The role of leaf superoxide dismutase and proline on intra-specific photosynthesis recovery of Schima superba following drought.

Understanding the physiological and biochemical responses of tree seedlings under extreme drought stress, along with recovery during rewatering, and potential intra-species differences, will allow us to more accurately predict forest responses under future climate change. Here, we selected seedlings from four provenances (AH (Anhui), JX (Jiangxi), HN (Hunan) and GX (Guangxi)) of Schima superba and carried out a simulated drought-rewatering experiment in a field-based rain-out shelter. Seedlings were progressively dried until they reached 50% and 88% loss of xylem hydraulic conductivity (PLC) (i.e. P50 and P88), respectively, before they were rehydrated and maintained at field capacity for 30 days. Leaf photosynthesis (Asat), water status, activity of superoxide dismutase (SOD), and proline (Pro) concentration were monitored and their associations were determined. Increasing drought significantly reduced Asat, relative water content (RWC) and SOD activity in all provenances, and Pro concentration was increased to improve water retention; all four provenances exhibited similar response patterns, associated with similar leaf ultrastructure at pre-drought. Upon rewatering, physiological and biochemical traits were restored to well-watered control values in P50-stressed seedlings. In P88-stressed seedlings, Pro was restored to control values, while SOD was not fully recovered. The recovery pattern differed partially among provenances. There was a progression of recovery following watering, with RWC firstly recovered, followed by SOD and Pro, and then Asat, but with significant associations among these traits. Collectively, the intra-specific differences of S. superba seedlings in recovery of physiology and biochemistry following rewatering highlight the need to consider variations within a given tree species coping with future more frequent drought stress.

Species-specific responses of C and N allocation to N addition: evidence from dual 13C and 15N labeling in three tree species.

Soil nitrogen (N) availability affects plant carbon (C) utilization. However, it is unclear how various tree functional types respond to N addition in terms of C assimilation, allocation, and storage. Here, a microcosm experiment with dual 13C and 15N labeling was conducted to study the effects of N addition (i.e., control, 0 g N kg-1; moderate N addition, 1.68 g N kg-1; and high N addition, 3.36 g N kg-1 soil) on morphological traits, on changes in nonstructural carbohydrates (NSC) in different organs, as well as on C and N uptake and allocation in three European temperate forest tree species (i.e., Acer pseudoplatanus, Picea abies and Abies alba). Our results demonstrated that root N uptake rates of the three tree species increased by N addition. In A. pseudoplatanus, N uptake by roots, N allocation to aboveground organs, and aboveground biomass allocation significantly improved by moderate and high N addition. In A. alba, only the high N addition treatment considerably raised aboveground N and C allocation. In contrast, biomass as well as C and N allocation between above and belowground tissues were not altered by N addition in P. abies. Meanwhile, NSC content as well as C and N coupling (represented by the ratio of relative 13C and 15N allocation rates in organs) were affected by N addition in A. pseudoplantanus and P. abies but not in A. alba. Overall, A. pseudoplatanus displayed the highest sensitivity to N addition and the highest N requirement among the three species, while P. abies had a lower N demand than A. alba. Our findings highlight that the responses of C and N allocation to soil N availability are species-specific and vary with the amount of N addition.

Divergent role of nutrient availability in determining drought responses of sessile oak and Scots pine seedlings: evidence from 13C and 15N dual labeling.

Increased soil nutrient availability can promote tree growth while drought impairs metabolic functioning and induces tree mortality. However, limited information is available about the role of nutrients in the drought responses of trees. A greenhouse experiment was conducted with sessile oak (Quercus petraea (Matt.) Liebl) and Scots pine (Pinus sylvestris L.) seedlings, which were subjected to three fertilization treatments in the first year and two water regimes in the second year. Old and newly fixed carbon (C) and nitrogen (N) allocation were traced by dual labeling with 13C and 15N tracers, respectively, at two time points. Leaf gas exchange, biomass, as well as N and nonstructural carbohydrate (NSC) concentrations of all organs were measured. Fertilization predisposed sessile oak to drought-induced mortality, mainly by prioritizing aboveground growth, C and N allocation, reducing root NSC concentrations and decreasing old C contribution to new growth of leaves. In contrast, fertilization did not additionally predispose Scots pine to drought, with minor effects of fertilization and drought on newly fixed and old C allocation, tissues N and NSC concentrations. The role of nutrients for drought responses of trees seems to be species-specific. Therefore, we suggest nutrient availability and species identity to be considered in the framework of physiological mechanisms affecting drought-induced mortality.

