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The efficiency-safety tradeoff has been thoroughly investigated in plants, especially concerning their capacity to transport water and avoid embolism. Stomatal regulation is a vital plant behaviour to respond to soil and atmospheric water limitation. Recently, a stomatal efficiency-safety tradeoff was reported where plants with higher maximum stomatal conductance (gmax) exhibited greater sensitivity to stomatal closure during soil drying, that is, less negative leaf water potential at 50% gmax (ψgs50). However, the underlying mechanism of this gmax-ψgs50 tradeoff remains unknown. Here, we utilized a soil-plant hydraulic model, in which stomatal closure is triggered by nonlinearity in soil-plant hydraulics, to investigate such tradeoff. Our simulations show that increasing gmax is aligned with less negative ψgs50. Plants with higher gmax (also higher transpiration) require larger quantities of water to be moved across the rhizosphere, which results in a precipitous decrease in water potential at the soil-root interface, and therefore in the leaves. We demonstrated that the gmax-ψgs50 tradeoff can be predicted based on soil-plant hydraulics, and is impacted by plant hydraulic properties, such as plant hydraulic conductance, active root length and embolism resistance. We conclude that plants may therefore adjust their growth and/or their hydraulic properties to adapt to contrasting habitats and climate conditions.

Plant function arises from a complex network of structural and physiological traits. Explicit representation of these traits, as well as their connections with other biophysical processes, is required to advance our understanding of plant-soil-climate interactions. We used the Terrestrial Regional Ecosystem Exchange Simulator (TREES) to evaluate physiological trait networks in maize. Net primary productivity (NPP) and grain yield were simulated across five contrasting climate scenarios. Simulations achieving high NPP and grain yield in high precipitation environments featured trait networks conferring high water use strategies: deep roots, high stomatal conductance at low water potential (“risky” stomatal regulation), high xylem hydraulic conductivity, and high maximal leaf area index. In contrast, high NPP and grain yield was achieved in dry environments with low late-season precipitation via water conserving trait networks: deep roots, high embolism resistance, and low stomatal conductance at low leaf water potential (“conservative” stomatal regulation). We suggest that our approach, which allows for the simultaneous evaluation of physiological traits and their interactions (i.e., networks), has potential to improve crop growth predictions in different environments. In contrast, evaluating single traits in isolation of other coordinated traits does not appear to be an effective strategy for predicting plant performance.Summary statementOur process-based model uncovered two beneficial but contrasting trait networks for maize which can be understood by their integrated effect on water use/conservation. Modification of multiple, physiologically aligned, traits were required to bring about meaningful improvements in NPP and yield.

Hydraulic failure resulting from drought-induced embolism in the xylem of plants is a key determinant of reduced productivity and mortality. Methods to assess this vulnerability are difficult to achieve at scale, leading to alternative metrics and correlations with more easily measured traits. These efforts have led to the longstanding and pervasive assumed mechanistic link between vessel diameter and vulnerability in angiosperms. However, there are at least two problems with this assumption that requires critical re-evaluation: (1) our current understanding of drought-induced embolism does not provide a mechanistic explanation why increased vessel width should lead to greater vulnerability, and (2) the most recent advancements in nanoscale embolism processes suggest that vessel diameter is not a direct driver. Here, we review data from physiological and comparative wood anatomy studies, highlighting the potential anatomical and physicochemical drivers of embolism formation and spread. We then put forward key knowledge gaps, emphasising what is known, unknown and speculation. A meaningful evaluation of the diameter–vulnerability link will require a better mechanistic understanding of the biophysical processes at the nanoscale level that determine embolism formation and spread, which will in turn lead to more accurate predictions of how water transport in plants is affected by drought.

In the stems of terrestrial vascular plants studied to date, the diameter of xylem water-conducting conduits D widens predictably with distance from the stem tip L approximating D ∝ Lb , with b ≈ 0.2. Because conduit diameter is central for conductance, it is essential to understand the cause of this remarkably pervasive pattern. We give reason to suspect that tip-to-base conduit widening is an adaptation, favored by natural selection because widening helps minimize the increase in hydraulic resistance that would otherwise occur as an individual stem grows longer and conductive path length increases. Evidence consistent with adaptation includes optimality models that predict the 0.2 exponent. The fact that this prediction can be made with a simple model of a single capillary, omitting much biological detail, itself makes numerous important predictions, e.g. that pit resistance must scale isometrically with conduit resistance. The idea that tip-to-base conduit widening has a nonadaptive cause, with temperature, drought, or turgor limiting the conduit diameters that plants are able to produce, is less consistent with the data than an adaptive explanation. We identify empirical priorities for testing the cause of tip-to-base conduit widening and underscore the need to study plant hydraulic systems leaf to root as integrated wholes.

