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Abstract PremiseHydraulic segmentation, caused by the difference in embolism resistance across plant organs, provides a sacrificial layer of cheaper plant organs, like leaves, to protect more costly organs, such as stems, during drought. Within‐leaf hydraulic segmentation has been observed in two compound‐leaved tree species, with leaflets being more vulnerable than the rachis or petiole. Many herbaceous species have compound leaves, and some species have leaflets that are associated with pulvini at the base of the lamina, which could provide an anatomical means of preventing embolism from spreading within a leaf because of the higher number of vessel endings in the pulvinus. MethodsWe used the optical vulnerability method to investigate whether differences in embolism resistance were observed across the leaf tissues of six herbaceous species and one deciduous tree species with compound leaves. Our species selection included both palmately and pinnately‐compound leaved species, one of each with a pulvinus at the base of the leaflets. ResultsWe found considerable variation in embolism resistance across the species measured, but no evidence of variation in embolism resistance within the leaf. In two species with pulvini, we observed major embolism events crossing the pulvinus, spreading from the rachis or petiole into the lamina, and embolizing both tissues at the same water potential. ConclusionsWe conclude that within‐leaf hydraulic segmentation, caused by variation in embolism resistance, is not a universal phenomenon to compound‐leaved species and that the presence of a pulvinus does not provide a barrier to embolism spread in compound leaves. 

ABSTRACT Freezing air temperatures kill most leaves, yet the leaves of some species can survive these events. Tracking the temporal and spatial dynamics of freezing remains an impediment to characterizing frost tolerance. Here we deploye time‐lapse imaging and image subtraction analysis, coupled with fine wire thermocouples, to discern the in situ spatial dynamics of freezing and thawing. Our method of analysis of pixel brightness reveals that ice formation in leaves exposed to natural frosts initiates in mesophyll before spreading to veins, and that while ex situ xylem sap freezes near 0°C, in situ xylem sap has a freezing point of −2°C in our model freezing‐resistant species ofLonicera. Photosynthetic rates in leaves that have been exposed to a rapid freeze or thaw do not recover, but leaves exposed to a slow, natural freezing and thawing to −10°C do recover. Using this method, we are able to quantify the spatial formation and timing of freezing events in leaves, and suggest that in situ and ex situ freezing points for xylem sap can differ by more than 4°C depending on the rate of temperature decline. 

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 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 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. 

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 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 Senescence vividly marks the onset of the final stages of the life of a leaf, yet the triggers and drivers of this process are still not fully understood. The hormone abscisic acid (ABA) is an important regulator of leaf senescence in model herbs, but the function of this hormone has not been widely tested in deciduous trees. Here we investigate the importance of ABA as a driver of leaf senescence in winter deciduous trees. In four diverse species we tracked leaf gas exchange, water potential, chlorophyll content, and leaf ABA levels from the end of summer until leaves were abscised or died. We found that no change in ABA levels occurred at the onset of chlorophyll decline or throughout the duration of leaf senescence. To test whether ABA could enhance leaf senescence, we girdled branches to disrupt ABA export in the phloem. Girdling increased leaf ABA levels in two of the species, and this increase triggered an accelerated rate of chlorophyll decline in these species. We conclude that an increase in ABA level may augment leaf senescence in winter deciduous species but that it is not essential for this annual process. 

Abstract Drought resistance is essential for plant production under water‐limiting environments. Abscisic acid (ABA) plays a critical role in stomata but its impact on hydraulic function beyond the stomata is far less studied. We selected genotypes differing in their ability to accumulate ABA to investigate its role in drought‐induced dysfunction. All genotypes exhibited similar leaf and stem embolism resistance regardless of differences in ABA levels. Their leaf hydraulic resistance was also similar. Differences were only observed between the two extreme genotypes:sitiens(sit; a strong ABA‐deficient mutant) andsp12(a transgenic line that constitutively overaccumulates ABA), where the water potential inducing 50% embolism was 0.25 MPa lower insp12than insit. Maximum stomatal and minimum leaf conductances were considerably lower in plants with higher ABA (wild type [WT] andsp12) than in ABA‐deficient mutants. Variations in gas exchange across genotypes were associated with ABA levels and differences in stomatal density and size. The lower water loss in plants with higher ABA meant that lethal water potentials associated with embolism occurred later during drought insp12plants, followed by WT, and then by the ABA‐deficient mutants. Therefore, the primary pathway by which ABA enhances drought resistance is via declines in water loss, which delays dehydration and hydraulic dysfunction.