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Turing's next steps: the mechanochemical basis of morphogenesis


Nearly 60 years ago, Alan Turing showed theoretically how two chemical species, termed morphogens, diffusing and reacting with each other can generate spatial patterns. Diffusion plays a crucial part in transporting chemical signals through space to establish the length scale of the pattern. When coupled to chemical reactions, mechanical processes — forces and flows generated by motor proteins — can also define length scales and provide a mechanochemical basis for morphogenesis. forces and flows generated by motor proteins — can also define length scales and provide a mechanochemical basis for morphogenesis.

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Figure 1: Patterning of biochemical species by reaction–diffusion mechanisms.
Figure 2: Patterning by motor-mediated transport.
Figure 3: Length scales set up by stresses.


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We thank N. Goehring, M. Mayer and F. Jülicher for discussions, and I. Tolic-Norrelykke for comments on the manuscript.

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Correspondence to Jonathon Howard, Stephan W. Grill or Justin S. Bois.

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The authors declare no competing financial interests.

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Active diffusion

Random motion caused by randomly directed active forces, such as those generated by motor proteins.


Directed transport driven by motor proteins or bulk fluid flow.


The randomly directed motion of a molecule or particle that causes both mixing and the flux of particles from regions of high concentration to low concentration. Diffusion can be caused by thermal forces — that is, collisions with molecules in solution — or by randomly directed active forces, such as those generated by motor proteins that randomly change their direction.

Diffusion coefficient

The constant of proportionality between the flux and the concentration gradient for a diffusing particle. Diffusion can be thermal or active.

Friction coefficient

The constant or proportionality between a stress gradient and velocity.

Length constant

The distance over which a quantity such as concentration decreases e-fold.


Substances, such as proteins or small molecules, that are non-uniformly distributed in space and can influence cell growth or differentiation.


The establishment of features that are much larger than those of the individual molecular components, and which are stereotyped from one cell to another or one organism to another.

Reaction–diffusion mechanism

A patterning process in which a diffusing morphogen undergoing chemical reactions (such as degradation or synthesis) forms a well-defined spatial distribution.


Force per unit area.

Viscoelastic material

A material that is both elastic (it can be stretched but returns to its original shape) and viscous (it deforms at a finite speed determined by the viscosity and the applied stress).


The constant of proportionality between rates of stress and strain (the relative deformation of a solid body due to a stress).

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Howard, J., Grill, S. & Bois, J. Turing's next steps: the mechanochemical basis of morphogenesis. Nat Rev Mol Cell Biol 12, 392–398 (2011).

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