Root relative water content is a potential signal for impending mortality of a subtropical conifer during extreme drought stress.

Adaptation to future climates characterized by more frequent severe droughts requires enhanced mechanistic understanding of tree mortality. However, our knowledge of the physiological limits to withstand extreme drought, and how the coordination between water and carbon traits enhances survival, is still limited. Potted seedlings of Pinus massoniana were dehydrated to three target droughts (percentage loss of stem hydraulic conductivity of ca. 50%, 85%, and 100%; PLC50 , PLC85 and PLC100 ) and then relieved from these target droughts by fully rewatering. Predawn and midday water potentials (Ψ), relative water content (RWC), PLC and nonstructural carbohydrates (NSC) were monitored. During drought, Ψ and RWC declined as PLC increased. Root RWC declined more rapidly than other organ RWCs, particularly after PLC50 stress. All organ NSC concentrations were above predrought values. During rewatering, water trait recovery declined as drought increased, with no mortality at PLC50 but 75% mortality at PLC85 . The observed stem hydraulic recovery at PLC50 following rewatering was not correlated to NSC dynamics. Collectively, our results highlighted the primary role of hydraulic failure in Pinus massoniana seedling mortality by assessing mortality threshold and links among water status and water supply. Root RWC can be considered as a potential warning signal of P. massoniana mortality.

Plant responses to high temperature and drought: A bibliometrics analysis.

Global climate change is expected to further increase the frequency and severity of extreme events, such as high temperature/heat waves as well as drought in the future. Thus, how plant responds to high temperature and drought has become a key research topic. In this study, we extracted data from Web of Science Core Collections database, and synthesized plant responses to high temperature and drought based on bibliometric methods using software of R and VOSviewer. The results showed that a stabilized increasing trend of the publications (1199 papers) was found during the period of 2008 to 2014, and then showed a rapid increase (2583 papers) from year 2015 to 2021. Secondly, the top five dominant research fields of plant responses to high temperature and drought were Plant Science, Agroforestry Science, Environmental Science, Biochemistry, and Molecular Biology, respectively. The largest amount of published article has been found in the Frontiers in Plant Science journal, which has the highest global total citations and H-index. We also found that the journal of Plant Physiology has the highest local citations. From the most cited papers and references, the most important research focus was the improvement of crop yield and vegetation stress resistance. Furthermore, "drought" has been the most prominent keyword over the last 14 years, and more attention has been paid to "climate change" over the last 5 years. Under future climate change, how to regulate growth and development of food crops subjected to high temperature and drought stress may become a hotspot, and increasing research is critical to provide more insights into plant responses to high temperature and drought by linking plant above-below ground components. To summarize, this research will contribute to a comprehensive understanding of the past, present, and future research on plant responses to high temperature and drought.

Dynamic genome evolution in a model fern.

The large size and complexity of most fern genomes have hampered efforts to elucidate fundamental aspects of fern biology and land plant evolution through genome-enabled research. Here we present a chromosomal genome assembly and associated methylome, transcriptome and metabolome analyses for the model fern species Ceratopteris richardii. The assembly reveals a history of remarkably dynamic genome evolution including rapid changes in genome content and structure following the most recent whole-genome duplication approximately 60 million years ago. These changes include massive gene loss, rampant tandem duplications and multiple horizontal gene transfers from bacteria, contributing to the diversification of defence-related gene families. The insertion of transposable elements into introns has led to the large size of the Ceratopteris genome and to exceptionally long genes relative to other plants. Gene family analyses indicate that genes directing seed development were co-opted from those controlling the development of fern sporangia, providing insights into seed plant evolution. Our findings and annotated genome assembly extend the utility of Ceratopteris as a model for investigating and teaching plant biology.

Limited hydraulic recovery in seedlings of six tree species with contrasting leaf habits in subtropical China.

