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Big Idea

The outer solution differs from the exact one only during the first microsecond. How large the error is depends on the observable: the discarded transient has height of order one and area εL\ve L, so the instantaneous velocity is 100%100\% wrong while the position is wrong by ε\ve of a body length for all time. Position is what we predict, so the reduction stands, and the equation of motion from here on is γ x˙=F\gamma\,\dot{\mathbf x} = \mathbf F.

The approximation error

From here on every model uses the reduced equation γx˙=F\gamma\dot x = F in place of mx¨=F−γx˙m\ddot x = F - \gamma\dot x. Because the exact solution is known, the error this commits, in velocity and in position, can be computed exactly rather than estimated.

Proposition 1 (The full error of the reduced model)

Let vexactv_\mathrm{exact} solve (2) with constant FF and v(0)=v0v(0) = v_0, and let vout=F/γv_\mathrm{out} = F/\gamma be the outer solution (2). Let xexactx_\mathrm{exact} and xoutx_\mathrm{out} be the corresponding positions, both starting from the same x(0)x(0). Then for every t≥0t \ge 0,

vexact(t)−vout=(v0−Fγ)e−t/τi,v_\mathrm{exact}(t) - v_\mathrm{out} = \Bigl(v_0 - \frac{F}{\gamma}\Bigr)e^{-t/\tau_i} ,
xexact(t)−xout(t)=(v0−Fγ)τi(1−e−t/τi),x_\mathrm{exact}(t) - x_\mathrm{out}(t) = \Bigl(v_0 - \frac{F}{\gamma}\Bigr)\tau_i\Bigl(1 - e^{-t/\tau_i}\Bigr) ,

and therefore

sup⁡t≥0∣xexact−xout∣=∣v0−Fγ∣ τi=∣v~0−F~∣ ε L.\sup_{t\ge0}\bigl|x_\mathrm{exact} - x_\mathrm{out}\bigr| = \Bigl|v_0 - \frac{F}{\gamma}\Bigr|\,\tau_i = \bigl|\tilde v_0 - \tilde F\bigr|\,\ve\,L .

Proof 1

Subtract F/γF/\gamma from (10) to get (1). Integrating it from 0 to tt, with the two positions agreeing at t=0t = 0, gives

xexact(t)−xout(t)=(v0−Fγ)∫0te−s/τi ds=(v0−Fγ)τi(1−e−t/τi),x_\mathrm{exact}(t) - x_\mathrm{out}(t) = \Bigl(v_0 - \frac{F}{\gamma}\Bigr)\int_0^t e^{-s/\tau_i}\,\dd s = \Bigl(v_0 - \frac{F}{\gamma}\Bigr)\tau_i\bigl(1 - e^{-t/\tau_i}\bigr) ,

which is (2). It is monotone in tt and tends to (v0−F/γ)τi(v_0 - F/\gamma)\tau_i, giving (3); dividing by LL and using ε=τi/T=τiv0/L\ve = \tau_i/T = \tau_i v_0/L puts it in dimensionless form.

Source
<Figure size 858x325 with 2 Axes>

The same reduction judged in two variables. Left: the velocity error (1) is O(1)O(1) at t=0t = 0, so inside the layer the reduced model is not approximately wrong but completely wrong. Right: the position error (2) accumulates only while the layer lasts and then stops, saturating at O(ε)O(\ve) of a cell diameter for all later time.

The two lines of Proposition 1 disagree by nine orders of magnitude about the same pair of trajectories, so “the error of the reduction” needs a precise meaning.

Definition 1 (Observable, and the error of a reduction in it)

An observable is a map QQ that takes a trajectory to a number: the position at a given time, the velocity at a given time, the displacement over an interval, the time spent in a region. The error of the reduction in QQ is

E[Q]=∣Q[xexact]−Q[xout]∣,E[Q] = \bigl|Q[x_\mathrm{exact}] - Q[x_\mathrm{out}]\bigr| ,

measured against the scale on which QQ itself is read. A reduction is not accurate or inaccurate on its own. It is accurate in an observable.

Nothing here refers to an instrument: both trajectories are known in closed form and E[Q]E[Q] is a computation. The two lines of Proposition 1 are E[Q]E[Q] for two choices of QQ:

QQE[Q]E[Q]read againstrelative, worst case
x(t)x(t)≤∣v0−F/γ∣ τi\le \bigl\lvert v_0 - F/\gamma\bigr\rvert\,\tau_iLLε\ve
x˙(t)\dot x(t)∣v0−F/γ∣ e−t/τi\bigl\lvert v_0 - F/\gamma\bigr\rvert\,e^{-t/\tau_i}∣v0−F/γ∣\bigl\lvert v_0 - F/\gamma\bigr\rvert1

Worst case means t=0t = 0 and ∣v~0−F~∣=1|\tilde v_0 - \tilde F| = 1, a cell that loses its traction while moving at v0v_0.

