematic reversibility of Stokes flow. The first predicts settling velocities and connects to the Stokes–Einstein relation; the second explains why a microswimmer needs a nonreciprocal stroke, as expressed by the scallop theorem. Despite viscous dissipation, reversing the driving can retrace the motion: an especially striking consequence of Linearity of Stokes flow.

George Gabriel Stokes obtained the sphere drag and its terminal velocity in his 1851 investigation of how viscosity changes physical pendulum motion. He also used the slow fall of small water droplets to discuss clouds. The paper was read on 9 December 1850; 1851 is its publication date. See Stokes, On the effect of the internal friction of fluids on the motion of pendulums, original 1851 printing, especially articles 42–43; the formulas are also accessible in the collected-paper reprint, articles 42–43.

The Taylor swimming sheet is an infinite two-dimensional microswimmer whose prescribed traveling deformation drives Stokes flow. A transverse wave $y_s=ϵ\sin (x-t)$ of small amplitude swims opposite to its direction of propagation, with speed $U={ϵ}^2/2+O({ϵ}^4)$ when lengths are scaled by inverse wavenumber and velocities by wave speed. Its biharmonic stream function for planar Stokes flow has first-order part ${\psi }_1=(1+y)e^{-y}\sin (x-t)$.