Loading regulation prevents phloem failure during drought and widens the
range of phloem and stomatal traits
- Ryan Stanfield,
- Megan Bartlett
Abstract
Plant carbon transport is controlled by a multitude of parameters both
internal and external to the sugar transporting phloem tissue. Sucrose
transporter kinetics, conduit hydraulic resistance, and xylem water
stress are all hypothesized to impact the amount of carbon delivered to
sink tissues. However, the most important traits determining carbon
export under drought are not well understood, especially for species
with active molecular regulation of sucrose transport. This in turn
limits our ability to assess species' resistances to phloem dysfunction
under drought. Here, we use an integrated xylem-phloem-stomatal model to
calculate leaf water potential from soil dryness, which is then used to
determine gas exchange and phloem pressure gradients. We quantitatively
compare the impacts of phloem loading kinetics, including feedbacks
between loading and phloem pressure, phloem conduit resistances, and
stomatal responses to water stress, on the total carbon export to sinks
during drought. Regulating sucrose transporter kinetics which
downregulates loading at high phloem pressures prevented runaway
viscosity in the phloem sap and was the most important determinant of
export rates under drought. In contrast to previous models, we found
this feedback mechanism decoupled stomatal traits from phloem export
efficiency during drought and increased the operational range of phloem
hydraulic resistances.