Elevated CO2 did not stimulate stem growth in 11 provenances of a globally important hardwood plantation species

Elevated atmospheric carbon dioxide (eCO2) often enhances rates of photosynthesis leading to increased productivity in trees. In their native habitats in Australia, eucalypts display considerable phenotypic plasticity in response to changes in environmental conditions. Little is known whether this plasticity can be harnessed effectively under future atmospheric eCO2 conditions and be used to identify provenances with superior growth. Here, we report two experiments that assessed the physiological and growth responses of Eucalyptus grandis—one of the world's most important hardwood plantation species—to eCO2. We used 11 provenances from contrasting climates. Our selection was based on site-specific information of long-term temperature and water availability. In Experiment 1, four provenances exhibited significant variation in light-saturated photosynthetic rates (Asat), stomatal conductance (gs), and concentrations of non-structural carbohydrates in leaves, stems and roots when grown under ambient CO2 (aCO2). Biomass of leaves, stems and roots varied significantly and were negatively correlated with mean annual temperature (MAT) at seed origin, indicating that provenances from cooler, wetter climates generally produced greater biomass. Yet, stem growth of these provenances was not stimulated by eCO2. Given the vast environmental gradient covered by provenances of E. grandis, we expanded the selection from four to nine provenances in Experiment 2. This allowed us to validate results from Experiment 1 with its small selection and detailed measurements of various physiological parameters by focusing on growth responses to eCO2 across a wider environmental gradient in Experiment 2. In Experiment 2, nine provenances also exhibited intraspecific differences in growth, but these were not related to climate of origin, and eCO2 had little effect on growth traits. Growth responses under eCO2 varied widely across provenances in both experiments, confirming phenotypic plasticity in E. grandis, though responses were not systematically correlated with climate of origin. These results indicate that selection of provenances for improved stem growth of E. grandis under future eCO2 cannot be based solely on climate of origin, as is common practice for other planted tree species

Aridity drives clinal patterns in leaf traits and responsiveness to precipitation in a broadly distributed Australian tree species

Aridity shapes species distributions and plant growth and function worldwide. Yet, plant traits often show complex relationships with aridity, challenging our understanding of aridity as a driver of evolutionary adaptation. We grew nine genotypes of Eucalyptus camaldulensis subsp. camaldulensis sourced from an aridity gradient together in the field for ~650 days under low and high precipitation treatments. Eucalyptus camaldulesis is considered a phreatophyte (deep-rooted species that utilizes groundwater), so we hypothesized that genotypes from more arid environments would show lower aboveground productivity, higher leaf gas-exchange rates, and greater tolerance/avoidance of dry surface soils (indicated by lower responsiveness) than genotypes from less arid environments. Aridity predicted genotype responses to precipitation, with more arid genotypes showing lower responsiveness to reduced precipitation and dry surface conditions than less arid genotypes. Under low precipitation, genotype net photosynthesis and stomatal conductance increased with home-climate aridity. Across treatments, genotype intrinsic water-use efficiency and osmotic potential declined with increasing aridity while photosynthetic capacity (Rubisco carboxylation and RuBP regeneration) increased with aridity. The observed clinal patterns indicate that E. camaldulensis genotypes from extremely arid environments possess a unique strategy defined by lower responsiveness to dry surface soils, low water-use efficiency, and high photosynthetic capacity. This strategy could be underpinned by deep rooting and could be adaptive under arid conditions where heat avoidance is critical and water demand is high

Two measures of leaf capacitance : insights into the water transport pathway and hydraulic conductance in leaves

The efficiency and stress tolerance of leaf water transport are key indicators of plant function, but our ability to assess these processes is constrained by gaps in our understanding of the water transport pathway in leaves. A major challenge is to understand how different pools of water in leaves are connected to the transpiration stream and, hence, determine leaf capacitance (Cleaf) to short- and medium-term fluctuations in transpiration. Here, we examine variation across an anatomically and phylogenetically diverse group of woody angiosperms in two measures of Cleaf assumed to represent bulk-leaf capacitance (Cbulk) and the capacitance of leaf tissues that influence dynamic changes in leaf hydration (Cdyn). Among species, Cbulk was significantly correlated with leaf mass per unit area, whereas Cdyn was independently related to leaf lignin content (%) and the saturated mass of leaf water per unit dry weight. Dynamic and steady-state measurements of leaf hydraulic conductance (Kleaf) agreed if Cdyn was used rather than Cbulk, suggesting that the leaf tissue in some species is hydraulically compartmentalised and that only a proportion of total leaf water is hydraulically well connected to the transpiration stream. These results indicate that leaf rehydration kinetics can accurately measure Kleaf with knowledge of the capacitance of the hydraulic pathway

