Soluble sugar drives plant leaf turgor, restricts plant growth, and regulates plant hydraulic capacity of Haloxylon ammodendron to different groundwater level

Anthropogenic withdraw of groundwater and climatic drought results in the decline of groundwater depth that, in turn, severely limits the water availability for phreatophytic vegetation in arid regions. In this study, a small xeric, phreatophytic tree Haloxylon ammodendron (C.A. Mey.) was investigated to understand the influence of depth to groundwater (DGW) on hydraulic traits and on the trade-off between drought tolerance and leaf area increment. A suite of traits including leaf water potential, pressure–volume (P–V) curves, Huber value, assimilation branch growth, and osmotic regulation substance were measured across five sites with DGW ranges from 3.45 to 15.91 m. Our results indicate that H. ammodendron was subject to greater water stress with increasing DGW, as indicated by decreased predawn (Ψpd) & midday (Ψmd) branch water potential. We also found that growth rate declined as Huber value increased with increasing DGW in the early growing season (EGS). Solute sugar, as a major osmotic substance, drives decreases in osmotic potential at full turgor, and thus constrains assimilation branch growth with increasing DGW in EGS. Therefore, osmotic adjustment accompanied with water potential regulation (Ψpd－Ψmd) and plasticity of Huber value allows this phreatophyte to absorb water from deeper soil layers and tolerate drought. However, these adaptive adjustments cannot fully compensate for nonoptimal water conditions as growth rate continued to decrease as DGW increased in EGS and even became negative in the late growing season (LGS) at almost all sites. Our results provide an insight into how H. ammodendron responds and adapts to changes DGW in a region experiencing hydrological and climatic drought. Greater depth of groundwater had a significant effect on H. ammodendron and may have similar effects for other non-riparian phreatophytic plants in arid regions.

Plant hydraulics coordinated with photosynthetic traits and climate

The coupling between water loss and carbon dioxide uptake drives the coordination of plant hydraulic and photosynthetic traits. Analysing multi-species measurements on a 3000 m elevation gradient, we found that hydraulic and leaf-economic traits were less plastic, and more closely associated with phylogeny, than photosynthetic traits. The two trait sets are linked by the sapwood-to-leaf area ratio (Huber value, vH), shown here to be codetermined by sapwood hydraulic conductance (KS), leaf mass-per-area (LMA) and photosynthetic capacity (Vcmax). Substantial hydraulic diversity was related to the trade-off between KS and vH. Leaf drought tolerance (inferred from turgor loss point, –πtlp) increased with wood density, but the trade-off between hydraulic efficiency (KS) and –πtlp was weak. The least-cost optimality framework was extended to predict trait (KS-dominated) and environmental (temperature-dominated) effects on vH. These results suggest an approach to include photosynthetic-hydraulic coordination in land-surface models; however, prediction of non-plastic trait distributions remains a challenge.

Vulnerability to xylem cavitation of Hakea species (Proteaceae) from a range of biomes and life histories predicted by climatic niche

Background and Aims Extreme drought conditions across the globe are impacting biodiversity, with serious implications for the persistence of native species. However, quantitative data on physiological tolerance are not available for diverse flora to inform conservation management. We quantified physiological resistance to cavitation in the diverse Hakea genus (Proteaceae) to test predictions based on climatic origin, life history and functional traits. Methods We sampled terminal branches of replicate plants of 16 species in a common garden. Xylem cavitation was induced in branches under varying water potentials (tension) in a centrifuge, and the tension generating 50 % loss of conductivity (stem P50) was characterized as a metric for cavitation resistance. The same branches were used to estimate plant functional traits, including wood density, specific leaf area and Huber value (sap flow area to leaf area ratio). Key Results There was significant variation in stem P50 among species, which was negatively associated with the species climate origin (rainfall and aridity). Cavitation resistance did not differ among life histories; however, a drought avoidance strategy with terete leaf form and greater Huber value may be important for species to colonize and persist in the arid biome. Conclusions This study highlights climate (rainfall and aridity), rather than life history and functional traits, as the key predictor of variation in cavitation resistance (stem P50). Rainfall for species origin was the best predictor of cavitation resistance, explaining variation in stem P50, which appears to be a major determinant of species distribution. This study also indicates that stem P50 is an adaptive trait, genetically determined, and hence reliable and robust for predicting species vulnerability to climate change. Our findings will contribute to future prediction of species vulnerability to drought and adaptive management under climate change.