Subtropical tree species may experience severe drought stress due to variable rainfall under future climates. However, the capacity to restore hydraulic function post-drought might differ among co-occurring species with contrasting leaf habits (e.g., evergreen and deciduous) and have implications for future forest composition. Moreover, the links between hydraulic recovery and physiological and morphological traits related to water-carbon availability are still not well understood. Here, potted seedlings of six tree species (four evergreen and two deciduous) were grown outdoors under a rainout shelter. They grew under favorable water conditions until they were experimentally subjected to a soil water deficit leading to losses of ca. 50% of hydraulic conductivity, and then soils were re-watered to field capacity. Traits related to carbon and water relations were measured. There were differences in drought responses and recovery between species, but not as a function of evergreen or deciduous groups. Sapindus mukorossi exhibited the most rapid drought response, which was associated with a suite of physiological and morphological traits (larger plant size, the lowest hydraulic capacitance (C branch), higher minimum conductance (g min) and lower HV (Huber value)). Upon re-watering, xylem water potential exhibited fast recovery in 1-3 days among species, while photosynthesis at saturating light (A sat) and stomatal conductance (g s) recovery lagged behind water potential recovery depending on species, with g s recovery being more delayed than A sat in most species. Furthermore, none of the six species exhibited significant hydraulic recovery during the 7 days re-watering period, indicating that xylem refilling was apparently limited; in addition, NSC availability had a minimal role in facilitating hydraulic recovery during this short-term period. Collectively, if water supply is limited by insignificant hydraulic recovery post-drought, the observed carbon assimilation recovery of seedlings may not be sustained over the longer term, potentially altering seedling regeneration and shifting forest species composition in subtropical China under climate change.

High N Storage but Low N Recovery After Long-Term N-Fertilization in a Subtropical Cunninghamia lanceolata Plantation Ecosystem: A 14-Year Case Study.

Forests are among the most important N pools of all terrestrial ecosystems. Elevated atmospheric N deposition in recent decades has led to increased interest in the influences of N application on forest N cycles. However, accurate assessments of N storage in forest ecosystems remain elusive. We used a 14-year experiment of a Chinese fir [Cunninghamia lanceolata (Lamb.) Hook] plantation to explore how long-term N fertilization affected N storage and recovery rates. Our study plots were located in a field that had been continuously fertilized over 14 years (2004-2017) with urea at rates of 0 (N0, control), 60 (N60, low-N), 120 (N120, medium-N), and 240 (N240, high-N) kg N hm-2a-1. Data were collected that included N content and biomass in the understory, litter, and various plant organs (i.e., leaves, branches, stems, roots, and bark), as well as soil N content and density at different depths. Results showed that the total ecosystem N storage in the N-fertilized plots was 1.1-1.4 times higher than that in the control plots. About 12.36% of the total ecosystem N was stored in vegetation (plant organs, litter, and understory) and 87.64% was stored in soil (0-60 cm). Plant organs, litter, and soil had higher N storage than the understory layer. Significantly higher plant N uptake was found in the medium-N (1.2 times) and high-N (1.4 times) treatments relative to the control. The N recovery rate of the understory layer in the N-fertilized treatments was negative and less than that in the control. Application of long-term N fertilizer to this stand led to a low N recovery rate (average 11.39%) and high loss of N (average 91.86%), which indicate low N use efficiency in the Chinese fir plantation ecosystem. Our findings further clarify the distribution of N in an important terrestrial ecosystem and improve our understanding of regional N cycles.

Testing the limits of plant drought stress and subsequent recovery in four provenances of a widely distributed subtropical tree species.

Drought-induced tree mortality may increase with ongoing climate change. Unraveling the links between stem hydraulics and mortality thresholds, and the effects of intraspecific variation, remain important unresolved issues. We conducted a water manipulation experiment in a rain-out shelter, using four provenances of Schima superba originating from a gradient of annual precipitation (1124-1796 mm) and temperature (16.4-22.4°C). Seedlings were droughted to three levels of percentage loss of hydraulic conductivity (i.e., P50 , P88  and P99) and subsequently rewatered to field capacity for 30 days; traits related to water and carbon relations were measured. The lethal water potential associated with incipient mortality was between P50 and P88 . Seedlings exhibited similar drought responses in xylem water potential, hydraulic conductivity and gas exchange. Upon rehydration, patterns of gas exchange differed among provenances but were not related to the climate at the origin. The four provenances exhibited a similar degree of stem hydraulic recovery, which was correlated with the magnitude of antecedent drought and stem soluble sugar at the end of the drought. Results suggest that there were intraspecific differences in the capacity of S. superba seedlings for carbon assimilation during recovery, indicating a decoupling between gas exchange recovery and stem hydraulics across provenances.

Global transcriptomic analysis reveals candidate genes associated with different phosphorus acquisition strategies among soybean varieties.

