(ROS) play a key role in the regulation of many biological processes in plants. Nonetheless, they are considered highly reactive and toxic to cells. Owing to their toxicity, as well as their important role in signaling, the level of ROS in cells needs to be tightly regulated. The ROS gene network, encoding a highly redundant arsenal of ROS scavenging mechanisms and an array of enzymes involved in ROS production, regulates ROS metabolism and signaling in plants. In this article, we review the role of the ROS gene network in plants and examine how it evolved. We identify key components of the ROS gene network in organisms that likely originated as early as 4.1-3.5 billion years ago,f reactive oxygen species metabolism.