Skip to main content
Sandbox Physics
Microswimming · A course with experiments

L4 · Chapter 8

Two swimmers feel each other through the timing of their strokes

A symmetric swimmer’s cycle-averaged flow is weak far away, falling as the inverse cube of distance. Pooley, Alexander and Yeomans showed that the interaction between two such swimmers is dominated by a different term, one that depends on how their strokes line up in time. Swimmers in phase can attract, repel, oscillate or travel on parallel paths depending on where they start; half a cycle out of phase, the sign of that term flips. Every stroke here runs on a fixed clock, so these results describe prescribed phases.

A pump up close, a weak field far away

Pooley, Alexander and Yeomans study the three-sphere swimmer with separate stroke amplitudes for its front and back arms. Averaged over a cycle, its far flow is a sum of point-force singularities. When the two amplitudes are equal, the stroke looks the same if you play it backward in time and flip the swimmer front to back, a property the paper calls TP invariance. Then the dipolar part of the mean flow vanishes and the far field falls as the inverse cube of distance, in line with Golestanian and Ajdari’s symmetric case.

Close to the swimmer the averaged flow looks different. Near the middle sphere the fluid moves backward: the swimmer acts as a pump that carries fluid from in front of it to behind it, and that is how it moves forward. The paper’s analytic far-field expression matches its numerical flow to within ten percent beyond about three swimmer lengths.

Dead and live terms

Now place a second swimmer nearby; the authors call the two Alice and Bob. Bob’s effect on Alice has two parts. The dead term is what Alice would feel if Bob’s cycle-averaged flow simply carried and turned her. It depends only on Bob’s flow and takes the expected dipolar form. Its size is proportional to the product of Bob’s front and back amplitudes times their difference, so it vanishes when they are equal.

The live term comes from both swimmers moving at once. Bob’s flow pushes on Alice’s spheres at particular moments of her own stroke, and the products of those instantaneous pushes leave a residue after averaging. Its size depends on the relative phase of the two strokes through a phase function that the paper plots in Figure 3. Equation 8 compares the two terms and finds the live term dominant.

ΔxAdead∝a2 ξB ξF (ξB−ξF)D2 r2,ΔxAdead=0  for  ξB=ξF\Delta\mathbf{x}_{A}^{\mathrm{dead}}\propto\frac{a^{2}\,\xi_{B}\,\xi_{F}\,(\xi_{B}-\xi_{F})}{D^{2}\,r^{2}},\qquad\Delta\mathbf{x}_{A}^{\mathrm{dead}}=\mathbf{0}\ \ \mathrm{for}\ \ \xi_{B}=\xi_{F}

Worked example

How many strokes before they meet

Figure 4(i) of the paper starts Alice at the origin and Bob five arm lengths above her, both aligned and in phase. All lengths are in units of the arm length: the sphere radius is one tenth and the stroke amplitude three tenths. The paper gives the step per cycle as seven twelfths of radius times amplitude squared over arm length squared, about 0.005 arm lengths.

This is the small-loop area law of Chapter 5. With both arms equal to the arm length, the coefficient is seven twelfths of radius over arm length squared, and the loop is a square whose side is the amplitude. The step comes out at 0.00525 arm lengths. The swimmers travel about 25 arm lengths before they collide, so the run lasts about 4,800 cycles, consistent with the paper’s figure of about five thousand. The trajectories stop when the swimmers come so close that the point-force approximation fails.

Now put in sizes. With ten-micrometre arms, one-micrometre spheres, three-micrometre strokes and arms moving at thirty micrometres per second, a cycle takes 0.4 seconds, so 4,800 cycles take about 32 minutes. In that time a single one-micrometre sphere in water wanders about 28 micrometres along any one direction by thermal motion. The three-sphere body wanders somewhat less, still several arm lengths. This estimate is ours, made with the diffusion coefficient from Chapter 2, and it shows how slowly the pair effects build compared with Brownian motion at this size.

Δ=712 a ξ2D2=712×0.1×0.09 D≈0.00525 D,N≈25 DΔ≈4.8×103\Delta=\frac{7}{12}\,\frac{a\,\xi^{2}}{D^{2}}=\frac{7}{12}\times0.1\times0.09\,D\approx0.00525\,D,\qquad N\approx\frac{25\,D}{\Delta}\approx4.8\times10^{3}

The phase decides the sign

For in-phase swimmers with equal amplitudes, Figure 4(a) divides Bob’s starting positions into four regions: attraction, repulsion, oscillation and parallel paths. Besides the attracting pair above, a repelling pair turns each swimmer through about seven degrees, mostly within 3,000 cycles. In the oscillating region the swimmers oscillate relative to each other, one oscillation taking about 10,000 cycles, with a slow drift on top. The authors trace the oscillation to the rotational part of the live term and the drift to its translational part.

Shift the swimmers half a cycle out of phase and the live term changes sign. In Figure 4(b) the attracting region becomes repelling, and the oscillating region turns into parallel paths, tilted slightly from the swimming direction, along which the two swimmers wobble exactly half a period apart. With a quarter-cycle lag, Figure 4(c), the lowest-order live term vanishes. The behaviour still differs for a partner in front and a partner behind, which the authors read as a sign that live terms still dominate.