A classic theory proposes that plant xylem cannot be both highly efficient in water transport and resistant to embolism, and therefore a hydraulic efficiency-safety trade-off should exist. However, the trade-off is weak, and many species exhibit both low efficiency and low safety, falling outside of the expected trade-off space. It remains unclear under what climatic conditions these species could maintain competitive fitness. We compiled hydraulic efficiency and safety traits for 682 observations of 499 woody species from 178 sites world-wide and measured the position of each observation within the proposed trade-off space. For both angiosperms and gymnosperms, observations from sites with high climatic seasonality, especially precipitation seasonality, tended to have higher hydraulic safety and efficiency than observations from sites with low seasonality. Specifically, high vapour pressure deficit, high solar radiation, and low precipitation during the wet season were driving factors. Strong climatic seasonality and drought in both dry and wet seasons appear to be ecological filters that select for species with co-optimized safety and efficiency, whereas the opposite environmental conditions may allow the existence of plants with low efficiency and safety.

Water released from wood during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twigs of 30 temperate angiosperm tree species, we measured day capacitance (between predawn and midday), water content, wood density, and anatomical traits, that is, vessel dimensions, tissue fractions, and vessel-tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, wood density (WD) and predawn lumen volumetric water content (VWCL-pd ) together were the strongest predictors of day capacitance (r2adj = .44). Vessel-tissue contact fractions explained an additional ~10% of the variation in day capacitance. Regression models were not improved by including tissue lumen fractions. Among diffuse-porous species, VWCL-pd and vessel-ray contact fraction together were the best predictors of day capacitance, whereas among semi/ring-porous species, VWCL-pd , WD and vessel-fibre contact fraction were the best predictors. At predawn, wood was less than fully saturated for all species (lumen relative water content = 0.52 ± 0.17). Our findings imply that day capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue.

Root-released carboxylates enhance the availability of manganese (Mn), which enters roots through transporters with low substrate specificity. Leaf Mn concentration ([Mn]) has been proposed as a signature for phosphorus (P)-mobilising carboxylates in the rhizosphere. Here we test whether leaf [Mn] provides a signature for root functional types related to P acquisition. Across 727 species at 66 sites in Australia and New Zealand, we measured leaf [Mn] as related to root functional type, while also considering soil and climate variables. To further assess the specific situations under which leaf [Mn] is a suitable proxy for rhizosphere carboxylate concentration, we studied leaf [Mn] along a strong gradient in water availability on one representative site. In addition, we focused on two systems where a species produced unexpected results. Controlling for background site-specific variation in leaf [Mn] with soil pH and mean annual precipitation, we established that mycorrhizal species have significantly lower leaf [Mn] than non-mycorrhizal species with carboxylate-releasing root structures, e.g., cluster roots. In exception to the general tendency, leaf [Mn] did not provide information about root functional types under seasonally waterlogged conditions, which increase iron availability and thereby interfere with Mn-uptake capacity. Two further exceptions were scrutinised, leading to the conclusion that they were ‘anomalous’ in not functioning like typical species in their families, as expected according to the literature. Leaf [Mn] variation provides considerable insights on differences in belowground functioning among co-occurring species. Using this approach, we concluded that, within typical mycorrhizal families, some species actually depend on a carboxylate-releasing P-mobilising strategy. Likewise, within families that are known to produce carboxylate-releasing cluster roots, some do not produce functional cluster roots when mature. An analysis of leaf [Mn] can alert us to such ‘anomalous’ species.

"Least-cost theory" posits that C3 plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO2 drawdown (lower ratio of leaf internal to ambient CO2 , Ci :Ca ) during light-saturated photosynthesis, and at higher leaf N per area (Narea ) and higher carboxylation capacity (Vcmax 25 ) for a given rate of stomatal conductance to water vapour, gsw . These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the Narea -gsw and Vcmax 25 -gsw slopes, and negative effects on Ci :Ca . The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.

Stem xylem-specific hydraulic conductivity (KS ) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in KS has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global-scale patterns of KS and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species-at-site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how KS varied with climatic variables, plant functional types, and biomes. Growing-season temperature and growing-season precipitation drove global variation in KS independently. Both the mean and the variation in KS were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for KS in predicting shifts in community composition in the face of climate change.

Copper (Cu) is important for many aspects of plant function including photosynthesis. It has been suggested that photosynthesis, especially in young leaves, is prioritized for Cu delivery after deficiency in hybrid poplar. To determine relative Cu delivery prioritization, we enriched hydroponic plant growth media of Cu-deficient poplar with 98% 65Cu and tracked Cu delivery after deficiency to young leaves, mature leaves and stems. Young leaves acquired ~58% more 65Cu on Day 1 and ~65% more 65Cu by Day 3 compared with mature leaves. Additionally, stomatal conductance (gs) was measured on leaves for 6 weeks and during a 3-day 65Cu pulse resupply period. During deficiency, mature leaves maintained a higher gs than younger leaves but 3 days after Cu resupply the younger leaves that had recovered showed the highest gs. In conclusion, these results provide a quantitative understanding of how Cu is systemically transported and distributed to photosynthetic and stem tissues.

