Patterns of PIP gene expression in Populus trichocarpa during recovery from xylem embolism suggest a major role for the PIP1 aquaporin subfamily as moderators of refilling process

Abstract

Embolism and the refilling of xylem vessels are intrinsic to the ability of plants to handle the transport of water under tension. Although the formation of an embolized vessel is an abiotic process, refilling against the pressure gradient requires biological activity to provide both the energy and the water needed to restore xylem transport capacity. Here, we present an analysis of the dynamics of embolism and refilling in Populus trichocarpa and follow temporal dynamics of co-occurring changes in expression level of aquaporins. Under mesic conditions, we found that the percent loss of conductance (PLC) varied diurnally by as much as 20%, suggesting a continuous embolism/refilling cycle. An increase in water stress tilted the balance between the two processes and increased the PLC to as much as 80%. Subsequent re-watering resulted in the reversal of water stress and recovery of PLC to pre-stress levels. Stem parenchyma cells responded to drought stress with considerable up-regulation of the PIP1 subfamily of water channels but not the PIP2 subfamily. Even more significant was the finding that PoptrPIP1.1 and PoptrPIP1.3 genes were up-regulated in response to embolism, but not to water stress, and were down-regulated after embolism removal, suggesting a local ability of plants to sense an embolism presence.