Remark 1 (One transient, two numbers)

Everything the reduction discards is the term (v0−F/γ)e−t/τi(v_0 - F/\gamma)e^{-t/\tau_i} in the velocity. It has height ∣v0−F/γ∣\lvert v_0 - F/\gamma\rvert, the full velocity scale, and width τi\tau_i, which is ε\ve of the time scale TT, so its area is small although its height is not:

∫0∞∣vexact−vout∣dt=∣v0−Fγ∣τi=∣v~0−F~∣ εL.\int_0^\infty \bigl\lvert v_\mathrm{exact} - v_\mathrm{out} \bigr\rvert \dd t = \Bigl\lvert v_0 - \frac{F}{\gamma}\Bigr\rvert \tau_i = \bigl\lvert \tilde v_0 - \tilde F \bigr\rvert\, \ve L .

An observable that reads the velocity at an instant sees the height. Every observable built by integrating the velocity over an interval, the position first among them, sees only the area.

Example 1 (What the reduction throws away)

A cell moving at v0v_0 loses all traction at t=0t = 0, so F=0F = 0 and ∣v~0−F~∣=1|\tilde v_0 - \tilde F| = 1, the largest value it can take. The distance the cell coasts before stopping, which is also the entire error the reduced model ever commits in position, is

xcoast=v0 τi≈(10−8 m/s)(10−6 s)=10−14 m,x_\mathrm{coast} = v_0\,\tau_i \approx \bigl(10^{-8}\ \mathrm{m/s}\bigr)\bigl(10^{-6}\ \mathrm{s}\bigr) = 10^{-14}\ \mathrm{m} ,

ten femtometres, about the diameter of a heavy atomic nucleus. Against the yardstick we chose for position,

xcoastL=v0τiL=τiL/v0=τiT=ε.\frac{x_\mathrm{coast}}{L} = \frac{v_0\tau_i}{L} = \frac{\tau_i}{L/v_0} = \frac{\tau_i}{T} = \ve .

This answers the question of From Newton’s Second Law to a Force Balance. A swimmer glides a body length or two; a cell glides 10-9 of one. The relative error in the observable is the small parameter, which is what a well-chosen set of scales does: ε\ve is exactly the fraction of a cell diameter that a cell coasts. The position is wrong by at most εL≈10−14\ve L \approx 10^{-14} m at any time, four orders of magnitude below the diameter of an atom, which no instrument and no mechanism inside the cell can resolve. There is no velocity memory to delete.

The velocity on a window

The two rows of the table sit at the two ends of one scale. The position averages the velocity over the whole of [0,t][0, t]; the instantaneous velocity averages it over nothing. The intermediate case averages over a window of length Δt\Delta t.

Definition 2 (Windowed velocity)

For Δt>0\Delta t > 0, the velocity of a trajectory averaged over a window Δt\Delta t is

vˉΔt(t)=1Δt∫tt+Δtx˙(s) ds=x(t+Δt)−x(t)Δt.\bar v_{\Delta t}(t) = \frac{1}{\Delta t}\int_t^{t + \Delta t} \dot x(s)\,\dd s = \frac{x(t + \Delta t) - x(t)}{\Delta t} .

It is a difference quotient of the position, defined for every trajectory and every Δt>0\Delta t > 0, and its Δt→0\Delta t \to 0 limit is x˙(t)\dot x(t).

Proposition 2 (The error in the windowed velocity)

Under the hypotheses of Proposition 1, vˉΔt\bar v_{\Delta t} of the outer solution is F/γF/\gamma for every tt and every Δt\Delta t, and

∣vˉΔt exact(t)−vˉΔt out∣∣v0−F/γ∣=τiΔt e−t/τi(1−e−Δt/τi)  ≤  min⁡{1, τiΔt}.\frac{\bigl\lvert \bar v_{\Delta t}^{\,\mathrm{exact}}(t) - \bar v_{\Delta t}^{\,\mathrm{out}} \bigr\rvert} {\bigl\lvert v_0 - F/\gamma \bigr\rvert} = \frac{\tau_i}{\Delta t}\,e^{-t/\tau_i} \bigl(1 - e^{-\Delta t/\tau_i}\bigr) \;\le\; \min\Bigl\{ 1,\ \frac{\tau_i}{\Delta t} \Bigr\} .

