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where
is an i-mer of Shh,
is the maximum number of Shh proteins which can associate into a single oligomer,
and
are respectively the association and dissociation rates in the reaction
, and the other parameters are the same as previously. Unspecified boundary conditions are zero-flux conditions, and unspecified initial conditions are 0.
The diffusion rates for oligomers were scaled from the rate for the monomer, according to the Stokes-Einstein law for spherical particles:
.