We introduce the AusTraits database - a compilation of values of plant traits for taxa in the Australian flora (hereafter AusTraits). AusTraits synthesises data on 448 traits across 28,640 taxa from field campaigns, published literature, taxonomic monographs, and individual taxon descriptions. Traits vary in scope from physiological measures of performance (e.g. photosynthetic gas exchange, water-use efficiency) to morphological attributes (e.g. leaf area, seed mass, plant height) which link to aspects of ecological variation. AusTraits contains curated and harmonised individual- and species-level measurements coupled to, where available, contextual information on site properties and experimental conditions. This article provides information on version 3.0.2 of AusTraits which contains data for 997,808 trait-by-taxon combinations. We envision AusTraits as an ongoing collaborative initiative for easily archiving and sharing trait data, which also provides a template for other national or regional initiatives globally to fill persistent gaps in trait knowledge., Measurement(s)plant traitTechnology Type(s)digital curationSample Characteristic - OrganismViridiplantaeSample Characteristic - LocationAustralia Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.14545755

Coordination between sapwood-specific hydraulic conductivity (Ks) and stomatal conductance (gs) has been identified in previous studies; however, coordination between leaf hydraulic conductance (Kleaf) and gs, as well as between Kleaf and Ks is not always consistent. This suggests that there is a need to improve our understanding of the coordination among hydraulic and gas exchange traits. In this study, hydraulic traits (e.g., Ks and Kleaf) and gas exchange traits, including gs, transpiration (E) and net CO2 assimilation (Aarea), were measured across 33 co-occurring subtropical woody species. Kleaf was divided into two components: leaf hydraulic conductance inside the xylem (Kleaf-x) and outside the xylem (Kleaf-ox). We found that both Kleaf-x and Kleaf-ox were coordinated with gs and E, but the correlations between Kleaf-ox and gs (or E) were substantially weaker, and that Ks was coordinated with Kleaf-x, but not with Kleaf-ox. In addition, we found that Ks, Kleaf-x and Kleaf-ox together explained 63% of the variation in gs and 42% of the variation in Aarea across species, with Ks contributing the largest proportion of explanatory power, whereas Kleaf-ox contributed the least explanatory power. Our results demonstrate that the coordination between leaf water transport and gas exchange, as well as the hydraulic linkage between leaf and stem, were weakened by Kleaf-ox. This highlights the possibility that water transport efficiencies of stem and leaf xylem, rather than that of leaf tissues outside the xylem, are important determinants of stomatal conductance and photosynthetic capacity across species.

We evaluated sixteen traits related to water acquisition and transport, stomatal conductance, and photosynthesis within a diverse panel of maize inbred lines, founders of the U.S. maize nested association mapping (NAM) population, with the aim to determine which traits confer improved growth under water deficit and well-watered conditions. Lasso regression revealed that three key traits explained meaningful and independent proportions of variation in total end-of-season biomass under deficit irrigation (multiple r2 = 0.86): 1) the maximal net CO2 assimilation rate (P = 0.007), 2) the achievable stomatal conductance during the hottest part of the day (P = 0.005), and 3) the width-to-depth ratio of the root system at the seedling stage (P = 0.060), i.e., initial deep root system development facilitated growth. Under well-watered conditions, maximal stomatal conductance in the morning (P = 0.014) and the width-to-depth ratio of the root system (P = 0.043) were identified as key traits contributing to improved performance (multiple r2 = 0.68). Structural equation models revealed that growth under water deficit was linked more strongly to stomatal conductance occurring during the middle of the day (std coef = 0.75; P = 0.006), rather than the maximal stomatal conductance (std coef = −0.25; P = 0.368). In turn, the maintenance of stomatal conductance through the middle of the day depended on the capacity of the xylem tissue to supply water (per unit cross-sectional area) (std coef = 0.48; P = 0.046). Aligned with the transport of water to the stomata and growth, root system depth ( r = 0.77; P = 0.003) and width-to-depth ratio ( r = −0.55; P = 0.064) at seedling stages were also correlated with the capacity of the xylem to transport water, thus suggesting close coordination between root, xylem, and stomatal traits to achieve greater growth under water deficit and well-watered conditions. We propose that maize performance under the drought conditions considered here, could likely be improved via lower stomatal sensitivity to hydraulic and atmospheric cues, greater xylem conductivity, and a deeper, but not necessarily more extensive, root system.