Proof 2

By (9) the error is a difference of two position errors divided by Δt\Delta t, so (2) gives

vˉΔt exact(t)−vˉΔt out=(v0−Fγ)τiΔt(e−t/τi−e−(t+Δt)/τi),\bar v_{\Delta t}^{\,\mathrm{exact}}(t) - \bar v_{\Delta t}^{\,\mathrm{out}} = \Bigl(v_0 - \frac{F}{\gamma}\Bigr)\frac{\tau_i}{\Delta t} \Bigl(e^{-t/\tau_i} - e^{-(t + \Delta t)/\tau_i}\Bigr) ,

which is (10) after factoring e−t/τie^{-t/\tau_i}. For the bound, e−t/τi≤1e^{-t/\tau_i} \le 1 and 1−e−u≤min⁡{1,u}1 - e^{-u} \le \min\{1, u\} with u=Δt/τiu = \Delta t/\tau_i.

The error (10) depends on the window only through Δt/τi\Delta t/\tau_i, and it contains both rows of the table:

There is one error, the fixed area (6) divided by the length of the window the observable averages over.

Source
<Figure size 598x351 with 1 Axes>

The relative error (10) at t=0t = 0 over eleven decades of window. The plateau on the left is the instantaneous velocity and its 100%100\% error, the falling branch is τi/Δt\tau_i/\Delta t, and the knee is at Δt=τi\Delta t = \tau_i. The marked windows are the layer itself, video rate, a tracking interval of one minute, and the migration time TT, whose value on the curve is ε\ve.

Example 2 (The windows a cell experiment works with)

What an experiment fixes is which Δt\Delta t is in play: positions recorded one frame apart give vˉΔt\bar v_{\Delta t} at that Δt\Delta t, and x˙\dot x itself is never available. With τi=10−6\tau_i = 10^{-6} s, at the worst time t=0t = 0:

Window Δt\Delta tRelative error τiΔt(1−e−Δt/τi)\dfrac{\tau_i}{\Delta t}\bigl(1 - e^{-\Delta t/\tau_i}\bigr)
Δt→0\Delta t \to 0, instantaneous1
τi=1 μ\tau_i = 1\ \mus, the layer itself0.63
1/301/30 s, video rate3×10−53\times10^{-5}
1 s10-6
1 min, typical for cell tracking2×10−82\times10^{-8}

Every window a cell experiment uses sits in the last three rows. To reach the top of the table the window would have to be shorter than a microsecond, over which a cell moves ten femtometres.

So a reported error is incomplete without the observable it belongs to, and for a velocity the observable is incomplete without its window. The number τi/Δt\tau_i/\Delta t is the same kind of number as ε=τi/T\ve = \tau_i/T: the cell’s own time divided by the time over which the observer averages. A reduction is justified by an error estimate in a named observable, not merely by a small term in the equation.

Summary

Definition 3 (Overdamped force balance)

For a cell in a viscous medium, on any time scale long compared with τi=m/γ\tau_i = m/\gamma, the equation of motion is

γ x˙=F(x,t).\gamma\,\dot{\mathbf x} = \mathbf F(\mathbf x, t).

Force sets velocity, not acceleration.

StepResult
Newton, with Stokes dragmx¨=F−γx˙m\ddot x = F - \gamma\dot x
set F=0F = 0 and solvea time appears: τi=m/γ≈1 μ\tau_i = m/\gamma \approx 1\ \mus
a length and a speeda second time: T=L/v0≈17T = L/v_0 \approx 17 min
scale, and their ratio appearsε=τi/T≈10−9\ve = \tau_i/T \approx 10^{-9}
set ε=0\ve = 0γx˙=F\gamma\dot x = F, plus a layer of width τi\tau_i
approximation errorposition error at most εL\ve L; velocity error τi/Δt\tau_i/\Delta t on a window Δt\Delta t

No value of the traction force was needed; the argument is about time scales. It used three techniques, none of them specific to cells: check units and find the times the constants form (Example 1); nondimensionalize so that the size of each term sits in its prefactor (Definition 4); and, when setting ε=0\ve = 0 lowers the order, construct outer and inner solutions, match them, and estimate the error in a named observable (Definition 3, Proposition 2).

What the equation says about the physics:

References
  1. Purcell, E. M. (1977). Life at low Reynolds number. American Journal of Physics, 45(1), 3–11.