Popis: |
Plants rely on diverse strategies to regulate water loss during drought. The phytohormone abscisic acid (ABA) is a critical mediator of stomatal closure during water stress in seed plants. Studies in conifers identified diverging strategies in long-term drought of ABA-mediated dynamics, particularly a peaking type dynamic during long term drought in some conifers. Few studies have reported this dynamic in angiosperms, and no study has revealed the mechanism driving declines in ABA levels as drought progresses in peaking type species. To understand peaking type dynamics, we exposed the model peaking type gymnosperm species Callitris rhomboidea and the highly drought resistant evergreen angiosperm Umbellularia californica to controlled long-term drought. We measured leaf water potentials (��l), stomatal conductance, ABA and the ABA catabolite phaseic acid (PA) levels in potted plants during a prolonged but non-fatal drought. We aimed to determine which of three potential drivers of peaking type dynamic were responsible for this response: (1) increased catabolism of ABA into PA at a threshold ��l , (2) ABA export from the leaf is enhanced under drought, and (3) ABA biosynthesis ceases at a threshold ��l. During long term drought, the evergreen angiosperm species U. californica demonstrated peaking type ABA dynamics like gymnosperms. In both species, PA levels did not increase significantly, in fact, PA levels tracked ABA levels, suggesting that ABA catabolism to PA may be a function of ABA levels. Girdling experiments to determine whether export from the leaf drove declines in ABA levels demonstrated that of the majority of ABA was likely converted to ABA glucose ester (ABA-GE), an inactive storage form of ABA, and exported from shoots during drought. Finally, by rapidly dehydrating branched collected at different timepoints during long-term drought we were able to determine that ABA biosynthesis is completely down regulated in leaves that have been dehydrated beyond leaf turgor loss point. The decline in ABA levels in peaking type species appears conserved across seed plants and is mediated by high export rates in the form of ABA-GE. Future work should assess a more diverse selection of species as well as study long-term drought in less tolerant species to test whether ABA biosynthesis is deactivated in all species once ��l declines below turgor loss point. |