Reactive Oxygen Species Signaling Pathways: Arbiters of Evolutionary Conflict?

In the history of life, cooperation between biological units has led to increased complexity, e.g., eukaryotic cells and multicellular organisms. Cooperation requires limiting the gains of “defectors” in favor of the cooperative higher-level unit. Early in an evolutionary transition, bioenergetics and reactive oxygen species (ROS) may play a large role in managing these evolutionary conflicts. Chemiosmosis can be thought of as a poorly insulated wire—when supply exceeds demand, electrons are cast off and can form ROS. ROS signaling may thus lead to the dispersal of the excess products into the environment. These products may lead to groups and the formation of higher-level units that can subsequently be targeted by selection. Examining modern symbioses such as those between corals and photosynthetic dinoflagellates provides useful insight in this context. While ROS are an important factor in coral bleaching, little is known of the function of ROS under other circumstances, although some data suggest that ROS may modulate cooperation. ROS may have functioned similarly in the origin of eukaryotes, involving chemiosmotic mitochondria and chloroplasts. ROS may act as “arbiters” of evolutionary conflict, leading to cooperation via signaling pathways that favor the emergence of the higher-level unit.