Leaf hydraulic vulnerability is related to conduit dimensions and drought resistance across a diverse range of woody angiosperms

Summary: Hydraulic dysfunction in leaves determines key aspects of whole-plant responses to water stress; however, our understanding of the physiology of hydraulic dysfunction and its relationships to leaf structure and ecological strategy remains incomplete. Here, we studied a morphologically and ecologically diverse sample of angiosperms to test whether the water potential inducing a 50% loss in leaf hydraulic conductance (P50leaf) is predicted by properties of leaf xylem relating to water tension-induced conduit collapse. We also assessed the relationships between P50leaf and other traits considered to reflect drought resistance and ecological strategy. Across species, P50leaf was strongly correlated with a theoretical predictor of vulnerability to cell collapse in minor veins (the cubed ratio of the conduit wall thickness to the conduit lumen breadth). P50leaf was also correlated with mesophyll traits known to be related to drought resistance, but unrelated to traits associated with carbon economy. Our data indicate a link between the structural mechanics of leaf xylem and hydraulic function under water stress. Although it is possible that collapse may contribute directly to dysfunction, this relationship may also be a secondary product of vascular economics, suggesting that leaf xylem is dimensioned to avoid wall collapse

Leaf hydraulic vulnerability influences species' bioclimatic limits in a diverse group of woody angiosperms

The ability of plants to maintain water flow through leaves under water stress-induced tension (assessed as the leaf hydraulic vulnerability; P50 leaf) is intimately linked with survival. We examined the significance of P50 leaf as an adaptive trait in influencing the dry-end distributional limits of cool temperate woody angiosperm species. We also examined differences in within-site variability in P50 leaf between two high-rainfall montane rainforest sites in Tasmania and Peru, respectively. A significant relationship between P50 leaf and the 5th percentile of mean annual rainfall across each species distribution was found in Tasmania, suggesting that P50 leaf influences species climatic limits. Furthermore, a strong correlation between P50 leaf and the minimum rainfall availability was found using five phylogenetically independent species pairs in wet and dry evergreen tree species, suggesting that rainfall is an important selective agent in the evolution of leaf hydraulic vulnerability. Greater within-site variability in P50 leaf was found among dominant montane rainforest species in Tasmania than in Peru and this result is discussed within the context of differences in spatial and temporal environmental heterogeneity and parochial historical ecology

Leaf hydraulics and drought stress : response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential (ψleaf), leaf hydraulic conductance (Kleaf), and midday transpiration (E) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of Kleaf to declining ψleaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and Kleaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of Kleaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to ψleaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery

Climate drives vein anatomy in Proteaceae

Premise of study: The mechanisms by which plants tolerate water defi cit are only just becoming clear. One key factor in drought tolerance is the ability to maintain the capacity to conduct water through the leaves in conditions of water stress. Recent work has shown that a simple feature of the leaf xylem cells, the cube of the thickness of cell walls divided by the lumen width (t/b) 3, is strongly correlated with this ability. • Methods: Using ecologically, phylogenetically, and anatomically diverse members of Proteaceae, we tested the relationships between (t/b) 3 and climate, leaf mass per unit area, leaf area, and vein density. To test relationships at high phylogenetic levels (mostly genus), we used phylogenetic and nonphylogenetic single and multiple regressions based on data from 50 species. We also used 14 within-genus species pairs to test for relationships at lower phylogenetic levels. • Key results: All analyses revealed that climate, especially mean annual precipitation, was the best predictor of (t/b) 3. The variation in (t/b) 3 was driven by variation in both lumen diameter and wall thickness, implying active control of these dimensions. Total vein density was weakly related to (t/b) 3 but unrelated to either leaf area or climate. Conclusions: We conclude that xylem reinforcement is a fundamental adaptation for water stress tolerance and, among evergreen woody plants, drives a strong association between rainfall and xylem anatomy. The strong association between (t/b) 3 and climate cannot be explained by autocorrelation with other aspects of leaf form and anatomy that vary along precipitation gradients

