Hierarchical Transitions and Fractal Wrinkling Drive Bacterial Pellicle Morphogenesis
Bacterial cells can self-organize into structured communities at fluid-fluid interfaces. These soft, living materials composed of cells and extracellular matrix are called pellicles. Cells residing in pellicles garner group-level survival advantages such as increased antibiotic resistance. The dynamics of pellicle formation and, more generally, how complex morphologies arise from active biomaterials confined at interfaces are not well understood. Here, using Vibrio cholerae as our model organism, a custom-built adaptive stereo microscope, fluorescence imaging, mechanical theory, and simulations, we report a fractal wrinkling morphogenesis program that differs radically from the well-known coalescence of wrinkles into folds that occurs in passive thin films at fluid-fluid interfaces. Four stages occur: growth of founding colonies, onset of primary wrinkles, development of secondary curved ridge instabilities, and finally the emergence of a cascade of finer structures with fractal-like scaling in wavelength. The time evolution of pellicle formation depends on the initial heterogeneity of the film microstructure. Changing the starting bacterial seeding density produces three variations in the sequence of morphogenic stages, which we term the bypass, crystalline, and incomplete modes. Despite these global architectural transitions, individual microcolonies remain spatially segregated, and thus the community maintains spatial and genetic heterogeneity. Our results suggest that the memory more »
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NSF-PAR ID:
10220721
Journal Name:
Proceedings of the National Academy of Sciences of the United States of America
ISSN:
1091-6490
3. Transition from laminar to turbulent flow occurring over a smooth surface is a particularly important route to chaos in fluid dynamics. It often occurs via sporadic inception of spatially localized patches (spots) of turbulence that grow and merge downstream to become the fully turbulent boundary layer. A long-standing question has been whether these incipient spots already contain properties of high-Reynolds-number, developed turbulence. In this study, the question is posed for geometric scaling properties of the interface separating turbulence within the spots from the outer flow. For high-Reynolds-number turbulence, such interfaces are known to display fractal scaling laws with a dimensionmore », where the 1/3 excess exponent above 2 (smooth surfaces) follows from Kolmogorov scaling of velocity fluctuations. The data used in this study are from a direct numerical simulation, and the spot boundaries (interfaces) are determined by using an unsupervised machine-learning method that can identify such interfaces without setting arbitrary thresholds. Wide separation between small and large scales during transition is provided by the large range of spot volumes, enabling accurate measurements of the volume–area fractal scaling exponent. Measurements show a dimension of$D=2.36±0.03$over almost 5 decades of spot volume, i.e., trends fully consistent with high-Reynolds-number turbulence. Additional observations pertaining to the dependence on height above the surface are also presented. Results provide evidence that turbulent spots exhibit high-Reynolds-number fractal-scaling properties already during early transitional and nonisotropic stages of the flow evolution.