Water shortages are responsible for the greatest crop losses around the world and are expected to worsen. In arid areas where agriculture is dependent on irrigation, various forms of deficit irrigation management have been suggested to achieve high yields with less water used by the crop (i.e. evapotranspiration, ET). This study of maize evaluated twelve treatments with varying levels of deficit irrigation during late vegetative and maturation (grain filling) growth stages in semi-arid Northern Colorado. In particular, application of greater deficit during the late vegetative state with full or nearly full ET during the rest of the season consistently resulted in yield similar to full ET treatments while saving approximately 15–17% of ET. Maize given 40% of full ET during the late vegetative period had slightly reduced leaf area index (LAI) with significant leaf curling, thus reduced light interception during vegetative growth. However, when plants were fully watered during anthesis, all treatments had full canopy cover with no differences in light interception. The efficiency of photosystem II (quantum yield) declined with water stress but recovered with re-watering. The ability of photosystem II and light interception to recover after stress when well-watered suggests that reductions in biomass and yield resulted from stomatal closure, reduced photochemistry, or loss of xylem conductance that was temporary. With little indication of permanent decline in carbon assimilation after reducing ET in vegetative stages, maize appears able to achieve high grain yield if soil water is readily available during the reproductive and maturation stages. However, plants given full or nearly full irrigation during the entire vegetative period followed by stress later on during the maturation period, had dramatically greater yield loss than ET savings. Thus, while strategic deficit irrigation might maintain yield with less water, it may be especially important for buffering crops against yield losses due to end of season water shortfalls in water limited environments.

While infrared thermometry and thermal imagery have potential to detect crop water stress and quantify evapotranspiration, both valuable in irrigation scheduling, it is often difficult to isolate plant canopy temperature from background temperatures. In this study, we demonstrate a simple technique that uses a homogeneous background temperature that contrasts with canopy temperature, thereby allowing the canopy temperature itself to be isolated in a thermal image. Analysis of pixel temperatures and their associated statistics demonstrate the potential of this method to measure small (ca. < 0.5°C) and rapid (ca. < 1 s) fluctuations in leaf energy balance. This technique has broad applicability in greenhouse, growth chamber, and other small-scale experiments where real time response of individual leaves or canopies is required. Keywords: Thermal imaging, Crop water stress, Infrared thermometry, Background masking, Deficit irrigation, Greenhouse.

There is increasing interest in understanding how trait networks can be manipulated to improve the performance of crop species. Working towards this goal, we have identified key traits linking the acquisition of water, the transport of water to the sites of evaporation and photosynthesis, stomatal conductance, and growth across eight maize hybrid lines grown under well-watered and water-limiting conditions in Northern Colorado. Under well-watered conditions, hybrids with higher end-of-season growth and grain yield exhibited higher leaf-specific conductance, lower operating water potentials, higher rates of midday stomatal conductance, higher rates of net CO2 assimilation, and greater leaf osmotic adjustment. This trait network was similar under water-limited conditions with the notable exception that linkages between water transport, midday stomatal conductance, and growth were even stronger than under fully watered conditions. The maintenance of high leaf-specific conductance throughout the day was achieved via higher maximal conductance rates rather than lower susceptibility to conductance loss. Our results suggest that efforts to improve maize performance in well-watered and water-limiting conditions would benefit from considering the physiological trait networks governing water and carbon flux rather than focusing on single traits independently of one another.

There is increasing interest in understanding how trait networks can be manipulated to improve the performance of crop species. Working towards this goal, we have identified key traits linking the acquisition of water, the transport of water to the sites of evaporation and photosynthesis, stomatal conductance, and growth across eight maize hybrid lines grown under well-watered and water-limiting conditions in Northern Colorado. Under well-watered conditions, well-performing hybrids exhibited high leaf-specific conductance, low operating water potentials, high rates of midday stomatal conductance, high rates of net CO2 assimilation, greater leaf osmotic adjustment, and higher end-of-season growth and grain yield. This trait network was similar under water-limited conditions with the notable exception that linkages between water transport, midday stomatal conductance, and growth were even stronger than under fully-watered conditions. The results of this experiment suggest that similar trait networks might confer improved performance under contrasting climate and soil conditions, and that efforts to improve the performance of crop species could possibly benefit by considering the water transport pathway within leaves, as well as within the whole-xylem, in addition to root-level and leaf-level traits.

Root pressure, also manifested as profusive sap flowing from cut stems, is a phenomenon in some species that has perplexed biologists for much of the last century. It is associated with increased crop production under drought, but its function and regulation remain largely unknown. In this study, we investigated the initiation, mechanisms, and possible adaptive function of root pressure in six genotypes of Sorghum bicolor during a drought experiment in the greenhouse. We observed that root pressure was induced in plants exposed to drought followed by re-watering but possibly inhibited by 100% re-watering in some genotypes. We found that root pressure in drought stressed and re-watered plants was associated with greater ratio of fine: coarse root length and shoot biomass production, indicating a possible role of root allocation in creating root pressure and adaptive benefit of root pressure for shoot biomass production. Using RNA-Seq, we identified gene transcripts that were up- and down-regulated in plants with root pressure expression, focusing on genes for aquaporins, membrane transporters, and ATPases that could regulate inter- and intra-cellular transport of water and ions to generate positive xylem pressure in root tissue.