Genetic adaptation and phenotypic plasticity contribute to greater leaf hydraulic tolerance in response to drought in warmer climates

The ability of plants to maintain an intact water transport system in leaves under drought conditions is intimately linked to survival and can been be seen as adaptive in shaping species climatic limits. Large differences in leaf hydraulic vulnerability to drought are known among species from contrasting climates, yet whether this trait varies among populations within a single species and, furthermore, whether it is altered by changes in growth conditions, remain unclear. We examined intraspecific variation in both leaf water transport capacity (Kleaf) and leaf hydraulic vulnerability to drought (P50leaf) among eight populations of Corymbia calophylla (R. Br.) K.D. Hill & L.A.S. Johnson (Myrtaceae) from both cool and warm climatic regions grown reciprocally under two temperature treatments representing the cool and warm edge of the species distribution. Kleaf did not vary between cool and warm-climate populations, nor was it affected by variable growth temperature. In contrast, population origin and growth temperature independently altered P50leaf. Using data pooled across growth temperatures, cool-climate populations showed significantly higher leaf hydraulic vulnerability (P50leaf = −3.55 ± 0.18 MPa) than warm-climate populations (P50leaf = −3.78 ± 0.08 MPa). Across populations, P50leaf decreased as population home-climate temperature increased, but was unrelated to rainfall and aridity. For populations from both cool and warm climatic regions, P50leaf was lower under the warmer growth conditions. These results provide evidence of trait plasticity in leaf hydraulic vulnerability to drought in response to variable growth temperature. Furthermore, they suggest that climate, and in particular temperature, may be a strong selective force in shaping intraspecific variation in leaf hydraulic vulnerability to drought

Coordination between leaf, stem, and root hydraulics and gas exchange in three arid‐zone angiosperms during severe drought and recovery

The ability to resist hydraulic dysfunction in leaves, stems, and roots strongly influences whether plants survive and recover from drought. However, the coordination of hydraulic function among different organs within species and their links to gas exchange during drought and recovery remains understudied. Here, we examine the interaction between gas exchange and hydraulic function in the leaves, stems, and roots of three semiarid evergreen species exposed to a cycle of severe water stress (associated with substantial cavitation) and recovery. In all species, stomatal closure occurred at water potentials well before 50% loss of stem hydraulic conductance, while in two species, leaves and/or roots were more vulnerable than stems. Following soil rewetting, leaf‐level photosynthesis (Anet) returned to prestress levels within 2–4 weeks, whereas stomatal conductance and canopy transpiration were slower to recover. The recovery of Anet was decoupled from the recovery of leaf, stem, and root hydraulics, which remained impaired throughout the recovery period. Our results suggest that in addition to high embolism resistance, early stomatal closure and hydraulic vulnerability segmentation confers drought tolerance in these arid zone species. The lack of substantial embolism refilling within all major organs suggests that vulnerability of the vascular system to drought‐induced dysfunction is a defining trait for predicting postdrought recovery

Leaf hydraulic vulnerability to drought is linked to site water availability across a broad range of species and climates

Background and Aims Vulnerability of the leaf hydraulic pathway to water-stress-induced dysfunction is a key component of drought tolerance in plants and may be important in defining species' climatic range. However, the generality of the association between leaf hydraulic vulnerability and climate across species and sites remains to be tested. Methods Leaf hydraulic vulnerability to drought (P50leaf, thewater potential inducing 50%loss in hydraulic function) was measured in a diverse group of 92 woody, mostly evergreen angiosperms from sites across a wide range of habitats. These new data together with some previously published were tested against key climate indices related to water availability. Differences in within-site variability in P50leaf between sites were also examined. Key Results Values of hydraulic vulnerability to drought in leaves decreased strongly (i.e. became more negative) with decreasing annual rainfall and increasing aridity across sites. The standard deviation in P50leaf values recorded within each site was positively correlated with increasing aridity. P50leaf was also a good indicator of the climatic envelope across each species' distributional range as well as their dry-end distributional limits within Australia, although this relationship was not consistently detectable within sites. Conclusions The findings indicate that species sorting processes have influenced distributional patterns of P50leaf across the rainfall spectrum, but alternative strategies for dealing with water deficit exist within sites. The strong link to aridity suggests leaf hydraulic vulnerability may influence plant distributions under future climates