Changes in epistatic green-beard alleles induce domain shift in hypostatic rock-paper-scissors-like green-beard competition

Myxobacteria with compatible TraA receptors can exchange materials located on the outer membrane. Such outer membrane exchange (OME) is an efficient strategy to help clonemates. If OME occurs between cells carrying different sitAI repertoires, however, transferred toxins may harm the recipient. While traA and sitAI can be categorized as green-beard genes, the functionality of sitAI is heavily reliant on traA. In other words, the harming effect of the sitAI is invalidated without OME. Hence, I termed traA and sitAI epistatic and hypostatic green-beard genes, respectively, and proposed a handshake-dependent rock-paper-scissors-like (RPSL) model. The simulation results imply that evolutionary pressure for traA diversification is context-dependent. The competition among aggressors, susceptibles, and freeloaders determined by hypostatic sitAI repertoire exhibits cyclic dominance, yet involves the possibility of divergent dynamics. By changing the epistatic green-beard traA alleles, susceptibles can avoid invasion of the aggressors, establishing another domain of the RPSL competition (domain shift). Furthermore, I mathematically analyzed the expected relatedness between myxobacteria from the compatibility of epistatic and hypostatic green-beard alleles. Not only can it explain the polymorphism of myxobacterial traA and sitAI alleles, but the model can also be expanded to investigate the social interactions of fusogenic cancer cells.