Vascular networks in plant leaves must provide for the safe and efficient transport of water, nutrients, and energy; however, the conditions whereby these networks lose and regain conductive capacity are still poorly understood. We measured the loss and recovery of leaf hydraulic conductance (Kleaf) in a tropical monocotyledon (Bambusa vulgaris) and dicotyledon species (Bauhinia blakeana) using Rehydration Kinetics Method (RKM) as well as a recently developed optical method. We found that both species lost ca 88% of their maximal Kleaf (measured by RKM) before any embolization was detected in their conductive elements (via optical observation). This suggests that the majority of loss in Kleaf, as measured with RKM, was associated with resistances other than embolization. Furthermore, embolism in B. vulgaris, a species known to generate root pressure, was reversed when rehydrated under positive pressure (120 kPa), but not under atmospheric pressure. In contrast, embolism was not reversed in B. blakeana under either elevated or atmospheric pressure. However, reductions in Kleaf that was not associated with embolization was recovered by this species when rehydrated under atmospheric conditions, whereas B. vulgaris did not exhibit any recovery under the same conditions. We suggest that root pressure is an adaptive mechanism allowing for the reversal of embolism and the recovery of extraxylary conductance. The absence of embolism reversal in B. blakeana suggests that embolization may be permanent without neutral or positive xylem pressure, but that the recovery of extraxylary conductance can be regained routinely in this species in the absence of root pressure.

The evolution of angiosperms was accompanied by the segregation and specialisation of their xylem tissues. This study aimed to determine whether the fraction and arrangement of parenchyma tissue influence the hydraulic efficiency-safety trade-off in the basal angiosperms. We examined xylem anatomical structure and hydraulic functioning of 28 woody species of Magnoliids in a tropical rainforest of Madagascar and reported, for the first time, quantitative measurements that support the relationship between vessel-to-xylem parenchyma connectivity and the hydraulic efficiency-safety trade-off. We also introduced a new measurement - the distance of species from the trade-off limit - to quantify the co-optimisation of hydraulic efficiency and safety. Although the basal angiosperms in this study had low hydraulic conductivity and safety, species with higher axial parenchyma fraction (APf) had significantly higher hydraulic conductivity. Hydraulic efficiency-safety optimisation was accompanied by higher APf and vessel-to-axial parenchyma connectivity. Conversely, species exhibiting high ray parenchyma fraction and high vessel-to-ray connectivity had lower Ks and were further away from the hydraulic trade-off limit line. Our results provide evidence that axial parenchyma fraction and paratracheal arrangement are associated with both enhanced hydraulic efficiency and safety.

Use of the term “functional" trait has increased exponentially in ecology. Although accounting for numerous ecological questions, this concept raises several issues. We propose that the term “functional” could be misleading because (1) no rigorous criteria exist to identify “functional” traits and (2) it suggests that only some traits (“functional” ones) can inform our understanding of species functioning, whatever the scale or discipline. Hence, the concept of "functional" trait in ecology is starting to be challenged and it remains unclear why some traits should be considered functional, whereas other traits should not. We argue that the most used “functional” traits are meaningful because they reflect important differences between populations or species, based on synchronic comparisons, that is, irrespective of time (hereafter “pattern” traits). Hence, they are useful for identifying trade‐offs and strategies across large numbers of observations, usually at rather coarse scales, and are most often used in analyses of “big data.” However, given that many ecological processes occur across short time scales and narrow gradients of climate and resource availability, the efficacy of these traits to inform us about these ecological processes appears questionable. We show that trait measurements that take time explicitly into account (hereafter “process” traits) differ from pattern traits because they quantify the flows of material and energy within a given environment across a defined period of time. Although pattern traits and process traits are both functional, it is important to understand the differences between the approaches. Moreover, better accounting of ontogeny, life form, plasticity, and genetic variability is required to enhance the convergence between pattern and process approaches. This revised framework allows more explicit connections between trait ecology and other biological sciences. It should enhance the study of processes at all scales in order to investigate efficiently the adaptive responses of biological organisms to climate change., Revising the concept of "functional" trait: In ecology, pattern approaches based on "functional" traits that do not explicitly take time into account, differ from process approaches which measure response curves at all scales over time. Using the term “functional” for traits meaningful for both pattern and process approaches, and better accounting of ontogeny, life form, plasticity and genetic variability should enhance the convergence across approaches within biological sciences.

