Coffee is one of the most important global crops and provides a livelihood to millions of people living in developing countries. Coffee species have been described as being highly sensitive to climate change, as largely deduced from modeling studies based on predictions of rising temperatures and changing rainfall patterns. Here, we discuss the physiological responses of the coffee tree in the context of present and ongoing climate changes, including drought, heat, and light stresses, and interactions between these factors. We also summarize recent insights on the physiological and agronomic performance of coffee at elevated atmospheric CO concentrations and highlight the key role of CO in mitigating the harmful effects of heat stress. Evidence is shown suggesting that warming, per se, may be less harmful to coffee suitability than previously estimated, at least under the conditions of an adequate water supply. Finally, we discuss several mitigation strategies to improve crop performance in a changing world.
The xylem cavitation threshold water potential establishes a hydraulic limit on the ability of woody species to survive in water-limiting environments, but herbs may be more plastic in terms of their ability to adapt to drying conditions. Here, we examined the capacity of sunflower (Helianthus annuus L.) leaves to adapt to reduced water availability by modifying the sensitivity of xylem and stomata to soil water deficit. We found that sunflower plants grown under water-limited conditions significantly adjusted leaf osmotic potential, which was linked to a prolongation of stomatal opening as soil dried and a reduced sensitivity of photosynthesis to water-stress-induced damage. At the same time, the vulnerability of midrib xylem to water-stress-induced cavitation was observed to be highly responsive to growth conditions, with water-limited plants producing conduits with thicker cell walls which were more resistant to xylem cavitation. Coordinated plasticity in osmotic potential and xylem vulnerability enabled water-limited sunflowers to safely extract water from the soil, while protecting leaf xylem against embolism. High plasticity in sunflower xylem contrasts with data from woody plants and may suggest an alternative strategy in herbs.
The driver of leaf mortality during drought stress is a critical unknown. We used the commercially important tree Persea americana, in which there is a large variation in the degree of drought-induced leaf death across the canopy, to test whether embolism formation in the xylem during drought drives this leaf mortality. A large range in the number of embolized vessels in the petioles of leaves was observed across the canopy of plants that had experienced drought. Despite considerable variation between leaves, the amount of embolized vessels in the xylem of the petiole strongly correlated with area of drought-induced tissue death in individual leaves. Consistent with this finding was a large interleaf variability in xylem resistance to embolism, with a 1.45 MPa variation in the water potential at which 50% of the xylem in the leaf midrib embolized across leaves. Our results implicate xylem embolism as a driver of leaf mortality during drought. Moreover, we propose that heterogeneity in drought-induced leaf mortality across a canopy is caused by high interleaf variability in xylem resistance to embolism, which may act as a buffer against complete canopy death during prolonged drought in P. americana.
Stomatal responses to changes in leaf water status are important for the diurnal regulation of gas exchange and the survival of plants during drought. These stomatal responses in angiosperm species are well characterized, yet in species of nonseed plants, an ongoing debate surrounds the role of metabolism, particularly the role of the hormone abscisic acid (ABA), in functionally regulating stomatal responses to changes in leaf water status. Here, we measured the stomatal response to changes in vapor pressure difference (VPD) in two natural forms of the fern species Athyrium filix-femina, recently suggested to have stomata that are regulated by ABA. The two forms measured had considerable differences in key hydraulic traits, including leaf hydraulic conductance and capacitance, as well as the kinetics of stomatal response to changes in VPD. In both forms, the stomatal responses to VPD could be accurately predicted by a dynamic, mechanistic model that assumes guard cell turgor changes in concert with leaf turgor in the light, and not via metabolic processes including the level of ABA. During drought, endogenous ABA did not play a role in stomatal closure, and exogenous ABA applied to live, intact leaves did not induce stomatal closure. Our results indicate that functional stomatal responses to changes in leaf water status in ferns are regulated by leaf hydraulics and not metabolism. With ferns being sister to seed plants, this result has implications for the evolutionary reconstruction of functional stomatal responses across vascular land plant lineages.
Rapid biosynthesis of abscisic acid (ABA) in the leaf, triggered by a decrease in cell volume, is essential for a functional stomatal response. However, it is not known whether rapid biosynthesis of ABA is also triggered in other plant tissues. Through the application of external pressure to flower, root, and leaf tissues, we test whether a reduction in cell volume can trigger rapid increases in ABA levels across the plant body in two species, Solanum lycopersicum and Passiflora tarminiana. Our results show that, in contrast to rapid ABA synthesis in the leaf, flower and root tissue did not show a significant, increase in ABA level in response to a drop in cell volume over a short time frame, suggesting that rapid ABA biosynthesis occurs only in leaf, not in flower or root tissues. A gene encoding the key, rate-limiting carotenoid cleavage enzyme (9-cis-epoxycarotenoid dioxygenase, NCED) in the ABA biosynthetic pathway in S. lycopersicum, NCED1, was upregulated to a lesser degree in flowers and roots compared with leaves in response to applied pressure. In both species, floral tissues contained substantially lower levels of the NCED substrate 9’-cis-neoxanthin than leaves, and this ABA precursor could not be detected in roots. Slow and minimal ABA biosynthesis was detected after 2 h in petals, indicating that floral tissue is capable of synthesizing ABA in response to sustained water deficit. Our results indicate that rapid ABA biosynthesis predominantly occurs in the leaves, and not in other tissues.
Investment in leaf veins (supplying xylem water) is balanced by stomatal abundance, such that sufficient water transport is provided for stomata to remain open when soil water is abundant. This coordination is mediated by a common dependence of vein and stomatal densities on cell size. Flowers may not conform to this same developmental pattern if they depend on water supplied by the phloem or have high rates of nonstomatal transpiration. We examined the relationships between veins, stomata and epidermal cells in leaves, sepals and petals of 27 angiosperms to determine whether common spacing rules applied to all tissues. Regression analysis found no evidence for different relationships within organ types. Both vein and stomatal densities were strongly associated with epidermal cell size within organs, but, for a given epidermal cell size, petals had fewer veins and stomata than sepals, which had fewer than leaves. Although our data support the concept of common scaling between veins and stomata in leaves and flowers, the large diversity in petal vein density suggests that, in some species, petal veins may be engaged in additional functions, such as the supply of water for high cuticular transpiration or for phloem delivery of water or carbohydrates.
Xylem resistance to embolism is a key metric determining plant survival during drought. Yet, we have a limited understanding of the degree of plasticity in vulnerability to embolism. Here, we tested whether light availability influences embolism resistance in leaves and stems. The optical vulnerability method was used to assess stem and leaf resistance to embolism in Phellodendron amurense and Ilex verticillata acclimated to sun and shade microenvironments within the same canopy. In both species, we found considerable segmentation in xylem resistance to embolism between leaves and stems, but only minor acclimation in response to light availability.With the addition of a third species, Betula pubescens, which shows no vulnerability segmentation, we sought to investigate xylem anatomical traits that might correlate with strong vulnerability segmentation. We found a correlation between the area fraction of vessels in the xylem and embolism resistance across species and tissue types. Our results suggest that minimal acclimation of embolism resistance occurs in response to light environment in the same individual and that the degree of vulnerability segmentation between leaves and stems might be determined by the vessel lumen fraction of the xylem.
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