Ethylene-regulated leaf lifespan explains divergent responses of plant productivity to warming among three hydrologically different growing seasons

Leaf senescence is known to be regulated by the plant hormone ethylene, but how leaf lifespan responds to global environmental change and links to ecosystem-level responses remains largely unexplored. Here we investigated the effects of climate warming and nitrogen addition on plant functional traits, plant hormone ethylene and net primary production in a 13-year field experiment in a desert steppe. Across the last 3 years of the experiment (2016-2018), plant productivity increased under warming only in 2016, when there was above normal precipitation, but consistently increased with nitrogen addition. Warming enhanced net photosynthesis, leaf nitrogen and ethylene production and reduced leaf lifespan in 2016 (a wet year), but not in 2017 (a drought year); the effect of warming in 2018 (a year with normal precipitation) was opposite to 2016, likely due to the below-normal precipitation in the mid-growing season in 2018. Nitrogen addition led to increases in leaf nitrogen, ethylene production and net photosynthesis, and declines in leaf lifespan in 2016 and 2018, but not in 2017. The ethylene-regulated lifespan was further evidenced by the addition of CoCl₂ (an ethylene biosynthesis inhibitor) that reduced ethylene production and prolonged lifespan. Structural equation modeling showed that leaf lifespan had a negative effect on plant productivity, both directly and indirectly via its negative effect on net photosynthesis, across all 3 years. Our results demonstrate the divergent responses of leaf lifespan and, in turn, plant productivity to warming under inter-annual and intra-annual precipitation variation, thus linking plant hormone production, functional traits and ecosystem functioning in the face of global environmental change.