Both empirical and theoretical models have been widely used to calculate a crop water stress index (CWSI) − a metric often used to describe crop water status. The purpose of this study was to determine the accuracy, limitation, and uncertainty of an empirical (CWSI-E) and two theoretical models compared with sap flow measurement in maize. One theoretical model used a calculated aerodynamic resistance (CWSI-T1), and the other theoretical model used seasonal average aerodynamic resistance (CWSI-T2). Considering the uncertainty of crop coefficient and sap flow measurement, CWSI-T2 and CWSI-E models gave reasonable overall estimates of water stress. The average root mean square deviation at each growth stage from each model ranged from 0.16 to 0.33. CWSI-T2 and the CWSI-E provided relatively accurate prediction of crop stress, both between growth stages and irrigation events. However, CWSI-T1 did not accurately predict water stress between growth stages or between irrigation events. By including climate factors, crop water stress estimated by CWSI-T2 showed less variation and uncertainty than CWSI-E. The uncertainty of both CWSI-T2 and CWSI-E decreased with increasing vapor pressure deficit (VPD), and CWSI-E show larger crop water stress prediction uncertainty. The intercept of non-water stress baseline was the main source of the uncertainty for CWSI-E and CWSI-T2. Considering both uncertainty and stability, we recommend CWSI-T2 model (i.e., seasonal average aerodynamic resistance) for maize water stress assessment.

Water transport in leaf vasculature is a fundamental process affecting plant growth, ecological interactions and ecosystem productivity, yet the architecture of leaf vascular networks is poorly understood. Although Murray's law and the West-Brown-Enquist (WBE) theories predict convergent scaling of conduit width and number, it is not known how conduit scaling is affected by habitat aridity or temperature. We measured the scaling of leaf size, conduit width and conduit number within the leaves of 36 evergreen Angiosperms spanning a large range in aridity and temperature in eastern Australia. Scaling of conduit width and number in midribs and 2° veins did not differ across species and habitats (P > 0.786), and did not differ from that predicted by Murray's law (P = 0.151). Leaf size was strongly correlated with the hydraulic radius of petiole conduits (r2 = 0.83, P 0.064), nor did the scaling exponent differ significantly from that predicted by hydraulic theory (P = 0.086). The maximum radius of conduits in petioles was positively correlated with the temperature of the coldest quarter (r2 = 0.67; P

Parenchyma represents a critically important living tissue in the sapwood of the secondary xylem of woody angiosperms. Considering various interactions between parenchyma and water transporting vessels, we hypothesize a structure-function relationship between both cell types. Through a generalized additive mixed model approach based on 2,332 woody angiosperm species derived from the literature, we explored the relationship between the proportion and spatial distribution of ray and axial parenchyma and vessel size, while controlling for maximum plant height and a range of climatic factors. When factoring in maximum plant height, we found that with increasing mean annual temperatures, mean vessel diameter showed a positive correlation with axial parenchyma proportion and arrangement, but not for ray parenchyma. Species with a high axial parenchyma tissue fraction tend to have wide vessels, with most of the parenchyma packed around vessels, whereas species with small diameter vessels show a reduced amount of axial parenchyma that is not directly connected to vessels. This finding provides evidence for independent functions of axial parenchyma and ray parenchyma in large vesselled species and further supports a strong role for axial parenchyma in long-distance xylem water transport.

The rate of elongation and thickening of individual branches (shoots) varies across plant species. This variation is important for the outcome of competition and other plant–plant interactions. Here, we compared rates of shoot growth across 44 species from tropical, warm temperate and cool temperate forests of eastern Australia. Shoot growth rate was found to correlate with a suite of traits including the potential height of the species, xylem-specific conductivity, leaf size, leaf area per xylem cross-section (LA/XA), twig diameter (at 40 cm length), wood density and modulus of elasticity. Within this suite of traits, maximum plant height was the clearest correlate of growth rates, explaining 50%–67% of the variation in growth overall (p

Present efforts to improve maize performance in water-limited environments are aggressively pursuing leaf-level traits, such as quantum efficiency, mesophyll osmoregulation, and stress-protein responses. However, it is possible that improvement of these traits will lead directly to hydraulic failure if hydraulics and photosynthesis are closely aligned. Here, we address the whole plant response to drought stress and ask if photosynthetic and hydraulic traits appear bundled together as a “coordinated” response, or if these traits operate largely independently of one another. Xylem conductance of leaves and stems, whole plant conductance, stomatal conductance, rate of electron transport (ETR), maximal catalytic rate of phosphoenolpyruvate carboxylase (V pmax ) (EC 4.1.1.31), and net CO 2 assimilation (A max ) were measured in maize plants subjected to contrasting levels of drought stress in greenhouse and field experiments. Photosynthetic traits (A max , ETR, V pmax ), hydraulic traits (whole plant and stem conductance) and stomatal conductance were all reduced by >80% as leaf water potentials declined below −3.0 MPa. Furthermore, 83% of the variation associated with the photosynthetic and hydraulic traits measured in this study was explained by a single principal component, revealing a remarkable degree of alignment among them. Whole plant transpiration rates recovered to ca 90% of maximal values 4 d after lifting severe drought stress (Ψ leaf ≈ −3.5 MPa). Closely aligned hydraulic, photosynthetic, and stomatal responses to drought stress suggest that improvements to individual traits, in isolation to each other, may lead to the loss of plant functioning (e.g. water transport) rather than leading to marked improvements in growth.

