Plant life needs cell death, but does plant cell death need Cys proteases?

Caspases are key regulators of apoptosis in animals. This correlation has driven plant researchers for decades to look for caspases regulating programmed cell death (PCD) in plants. These studies revealed caspase-like activities, caspase-related proteases, and cysteine (Cys) proteases regulating PCD in plants, but identified no caspases and no conserved, apoptosis-like death pathway. Here, we critically review the evidence for Cys proteases implicated in PCD in plants. We discuss the role of papain-like Cys proteases, vacuolar processing enzymes, and metacaspases in PCD during the development of tracheary elements, seed coat, suspensor, and tapetum, and during the hypersensitive response. There are several convincing cases where these Cys proteases are required for PCD, but this requirement is often not conserved across different plant species. There are also cases where Cys proteases contribute to the speed, but not the timing of PCD, while other Cys proteases are nonessential for PCD, but have other roles, e.g., in the clearance of cell remains after PCD. These data illustrate the need for caution when generalizing the role of Cys proteases in regulating PCD in plants, and call for studies that further investigate plant Cys proteases and other PCD regulators.