Search for: All records

Abstract By regulating carbon uptake and water loss by plants, stomata are not only responsible for productivity but also survival during drought. The timing of the onset of stomatal closure is crucial for preventing excessive water loss during drought, but is poorly explained by plant hydraulics alone and what triggers stomatal closure remains disputed. We investigated whether the hormone abscisic acid (ABA) was this trigger in a highly embolism‐resistant tree speciesUmbellularia californica. We tracked leaf ABA levels, determined the leaf water potential and gravimetric soil water content (gSWC) thresholds for stomatal closure and transpiration decline during a progressive drought. We found thatU. californicaplants have a peaking‐type ABA dynamic, where ABA levels rise early in drought and then decline under prolonged drought conditions. The early increase in ABA levels correlated with the closing of stomata and reduced transpiration. Furthermore, we found that transpiration declined before any large decreases in predawn plant water status and could best be explained by transient drops in midday water potentials triggering increased ABA levels. Our results indicate that ABA‐mediated stomatal regulation may be an integral mechanism for reducing transpiration during drought before major drops in bulk soil and plant water status. 

Summary The onset of stomatal closure reduces transpiration during drought. In seed plants, drought causes declines in plant water status which increases leaf endogenous abscisic acid (ABA) levels required for stomatal closure. There are multiple possible points of increased belowground resistance in the soil–plant atmospheric continuum that could decrease leaf water potential enough to trigger ABA production and the subsequent decreases in transpiration.We investigate the dynamic patterns of leaf ABA levels, plant hydraulic conductance and the point of failure in the soil–plant conductance in the highly embolism‐resistant speciesCallitris tuberculatausing continuous dendrometer measurements of leaf water potential during drought.We show that decreases in transpiration and ABA biosynthesis begin before any permanent decreases in predawn water potential, collapse in soil–plant hydraulic pathway and xylem embolism spread.We find that a dynamic but recoverable increases in hydraulic resistance in the soil in close proximity to the roots is the most likely driver of declines in midday leaf water potential needed for ABA biosynthesis and the onset of decreases in transpiration. 

Abstract Stomatal opening in the light, observed in nearly all vascular land plants, is essential for providing access to atmospheric CO2 for photosynthesis. The speed of stomatal opening in the light is critical for maximizing carbon gain in environments in which light intensity changes, yet we have little understanding of how other environmental signals, particularly evaporative demand driven by vapor pressure deficit (VPD) influences the kinetics of this response. In angiosperms, and some fern species from the family Marsileaceae, a mechanical interaction between the guard cells and the epidermal cells determines the aperture of the pore. Here, we examine whether this mechanical interaction influences the speed of stomatal opening in the light. To test this, we investigated the speed of stomatal opening in response to light across a range of VPDs in seven plant species spanning the evolutionary diversity of guard cell and epidermal cell mechanical interactions. We found that stomatal opening speed is a function of evaporative demand in angiosperm species and Marsilea, which have guard cell and epidermal cell mechanical interactions. Stomatal opening speeds did not change across a range of VPD in species of gymnosperm and fern, which do not have guard cell mechanical interactions with the epidermis. We find that guard cell and epidermal cell mechanical interactions may play a key role in regulating stomatal responsiveness to light. These results provide valuable insight into the adaptive relevance of mechanical advantage. 

Abstract The phytohormone abscisic acid (ABA) is synthesised by plants during drought to close stomata and regulate desiccation tolerance pathways. Conifers and some angiosperms with embolism‐resistant xylem show a peaking‐type (p‐type) response in ABA levels, in which ABA levels increase early in drought then decrease as drought progresses, declining to pre‐stressed levels. The mechanism behind this dynamic remains unknown. Here, we sought to characterise the mechanism driving p‐type ABA dynamics in the coniferCallitris rhomboideaand the highly drought‐resistant angiospermUmbellularia californica. We measured leaf water potentials (Ψ_l), stomatal conductance, ABA, conjugates and phaseic acid (PA) levels in potted plants during a prolonged but non‐fatal drought. Both species displayed a p‐type ABA dynamic during prolonged drought. In branches collected before and after the peak in endogenous ABA levels in planta, that were rehydrated overnight and then bench dried, ABA biosynthesis was deactivated beyond leaf turgor loss point. Considerable conversion of ABA to conjugates was found to occur during drought, but not catabolism to PA. The mechanism driving the decline in ABA levels in p‐type species may be conserved across embolism‐resistant seed plants and is mediated by sustained conjugation of ABA and the deactivation of ABA accumulation asΨ_lbecomes more negative than turgor loss. 