This article comments on: Water relations of Calycanthus flowers: Hydraulic conductance, capacitance, and embolism resistance.

SUMMARYWater released from wood tissue during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twig wood of 30 temperate angiosperm tree species, we measured capacitance, water content, wood density, and anatomical traits, i.e., vessel dimensions, tissue fractions, and vessel-tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, the strongest predictors of capacitance were wood density (WD) and predawn lumen volumetric water content (VWCL-pd, r2adj=0.44, Ppd) or tissue lumen fractions. Among diffuse-porous species, VWCL-pd and vessel-ray contact fraction were the best predictors of capacitance, whereas among ring/semi-ring-porous species, VWCL-pd, WD and vessel-fibre contact fraction were the best predictors. Mean RWCpd was 0.65±0.13 and uncorrelated with WD. VWCL-pd was weakly negatively correlated with WD. Our findings imply that capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue.

Coordination between plant height and xylem traits is aligned with habitat aridity across the Earth’s terrestrial biomes., Water must be transported long distances in tall plants, resulting in increasing hydraulic resistance, which may place limitations on the maximum plant height (Hmax) in a given habitat. However, the coordination of hydraulic traits with Hmax and habitat aridity remains poorly understood. To explore whether Hmax modifies the trade-off between hydraulic efficiency and safety or how water availability might influence the relationship between Hmax and other hydraulic traits, we compiled a dataset including Hmax and 11 hydraulic traits for 1281 woody species from 369 sites worldwide. We found that taller species from wet habitats exhibited greater xylem efficiency and lower hydraulic safety, wider conduits, lower conduit density, and lower sapwood density, which were all associated with habitat water availability. Plant height and hydraulic functioning appear to represent a single, coordinated axis of variation, aligned primarily with water availability, thus suggesting an important role for this axis in species sorting processes.

Although CO2 and H2O exchange rates are often measured in experiments as indicators of physiological plant responses these gas exchange measurements are prone to large experimental error. Gas exchange equipment and technology have improved greatly over the past two decades which supports scrutinizing current issues of experimental error in measuring plant photosynthesis and stomatal conductance. This report shows results of a greenhouse experiment with the goal of identifying lessor understood sources of experimental error and variation in measurements with the LI-COR 6400XT gas exchange system. A variety of plant types were used to encompass differing species variation. We found significant sources of experimental error in 1) the time for initial adjustment when placing a leaf in the leaf chamber 2) the time-of-day when measuring 3) leaf age 4) having the chamber window full vs. partially full with leaf tissue 5) using a leaf chamber environment that greatly diverges from the whole plant environment 6) differing degree of experimental error depending upon plant species. A situation with multiple contributors to error would result in useless gas-exchange data. Recommendations for minimizing these experimental errors are given.

Research in plant hydraulics has provided important insights into plant responses to drought and species absolute drought tolerance. However, our ability to predict when plants will die from hydraulic failure under extreme drought is limited by a lack of knowledge with regards to the dynamics of plant desiccation following stomatal closure. Thus, we develop a simple hydraulics model based on branch-level traits that incorporates key aspects of allometry, rates of water loss and resistance to embolism thresholds in order to define species differences in the time it takes plants to desiccate from stomatal closure to lethal levels of drought stress.

BACKGROUND AND AIMS: The structural properties of leaf venation and xylem anatomy strongly influence leaf hydraulics, including the ability of leaves to maintain hydraulic function during drought. Here we examined the strength of the links between different leaf venation traits and leaf hydraulic vulnerability to drought (expressed as P50(leaf) by rehydration kinetics) in a diverse group of 26 woody angiosperm species, representing a wide range of leaf vulnerabilities, from four low-nutrient sites with contrasting rainfall across eastern Australia. METHODS: For each species we measured key aspects of leaf venation design, xylem anatomy and leaf morphology. We also assessed for the first time the scaling relationships between hydraulically weighted vessel wall thickness (t(h)) and lumen breadth (b(h)) across vein orders and habitats. KEY RESULTS: Across species, variation in P50(leaf) was strongly correlated with the ratio of vessel wall thickness (t(h)) to lumen breadth (b(h)) [(t/b)(h); an index of conduit reinforcement] at each leaf vein order. Concomitantly, the scaling relationship between t(h) and b(h) was similar across vein orders, with a log–log slope less than 1 indicating greater xylem reinforcement in smaller vessels. In contrast, P50(leaf) was not related to t(h) and b(h) individually, to major vein density (Dv(major)) or to leaf size. Principal components analysis revealed two largely orthogonal trait groupings linked to variation in leaf size and drought tolerance. CONCLUSIONS: Our results indicate that xylem conduit reinforcement occurs throughout leaf venation, and remains closely linked to leaf drought tolerance irrespective of leaf size.