Drought tolerance of Hakea species (Proteaceae) from a range of biomes and life-histories predicted by climatic niche

Extreme drought conditions across the globe are impacting biodiversity with serious implications for the persistence of native species. However, quantitative data on drought tolerance is not available for diverse flora to inform conservation management. We quantified physiological drought tolerance in the diverse Hakea genus (Proteaceae) to test predictions based on climatic-origin, life history and functional traits. We sampled terminal branches of replicate plants of 16 species in a common garden. Xylem cavitation was induced in branches under varying water potential (tension) in a centrifuge and the tension generating 50% loss of conductivity (stem P50) was characterized as a metric for drought tolerance. The same branches were used to estimate plant functional traits, including wood density, specific leaf area, and Huber value (sap flow area to leaf area ratio). There was significant variation in stem P50 among species, which was negatively associated with the species climate-origin (rainfall and aridity). Drought tolerance did not differ among life histories; however, a drought avoidance strategy with terete leaf form and greater Huber value may be important for species to colonize and persist in the arid biome. Our findings will contribute to future prediction of species vulnerability to drought and adaptive management under climate change.

Hydraulic traits vary as the result of tip-to-base conduit widening in vascular plants

Plant hydraulic traits are essential metrics for characterizing variation in plant function, but they vary markedly with plant size and position in a plant. We explore the potential effect of conduit widening on variation in hydraulic traits along the stem. We examined three species that differ in conduit diameter at the stem base for a given height (Moringa oleifera, Casimiroa edulis, and Pinus ayacahuite). We made anatomical and hydraulic measurements at different distances from the stem tip, constructed vulnerability curves, and examined the safety–efficiency trade-off with height-standardized data. Our results showed that segment-specific hydraulic resistance varied predictably along the stem, paralleling changes in mean conduit diameter and total number of conduits. The Huber value and leaf specific conductivity also varied depending on the sampling point. Vulnerability curves were markedly less noisy with height standardization, making the vulnerability–efficiency trade-off clearer. Because conduits widen predictably along the stem, taking height and distance from the tip into account provides a way of enhancing comparability and interpretation of hydraulic traits. Our results suggest the need for rethinking hydraulic sampling for comparing plant functional differences and strategies across individuals.

Aridity drives coordinated trait shifts but not decreased trait variance across the geographic range of eight Australian trees

ContextLarge intraspecific functional trait variation strongly impacts many aspects of natural communities and ecosystems, yet is inconsistent across traits and species.ApproachWe measured within-species variation in leaf mass per area (LMA), leaf dry matter content (LDMC), branch wood density (WD), and allocation to stem area vs. leaf area in branches (branch Huber value, HV) across the aridity range of seven Australian eucalypts and an Acacia species to explore how traits and their variances change with aridity.Results and ConclusionsWithin-species, we found consistent increases in LMA, LDMC and WD, and HV with increasing aridity, resulting in consistent trait coordination across tissues. However, this coordination only emerged across sites with large climate differences. Unlike trait means, patterns of trait variance with aridity were mixed across populations and species and showed limited support for constrained trait variation in dryer populations or more xeric species.SynthesisOur results highlight that climate can drive consistent within-species trait patterns, but that these patterns might often be obscured by the complex nature of morphological traits and sampling incomplete species ranges or sampling confounded stress gradients.

Phenotypic plasticity and genetic adaptation of functional traits influences intra-specific variation in hydraulic efficiency and safety

Understanding which hydraulic traits are under genetic control and/or are phenotypically plastic is essential in understanding how tree species will respond to rapid shifts in climate. We quantified hydraulic traits in Eucalyptus obliqua L'Her. across a precipitation gradient in the field to describe (i) trait variation in relation to long-term climate and (ii) the short-term (seasonal) ability of traits to adjust (i.e., phenotypic plasticity). Seedlings from each field population were raised under controlled conditions to assess (iii) which traits are under strong genetic control. In the field, drier populations had smaller leaves with anatomically thicker xylem vessel walls, a lower leaf hydraulic vulnerability and a lower water potential at turgor loss point, which likely confers higher hydraulic safety. Traits such as the water potential at turgor loss point and ratio of sapwood to leaf area (Huber value) showed significant adjustment from wet to dry conditions in the field, indicating phenotypic plasticity and importantly, the ability to increase hydraulic safety in the short term. In the nursery, seedlings from drier populations had smaller leaves and a lower leaf hydraulic vulnerability, suggesting that key traits associated with hydraulic safety are under strong genetic control. Overall, our study suggests a strong genetic control over traits associated with hydraulic safety, which may compromise the survival of wet-origin populations in drier future climates. However, phenotypic plasticity in physiological and morphological traits may confer sufficient hydraulic safety to facilitate genetic adaptation.

