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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. 

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 The cause of reduced leaf-level transpiration under elevated CO2 remains largely elusive. Here, we assessed stomatal, hydraulic, and morphological adjustments in a long-term experiment on Aleppo pine (Pinus halepensis) seedlings germinated and grown for 22–40 months under elevated (eCO2; c. 860 ppm) or ambient (aCO2; c. 410 ppm) CO2. We assessed if eCO2-triggered reductions in canopy conductance (gc) alter the response to soil or atmospheric drought and are reversible or lasting due to anatomical adjustments by exposing eCO2 seedlings to decreasing [CO2]. To quantify underlying mechanisms, we analyzed leaf abscisic acid (ABA) level, stomatal and leaf morphology, xylem structure, hydraulic efficiency, and hydraulic safety. Effects of eCO2 manifested in a strong reduction in leaf-level gc (−55%) not caused by ABA and not reversible under low CO2 (c. 200 ppm). Stomatal development and size were unchanged, while stomatal density increased (+18%). An increased vein-to-epidermis distance (+65%) suggested a larger leaf resistance to water flow. This was supported by anatomical adjustments of branch xylem having smaller conduits (−8%) and lower conduit lumen fraction (−11%), which resulted in a lower specific conductivity (−19%) and leaf-specific conductivity (−34%). These adaptations to CO2 did not change stomatal sensitivity to soil or atmospheric drought, consistent with similar xylem safety thresholds. In summary, we found reductions of gc under elevated CO2 to be reflected in anatomical adjustments and decreases in hydraulic conductivity. As these water savings were largely annulled by increases in leaf biomass, we do not expect alleviation of drought stress in a high CO2 atmosphere. 

Abstract The resistance of xylem conduits to embolism is a major factor defining drought tolerance and can set the distributional limits of species across rainfall gradients. Recent work suggests that the proximity of vessels to neighbors increases the vulnerability of a conduit. We therefore investigated whether the relative vessel area of xylem correlates with intra- and inter-generic variation in xylem embolism resistance in species pairs or triplets from the genera Acer, Cinnamomum, Ilex, Quercus and Persea, adapted to environments differing in aridity. We used the optical vulnerability method to assess embolism resistance in stems and conducted anatomical measurements on the xylem in which embolism resistance was quantified. Vessel lumen fraction (VLF) correlated with xylem embolism resistance across and within genera. A low VLF likely increases the resistance to gas movement between conduits, by diffusion or advection, whereas a high VLF enhances gas transport thorough increased conduit-to-conduit connectivity and reduced distances between conduits and therefore the likelihood of embolism propagation. We suggest that the rate of gas movement due to local pressure differences and xylem network connectivity is a central driver of embolism propagation in angiosperm vessels. 

Maintaining water transport in the xylem is critical for vascular plants to grow and survive. The drought-induced accumulation of embolism, when gas enters xylem conduits, causes declines in hydraulic conductance (K) and is ultimately lethal. Several methods can be used to estimate the degree of embolism in xylem, from measuring K in tissues to directly visualising embolism in conduits. One method allowing a direct quantification of embolised xylem area is the optical vulnerability (OV) technique. This method has been used across different organs and has a high spatial and temporal resolution. Here, we review studies using the OV technique, discuss the main advantages and disadvantages of this method, and summarise key advances arising from its use. Vulnerability curves generated by the OV method are regularly comparable to other methods, including the centrifuge and X-ray microtomography. A major advantage of the OV technique over other methods is that it can be simultaneously used to determine in situ embolism formation in leaves, stems and roots, in species spanning the phylogeny of land plants. The OV method has been used to experimentally investigate the spreading of embolism through xylem networks, associate embolism with downstream tissue death, and observe embolism formation in the field.