QWOP's real idea was never "hard controls" — it was taking away the verb you expected. You asked to run and it handed you the layer underneath, so locomotion stopped being a button and became a skill. The flagella model already had that layer, and the organism came with a gift: Pterosperma has four cilia. One per key.
| Key | Cilium | What it does |
|---|---|---|
| Q | outer left | Hold and dynein engages — that cilium beats, ramping from its idle 10 Hz up to a 95 Hz power stroke. Release and it unfurls and slows again. |
| W | inner left | |
| O | inner right | |
| P | outer right |
Tapping is different from holding. A key press resets that cilium to the start of its power stroke. That is the whole game, because of the thing sitting between hold and tap:
The cilia are detuned. Each runs at a slightly different intrinsic frequency, as real cilia do. Hold all four down and within a second or two they have drifted out of phase, their thrust vectors start pointing in different directions, they partly cancel, and the leftover torque spins you. Four cilia in phase pull together into one compound cilium — which is the thing the real organism does to swim — and that is far faster than four filaments arguing.
So you cannot just hold everything down. You have to re-tap, in rhythm, to drag them back into phase. Watch the four phase dials under the keys: when they converge you are bundled, and the cell glows. Roughly a quarter of a second between taps is a good place to start.
Four buttons appear under the game, one per cilium, in the same colours as the filaments they drive. They are real multi-touch — press two, three or all four at once, which you will need to, because that is the game. Tap anywhere on the title or the death screen to start a run. The on-canvas letter badges drop out when the buttons are showing, leaving just the four phase dials, since those are the part the buttons cannot tell you.
Steering falls out of the same mechanic. Drive only the left pair and the thrust acts off-centre at the front of the cell, so you turn — and because you have stopped driving the other two, your bundle collapses while you do it. Every dodge is paid for in the speed you spent effort building. That is the decision the game is made of.
This part is straight out of the paper. A real Pterosperma is stopped 96.6% of the time, and the authors read it as a sit-and-wait animal. Here that statistic becomes a tactic: a beating cell pushes a flow signature that predators feel from a long way off, and a stopped one is nearly invisible. The dashed ring around you is how far you can currently be heard. Sprinting is how you make distance and how you get eaten.
| Hunts by | Counterplay | |
|---|---|---|
| copepod | Patrols, hears a moderate distance, gives up quickly | You can outswim it. Just go. |
| medusa | Sits still, hears a very long way, slow to start | Go quiet early and drift past. Do not sprint near one. |
| arrow worm | Short-sighted but faster than you, and relentless | It must not hear you at all. Once it has you, it has you. |
The cilia are not animation. Each one is a full instance of the model in
../proteus/flagella.js — the Chebyshev waveform, the linear
dispersion relation setting its wavelength from its frequency, and Gray &
Hancock resistive force theory turning its travelling wave into thrust. The
body is overdamped, as anything this small must be: there is no inertia and
nothing coasts, so velocity is force over drag and releasing every key stops
you almost at once. Everything comes from
Embodied behavioural complexity in a ciliated microorganism,
Nature Communications 17, 8445 (2026).
Four things are the game's and not the paper's, and are marked as such in
game.js: how much a bundle helps (set by the skill gradient it
produces), how readily the cell turns (the true value is far too sluggish to
dodge with), a symmetric four-cilium arrangement in place of the real 3+1 one
so that left and right steer alike, and the predators, which are invented
entirely. The detune, the mirrored left/right beating, and the phase-reset are
all real ciliary behaviour — Chlamydomonas swims straight precisely
because its flagella beat as a mirrored pair.
There is a variant at /graze/ where the beating costs ATP and there is prey as well as predators — and where the paper's 96.6% is left to emerge from the economics instead of being quoted at you.
The instrument at /flag/ shows the same model with every measurement it came from; the amoeba at /proteus/ hangs it off a crawling cell instead.
And two more keep the four keys but throw the biology away: /qgol/ is Conway's Life with a generation split across Q W O P, three keys that mark and one that commits; /griddle/ is a pancake station where the face that is cooking is the one you cannot see.