It is not currently well-understood how much xylem conductance is lost in maize plants during the day, if conductance is recovered during the night, or what soil water conditions are required for recovery to take place. To answer these questions we designed a greenhouse experiment whereby two genetically dissimilar maize genotypes were subjected to a level of water stress commonly experienced in the field (Ψxylem ∼-2 MPa). We then measured the loss of stem-specific conductivity associated with this level of stress, as well as the overnight recovery following three re-watering treatments: Ψsoil ∼ 0 MPa, Ψsoil ∼-0.40 MPa, and Ψsoil ∼-1.70 MPa. Mid-day leaf water potentials of -1.98 MPa resulted in stem-specific conductivity (KS) values that were 31.5% of maximal (i.e., 68% loss). Returning soils to field capacity (Ψsoil ∼ 0 MPa) overnight allowed for the significant recovery of KS (76% of maximal), whereas partial watering (Ψsoil ∼-0.40 MPa) resulted KS values that were 51.7% of maximal values, whereas not watering resulted in no recovery (35.4% of maximal; Ψsoil ∼-1.7 MPa). Recovery of KS was facilitated by the generation of root pressure and low rates of nighttime transpiration.

Low water potentials in xylem can result in damaging levels of cavitation, yet little is understood about which hydraulic traits have most influence in delaying the onset of hydraulic dysfunction during periods of drought. We examined three traits contributing to longer desiccation times in excised shoots of 11 species from two sites of contrasting aridity: (i) the amount of water released from plant tissues per decrease in xylem water potential (WΨ); (ii) the minimum xylem water potential preceding acute water stress (defined as P50L; water potential at 50% loss of leaf conductance); and (iii) the integrated transpiration rate between the points of full hydration and P50L (Wtime). The time required for species to reach P50L varied markedly, ranging from 1.3 h to nearly 3 days. WΨ, P50L and Wtime all contributed significantly to longer desiccation times, explaining 28, 22 and 50% of the variance in the time required to reach P50L. Interestingly, these three traits were nearly orthogonal to one another, suggesting that they do not represent alternative hydraulic strategies, but likely trade off with other ecological strategies not evaluated in this study. The majority of water lost during desiccation (60-91%) originated from leaves, suggesting an important role for leaf capacitance in small plants when xylem water potentials decrease below -2 MPa.

Premise of research. Plants are faced with a challenge across all climates they inhabit—they must transport water to their leaves so that photosynthesis can take place. Although this is simple in concept, it can be achieved by different arrangements of root, stem, and leaf traits. The hydraulic functioning of species across aridity gradients is determined by the coordination of these traits. Nevertheless, we have an imperfect understanding of which trait shifts are favored across aridity gradients as well as the alignment of trait shifts with climate.Methodology. We measured hydraulic traits relating to Darcy’s law for 120 angiosperm species across a broad range of climates in eastern Australia; nearly one-third of all biome space on Earth was represented. We then determined which hydraulic trait shifts have been favored across aridity gradients and which climate characteristics these trait shifts aligned with.Pivotal results. Increasing aridity, from climates with similar precipitation and evaporation to...

The causes underlying bark diversity are unclear. Variation has been frequently attributed to environmental differences across sites. However, variation may also result from tradeoffs and coordination between bark's multiple functions. Bark traits may also covary with wood and leaf traits as part of major dimensions of plant variation. To assess hypotheses regarding tradeoffs and functional coordination, we measured bark traits reflecting protection, storage, mechanics, and photosynthesis in branches of 90 species spanning a wide phylogenetic and environmental range. We also tested associations between bark, wood, and leaf traits. We partitioned trait variation within species, and within and across communities to quantify variation associated with across-site differences. We observed associations between bark mechanics and storage, density and thickness, and thickness and photosynthetic activity. Increasing bark thickness contributed significantly to stiffer stems and greater water storage. Bark density, water content, and mechanics covaried strongly with the equivalent wood traits, and to a lesser degree with leaf size, xylem conductivity, and vessel diameter. Most variation was observed within sites and had low phylogenetic signal. Compared with relatively minor across-site differences, tradeoffs and coordination among functions of bark, leaves, and wood are likely to be major and overlooked factors shaping bark ecology and evolution.