Effect of Long-Term vs. Short-Term Ambient Ozone Exposure on Radial Stem Growth, Sap Flux and Xylem Morphology of O3-Sensitive Poplar Trees

High ozone (O3) pollution impairs the carbon and water balance of trees, which is of special interest in planted forests. However, the effect of long-term O3 exposure on tree growth and water use, little remains known. In this study, we analysed the relationships of intra-annual stem growth pattern, seasonal sap flow dynamics and xylem morphology to assess the effect of long term O3 exposure of mature O3-sensitive hybrid poplars (‘Oxford’ clone). Rooted cuttings were planted in autumn 2007 and drip irrigated with 2 liters of water as ambient O3 treatment, or 450 ppm ethylenediurea (N-[2-(2-oxo-1-imidazolidinyl)ethyl]-N0-phenylurea, abbreviated as EDU) solution as O3 protection treatment over all growing seasons. During 2013, point dendrometers and heat pulses were installed to monitor radial growth, stem water relations and sap flow. Ambient O3 did not affect growth rates, even if the seasonal culmination point was 20 days earlier on average than that recorded in the O3 protected trees. Under ambient O3, trees showed reduced seasonal sap flow, however, the lower water use was due to a decrease of Huber value (decrease of leaf area for sapwood unit) rather than to a change in xylem morphology or due to a direct effect of sluggish stomatal responses on transpiration. Under high evaporative demand and ambient O3 concentrations, trees showed a high use of internal stem water resources modulated by stomatal sluggishness, thus predisposing them to be more sensitive water deficit during summer. The results of this study help untangle the compensatory mechanisms involved in the acclimation processes of forest species to long-term O3 exposure in a context of global change.

The hydraulic architecture of Eucalyptus trees growing across a gradient of depth-to-groundwater

Heterogeneity in water availability acts as an important driver of variation in plant structure and function. Changes in hydraulic architecture represent a key mechanism by which adaptation to changes in water availability can be expressed in plants. The aim of this study was to investigate whether differences in depth-to-groundwater influence the hydraulic architecture of Eucalyptus trees in remnant woodlands within mesic environments. Hydraulic architecture of trees was examined in winter and summer by measuring the following traits: Huber value (HV: the ratio between sapwood area and leaf area), branch hydraulic conductivity (leaf and sapwood area specific), sapwood density, xylem vulnerability (P50 and Pe) and hydraulic safety margins across four sites where depth-to-groundwater ranged from 2.4 to 37.5 m. Huber value increased significantly as depth-to-groundwater increased. Neither sapwood density nor branch hydraulic conductivity (sapwood and leaf area specific) varied significantly across sites. Xylem vulnerability to embolism (represented by P50 and Pe) in both seasons was significantly and negatively correlated with depth-to-groundwater. Hydraulic safety margins increased with increasing depth-to-groundwater and therefore trees growing at sites with deeper water tables were less sensitive to drought induced embolism. These results showed plasticity in some, but not all, hydraulic traits (as reflected in HV, P50, Pe and hydraulic safety margin) in response to increase in depth-to-groundwater in a mesic environment.

The influence of depth-to-groundwater on structure and productivity of Eucalyptus woodlands

Although it is well documented that access to groundwater can help plants survive drought in arid and semiarid areas, there have been few studies in mesic environments that have evaluated variation of vegetation characteristics across a naturally occurring gradient in depth-to-groundwater (DGW). The aim of this study was to determine whether differences in groundwater depth influence structural attributes and productivity of remnant woodlands in south-eastern Australia. The study area was located in the Kangaloon bore-field area of New South Wales, where DGW varies from 2.4 m to 37.5 m and rainfall is plentiful. We examined structural (leaf-area index, basal area, stem density, tree height, Huber value (HV) and aboveground biomass) and functional (aboveground net primary productivity (ANPP)) attributes of seven woodland sites differing in DGW. We also used ∂13C analysis of sapwood across six sites, along with observed non-linear changes in structural attributes, to infer groundwater use by trees. Significant differences in structural attributes and ANPP were observed across sites. The three shallowest sites with 2.4 m, 4.3 m and 5.5 m DWG had significantly larger aboveground biomass and ANPP than did the four deepest sites (DGW ≥9.8 m). Across all attributes (except HV in the summer, where the mean values were significantly larger at sites where DGW was 5.5 m or less and across the four deeper sites (DGW ≥9.8 m), there were no differences in these three structural traits, nor in ANPP. Despite finding no significant differences in HV across sites in the summer, in winter, the two deepest sites had a significantly larger HV than did the two shallowest sites. Significant increases in ∂13C of sapwood occurred across five of the six sites, consistent with increasing water-use efficiency as DGW increased, reflecting the declining availability of groundwater with increasing DGW. This study has demonstrated that even in a mesic environment, putative access to groundwater can have important impacts on structural and functional traits of trees and, consequently, on woodland productivity.