Widespread biochemical reaction networks enable Turing patterns without imposed feedback

Understanding self-organized pattern formation is fundamental to biology. In 1952, Alan Turing proposed a pattern-enabling mechanism in reaction-diffusion systems containing chemical species later conceptualized as activators and inhibitors that are involved in feedback loops. However, identifying pattern-enabling regulatory systems with the concept of feedback loops has been a long-standing challenge. To date, very few pattern-enabling circuits have been discovered experimentally. This is in stark contrast to ubiquitous periodic patterns and symmetry in biology. In this work, we systematically study Turing patterns in 23 elementary biochemical networks without assigning any activator or inhibitor. These mass action models describe post-synthesis interactions applicable to most proteins and RNAs in multicellular organisms. Strikingly, we find ten simple reaction networks capable of generating Turing patterns. While these network models are consistent with Turing's theory mathematically, there is no apparent connection between them and commonly used activator-feedback intuition. Instead, we identify a unifying network motif that enables Turing patterns via regulated degradation pathways with flexible diffusion rate constants of individual molecules. Our work reveals widespread biochemical systems for pattern formation, and it provides an alternative approach to tackle the challenge of identifying pattern-enabling biological systems.