Pattern Formation and Morphogenesis ( 1998 ) Modeling Branching and ChiralColonial Patterning

In nature, microorganisms must often cope with hostile environmental conditions. To do so they have developed sophisticated cooperative behavior and intricate communication capabilities, such as: direct cell-cell physical interactions via extra-membrane polymers, collective production of extracellular "wetting" uid for movement on hard surfaces, long range chemical signal-ing such as quorum sensing and chemotactic (bias of movement according to gradient of chemical agent) sig-naling, collective activation and deactivation of genes and even exchange of genetic material. Utilizing these 1 capabilities, the colonies develop complex spatio-temporal patterns in response to adverse growth conditions. We present a wealth of branching and chiral patterns formed during colonial development of lubricating bacteria (bacteria which produce a wetting layer of uid for their movement). Invoking ideas from pattern formation in non-living systems and using \generic" modeling we are able to reveal novel survival strategies which account for the salient features of the evolved patterns. Using the models, we demonstrate how communication leads to self-organization via cooperative behavior of the cells. In this regard, pattern formation in microorganisms can be viewed as the result of the exchange of information between the micro-level (the individual cells) and the macro-level (the colony). We mainly review known results , but include a new model of chiral growth, which enables us to study the eeect of chemotactic signaling on the chiral growth. We also introduce a measure for weak chirality and use this measure to compare the results of model simulations with experimental observations.