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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 of the original microstructure is critical in setting the morphogenic dynamics of a pellicle as an active biomaterial. 

Cyanobacterial harmful algal blooms (CyanoHABs) are an increasingly common feature of large, eutrophic lakes. Non-N2-fixing CyanoHABs (e.g., Microcystis) appear to be proliferating relative to N2-fixing CyanoHABs in systems receiving increasing nutrient loads. This shift reflects increasing external nitrogen (N) inputs, and a[50-year legacy of excessive phosphorus (P) and N loading. Phosphorus is effectively retained in legacy-impacted systems, while N may be retained or lost to the atmosphere in gaseous forms (e.g., N2, NH3, N2O). Biological control on N inputs versus outputs, or the balance between N2 fixation versus denitrification, favors the latter, especially in lakes undergoing accelerating eutrophication, although denitrification removal efficiency is inhibited by increasing external N loads. Phytoplankton in eutrophic lakes have become more responsive to N inputs relative to P, despite sustained increases in N loading. From a nutrient management perspective, this suggests a need to change the freshwater nutrient limitation and input reduction paradigms; a shift from an exclusive focus on P limitation to a dual N and P colimitation and management strategy. The recent proliferation of toxic non-N2-fixing CyanoHABs, and ever-increasing N and P legacy stores, argues for such a strategy if we are to mitigate eutrophication and CyanoHAB expansion globally.