We present experimental results of colony formation in bacteria as an example of pattern formation resulting from reproduction and movement in biological populations. The bacterium Bacillus subtilis is known to exhibit at least five distinct types of colony pattern, depending on the substrate softness and nutrient concentration: diffusion-limited aggregation (DLA), compact Eden-like, dense branching morphology (DBM), concentric ring-like, and disk. We established a morphological diagram of the colony patterns, and then examined and characterized both macroscopically and microscopically how the the colonies grow. There seem to be two kinds of bacterial cells – active and inactive – and the active form drives the colony interfaces outwards. The active cells may be clearly distinguished from the inactive ones as they form the characteristic fingernail-like structure at the tips of growing branches of the DBM colony. The concentric ring-like colony is formed as a consequence of repeated alternate migration and resting of the growing interface, the cycle time for which seems to be independent of the substrate softness and nutrient concentration. So far there have been several phenomenological models proposed to qualitatively explain or reproduce the patterns observed in bacterial colonies. A few of them are reviewed here, systematically and critically, in light of our experimental results.

The formation of concentric ring-like colonies by bacterial species Bacillus subtilis has been investigated, focusing our attention on the effect of local cell density upon the bacterial motility: (i) Neither any chemicals nor a pacemaker at the center of the ring takes part in the concentric ring formation. (ii) Phase entrainment between two colonies having different phase of concentric ring does not occur. (iii) From the measurement of lag-phase time when varying the initial cell density, the start of the first migration phase is found to depend on the cell density. (iv) When cutting the part of a colony which is behind a migration phase just after the start of migration, the migration phase becomes shorter. On the other hand, the following consolidation phase becomes longer. (v) By the replica-printing method, active bacteria move collectively from inside to outside of the outermost consolidation terrace. Our present experimental results are qualitatively consistent with the results of the other bacterial species P. mirabilis, although the individual cell motility is quite different from each other. The present results suggest that the essential factor of the change of the bacterial motility of B. subtilis during concetric ring formation is the local cell density.

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Characteristic patterns of expansion were generated by bacterial cell populations of Serratia marcescens on media with different concentrations of nutrient and agar. These patterns were classified as Eden-like, densebranching-morphology-like, flower-like, concentric ring-like and diffusion-limited aggregation-like. Although flower-like was specific to Serratia marcescens, the other patterns were exhibited by Bacillus subtilis. Through macro-and microscopic tracing of the processes generating these patterns, physico-chemical principles of bacterial growth, collaborative and independent properties of bacteria, structural organization for population expansion, and the division of labor among bacterial cells (i.e., wall composer, pressure generator, and logistic supporter) were brought to light.