Phototaxis as a Collective Phenomenon in Cyanobacterial Colonies

Cyanobacteria are a widely distributed, diverse group of photosynthetic bacteria that exhibit phototaxis, or motion in response to light. Cyanobacteria such as Synechocystis sp. secrete a mixture of complex polysaccharides that facilitate cell motion, while their type 4 pili allow them to physically attach to each other. Even though cells can respond individually to light, colonies of such bacteria are observed to move collectively towards the light source in dense finger-like projections. Agent-based models are especially useful in connecting individual cell behaviour with the emergent collective phenomena that arise out of their interactions. We present an agent-based model for cyanobacterial phototaxis that accounts for slime deposition as well as for direct physical links between bacteria, mediated through their type 4 pili. We reproduce the experimentally observed aggregation of cells at the colony boundary as a precursor to finger formation. Our model also describes the changes in colony morphology that occur when the location of the light source is abruptly changed. We find that the overall motion of cells toward light remains relatively unimpaired even if a fraction of them do not sense light, allowing heterogeneous populations to continue to mount a robust collective response to stimuli. Our work suggests that in addition to bio-chemical signalling via diffusible molecules in the context of bacterial quorum-sensing, short-ranged physical interactions may also contribute to collective effects in bacterial motility.