Unequal amplitudes break the perfect periodicity of the in-phase oscillations. A smaller front amplitude makes the oscillations grow until the swimmers diverge. A larger front amplitude makes them shrink and the swimmers close in, which suggests that in-phase suspensions might form chains. The authors add that attraction between two swimmers is insufficient evidence for clustering in larger groups.

What the pair Lab reproduces

Our calculation: the pair Lab uses the sizes above and couples all six spheres of the two swimmers as point forces. Side by side, five arm lengths apart and in step, the two swimmers turn toward each other and meet after about 2,700 strokes. For a stroke of three tenths of an arm, the full point-force step is 0.0824 micrometres per cycle, about 1.6 times the small-loop estimate. At that step, 25 arm lengths take about 3,000 cycles, close to the Lab’s count. The paper quotes the small-loop step for its runs, and we have not traced how its figure of five thousand was computed.

Half a cycle out of step, the same pair turns apart, as the sign flip of the live term predicts. Far apart, Alice’s turn per stroke varies with Bob’s bearing, measured from her swimming direction, with the angular shape of the leading-order result of Pooley and colleagues. The Lab plots the two side by side.

ΔθA∝sin⁡α(3−30cos⁡2α+35cos⁡4α)\Delta\theta_{A}\propto\sin\alpha\left(3-30\cos^{2}\alpha+35\cos^{4}\alpha\right)

Fixed clocks and real synchrony

Every stroke in this paper runs on its own fixed clock. The calculations show what a given phase does to the trajectories, and the phases themselves stay put. Pooley and colleagues note that externally driven artificial swimmers could be held strictly in phase, which makes their predictions testable, while autonomous organisms would let their phases drift, perhaps toward phase locking.

Lauga and Powers review when phases do lock. Taylor showed that two swimming sheets with the same prescribed waveform dissipate least when they beat in phase. Later simulations found that such sheets drift until they are exactly in phase or exactly out of phase, with both states stable. For real flagella the waveform itself responds to the forces from a neighbour, and that feedback is expected to allow perfect phase locking. Rotating helices tell a similar story: perfectly rigid ones fail to synchronise, and some flexibility is needed. A model of synchrony therefore lets each stroke respond to the force it meets.

An experiment to think through

Predict, change, explain

  1. Two identical swimmers start side by side, five arm lengths apart, with their strokes in step. Predict whether the gap shrinks or grows, and what changes when one stroke runs half a cycle ahead.
  2. Run “In step side by side: drawn together” and watch Sideways gap and Strokes so far. Then run “Opposite phase side by side: pushed apart” and compare Alice’s heading with Bob’s heading.
  3. Explain, with the live term, why shifting one stroke by half a cycle reverses the outcome.

The pair Lab couples the six spheres of two swimmers in unbounded water and holds their strokes at a fixed phase.

Try explaining it yourself

1. Compute the step per cycle for Pooley’s swimmer with a sphere radius of 0.1 and a stroke amplitude of 0.3 arm lengths, and the number of cycles needed to cover 25 arm lengths.

Read an explained answer

Seven twelfths times 0.1 times 0.09 gives 0.00525 arm lengths per cycle. Twenty-five arm lengths then take about 4,800 cycles.

2. Why does the dead term vanish for a swimmer with equal front and back amplitudes?

Read an explained answer

The dead term is proportional to the product of the two amplitudes times their difference. Equal amplitudes make the difference zero, and the cycle-averaged flow then has no dipolar part.

3. Two identical in-phase swimmers attract. What does the paper predict when one of them is shifted by half a cycle?

Read an explained answer

The live term changes sign, so the attracting configuration becomes a repelling one.

4. Can these calculations show two swimmers falling into step on their own?

Read an explained answer

The phases are fixed inputs, so phase locking lies outside these calculations. Lauga and Powers point to strokes or waveforms that respond to hydrodynamic forces, and to flexibility, as the ingredients synchrony needs.

Sources to read alongside

  1. Lauga & Powers, The hydrodynamics of swimming microorganisms, arXiv:0812.2887v1 (2008); Reports on Progress in Physics 72, 096601 (2009)

    Preprint version read. Sections 3 and 4 supply the Stokes equations, the point-force flow and the proofs of the scallop theorem; Sections 5 to 7 cover resistive-force theory, flexible filaments, walls and synchronisation.

  2. Golestanian & Ajdari, Three-sphere low Reynolds number swimmer revisited, arXiv:0711.3700v1 (2007); Physical Review E 77, 036308 (2008)

    Preprint version 1 read; the journal version was not checked. Equations 1 to 6, 9, 12, 13 and 16, Section IV on the far-field flow and footnote 23 are used.

  3. Pooley, Alexander & Yeomans, Swimming with a friend at low Reynolds number, arXiv:0705.3612v1 (2007); Physical Review Letters 99, 228103 (2007)

    Preprint version 1 read; the journal version was not checked. Equations 1 and 4 to 8, Figure 3 and the trajectories of Figure 4 are described in words; no curve was digitised.

FeedbackSomething broken, confusing, or missing on this page?Tell us
What kind of feedback?

Sent with this page’s address, its Lab ID, the site language and your device type. Please leave out personal details. Feedback is kept for up to 12 months.Privacy policy