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During development, organisms acquire three-dimensional shapes with importantphysiological consequences. While the basic mechanisms underlying morphogenesisare known in eukaryotes, it is often difficult to manipulate them in vivo. Tocircumvent this issue, here we present a study of developing Vibrio choleraebiofilms grown on agar substrates in which the spatiotemporal morphologicalpatterns were altered by varying the agar concentration. Expanding biofilms areinitially flat, but later experience a mechanical instability and becomewrinkled. Whereas the peripheral region develops ordered radial stripes, thecentral region acquires a zigzag herringbone-like wrinkle pattern. Depending onthe agar concentration, the wrinkles initially appear either in the peripheralregion and propagate inward (low agar concentration) or in the central regionand propagate outward (high agar concentration). To understand theseexperimental observations, we developed a model that considers diffusion ofnutrients and their uptake by bacteria, bacterial growth/biofilm matrixproduction, mechanical deformation of both the biofilm and the agar, and thefriction between them. Our model demonstrates that depletion of nutrientsbeneath the central region of the biofilm results in radially-dependent growthprofiles, which in turn, produce anisotropic stresses that dictate themorphology of wrinkles. Furthermore, we predict that increasing surfacefriction (agar concentration) reduces stress anisotropy and shifts the locationof the maximum compressive stress, where the wrinkling instability firstoccurs, toward the center of the biofilm, in agreement with our experimentalobservations. Our results are broadly applicable to bacterial biofilms withsimilar morphologies and also provide insight into how other bacterial biofilmsform distinct wrinkle patterns.

Non-uniform growth and surface friction determine bacterial biofilm morphology on soft substrates