Biofilms are communities of bacteria adhered to surfaces. Recently, biofilms
of rod-shaped bacteria were observed at single-cell resolution and shown to
develop from a disordered, two-dimensional layer of founder cells into a
three-dimensional structure with a vertically-aligned core. Here, we elucidate
the physical mechanism underpinning this transition using a combination of
agent-based and continuum modeling. We find that verticalization proceeds
through a series of localized mechanical instabilities on the cellular scale.
For short cells, these instabilities are primarily triggered by cell division,
whereas long cells are more likely to be peeled off the surface by nearby
vertical cells, creating an "inverse domino effect". The interplay between cell
growth and cell verticalization gives rise to an exotic mechanical state in
which the effective surface pressure becomes constant throughout the growing
core of the biofilm surface layer. This dynamical isobaricity determines the
expansion speed of a biofilm cluster and thereby governs how cells access the
third dimension. In particular, theory predicts that a longer average cell
length yields more rapidly expanding, flatter biofilms. We experimentally show
that such changes in biofilm development occur by exploiting chemicals that
modulate cell length.Comment: Main text 10 pages, 4 figures; Supplementary Information 35 pages, 15
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