Abstract Vapor pressure difference between the leaf and atmosphere (VPD) is the most important regulator of daytime transpiration, yet the mechanism driving stomatal responses to an increase in VPD in angiosperms remains unresolved. Here, we sought to characterize the mechanism driving stomatal closure at high VPD in an angiosperm species, particularly testing whether abscisic acid (ABA) biosynthesis could explain the observation of a trigger point for stomatal sensitivity to an increase in VPD. We tracked leaf gas exchange and modeled leaf water potential (Ψl) in leaves exposed to a range of step-increases in VPD in the herbaceous species Senecio minimus Poir. (Asteraceae). We found that mild increases in VPD in this species did not induce stomatal closure because modeled Ψl did not decline below a threshold close to turgor loss point (Ψtlp), but when leaves were exposed to a large increase in VPD, stomata closed as modeled Ψl declined below Ψtlp. Leaf ABA levels were higher in leaves exposed to a step-increase in VPD that caused Ψl to transiently decline below Ψtlp and in which stomata closed compared with leaves in which stomata did not close. We conclude that the stomata of S. minimus are insensitive to VPD until Ψl declines to a threshold that triggers the biosynthesis of ABA and that this mechanism might be common to angiosperms. 

Abstract The phytohormone abscisic acid (ABA) plays a major role in closing the stomata of angiosperms. However, recent reports of some angiosperm species having a peaking-type ABA dynamic, in which under extreme drought ABA levels decline to pre-stressed levels, raises the possibility that passive stomatal closure by leaf water status alone can occur in species from this lineage. To test this hypothesis, we conducted instantaneous rehydration experiments in the peaking-type species Umbellularia californica through a long-term drought, in which ABA levels declined to pre-stress levels, yet stomata remain closed. We found that when ABA levels were lowest during extreme drought, stomata reopen rapidly to maximum rates of gas exchange on instantaneous rehydration, suggesting that the stomata of U. californica were passively closed by leaf water status alone. This contrasts with leaves early in drought, in which ABA levels were highest and stomata did not reopen on instantaneous rehydration. The transition from ABA-driven stomatal closure to passively driven stomatal closure as drought progresses in this species occurs at very low water potentials facilitated by highly embolism-resistant xylem. These results have important implications for understanding stomatal control during drought in angiosperms. 

Summary The hydraulic system of vascular plants and its integrity is essential for plant survival. To transport water under tension, the walls of xylem conduits must approximate rigid pipes. Against this expectation, conduit deformation has been reported in the leaves of a few species and hypothesized to function as a ‘circuit breaker’ against embolism. Experimental evidence is lacking, and its generality is unknown.We demonstrated the role of conduit deformation in protecting the upstream xylem from embolism through experiments on three species and surveyed a diverse selection of vascular plants for conduit deformation in leaves.Conduit deformation in minor veins occurred before embolism during slow dehydration. When leaves were exposed to transient increases in transpiration, conduit deformation was accompanied by large water potential differences from leaf to stem and minimal embolism in the upstream xylem. In the three species tested, collapsible vein endings provided clear protection of upstream xylem from embolism during transient increases in transpiration.We found conduit deformation in diverse vascular plants, including 11 eudicots, ginkgo, a cycad, a fern, a bamboo, and a grass species, but not in two bamboo and a palm species, demonstrating that the potential for ‘circuit breaker’ functionality may be widespread across vascular plants.