Introduction: Soybean adapts to phosphorus-deficient soils through three important phosphorus acquisition strategies, namely altered root conformation, exudation of carboxylic acids, and symbiosis with clumping mycorrhizal fungi. However, the trade-offs and regulatory mechanisms of these three phosphorus acquisition strategies in soybean have not been researched. Methods: In this study, we investigated the responses of ten different soybean varieties to low soil phosphorus availability by determining biomass, phosphorus accumulation, root morphology, exudation, and mycorrhizal colonization rate. Furthermore, the molecular regulatory mechanisms underlying root phosphorus acquisition strategies were examined among varieties with different low-phosphorus tolerance using transcriptome sequencing and weighted gene co-expression network analysis. Results and discussion: The results showed that two types of phosphorus acquisition strategies-"outsourcing" and "do-it-yourself"-were employed by soybean varieties under low phosphorus availability. The "do-it-yourself" varieties, represented by QD11, Zh30, and Sd, obtained sufficient phosphorus by increasing their root surface area and secreting carboxylic acids. In contrast, the "outsourcing" varieties, represented by Zh301, Zh13, and Hc6, used increased symbiosis with mycorrhizae to obtain phosphorus owing to their large root diameters. Transcriptome analysis showed that the direction of acetyl-CoA metabolism could be the dividing line between the two strategies of soybean selection. ERF1 and WRKY1 may be involved in the regulation of phosphorus acquisition strategies for soybeans grown under low P environments. These findings will enhance our understanding of phosphorus acquisition strategies in soybeans. In addition, they will facilitate the development of breeding strategies that are more flexible to accommodate a variety of production scenarios in agriculture under low phosphorus environments. Background: Association of gastric atrophy or cancer with levels of serum pepsinogens, gastrin-17 and anti-Helicobacter pylori IgG antibody have been extensively studied. However, the association of serum pepsinogen and gastrin-17 with H. pylori infection has not been studied in a large population.

Stoichiometric traits (N:P) of understory plants contribute to reductions in plant diversity following long-term nitrogen addition in subtropical forest.

Nitrogen enrichment is pervasive in forest ecosystems, but its influence on understory plant communities and their stoichiometric characteristics is poorly understood. We hypothesize that when forest is enriched with nitrogen (N), the stoichiometric characteristics of plant species explain changes in understory plant diversity. A 13-year field experiment was conducted to explore the effects of N addition on foliar carbon (C): N: phosphorus (P) stoichiometry, understory plant species richness, and intrinsic water use efficiency (iWUE) in a subtropical Chinese fir forest. Four levels of N addition were applied: 0, 6, 12, and 24 g m-2 year-1. Individual plant species were categorized into resistant plants, intermediate resistant plants, and sensitive plants based on their response to nitrogen addition. Results showed that N addition significantly decreased the number of species, genera, and families of herbaceous plants. Foliar N:P ratios were greater in sensitive plants than resistant or intermediate resistant plants, while iWUE showed an opposite trend. However, no relationship was detected between soil available N and foliar N, and soil N:P and foliar N:P ratios. Our results indicated that long-term N addition decreased the diversity of understory plants in a subtropical forest. Through regulating water use efficiency with N addition, sensitive plants change their N:P stoichiometry and have a higher risk of mortality, while resistant plants maintain a stable N:P stoichiometry, which contributes to their survival. These findings suggest that plant N:P stoichiometry plays an important role in understory plant performance in response to environmental change of N.

Reduced photosynthetic thermal acclimation capacity under elevated ozone in poplar (Populus tremula) saplings.

The sensitivity of photosynthesis to temperature has been identified as a key uncertainty for projecting the magnitude of the terrestrial carbon cycle response to future climate change. Although thermal acclimation of photosynthesis under rising temperature has been reported in many tree species, whether tropospheric ozone (O3 ) affects the acclimation capacity remains unknown. In this study, temperature responses of photosynthesis (light-saturated rate of photosynthesis (Asat ), maximum rates of RuBP carboxylation (Vcmax ), and electron transport (Jmax ) and dark respiration (Rdark ) of Populus tremula exposed to ambient O3 (AO3 , maximum of 30 ppb) or elevated O3 (EO3 , maximum of 110 ppb) and ambient or elevated temperature (ambient +5°C) were investigated in solardomes. We found that the optimum temperature of Asat (ToptA ) significantly increased in response to warming. However, the thermal acclimation capacity was reduced by O3 exposure, as indicated by decreased ToptA , and temperature optima of Vcmax (ToptV ) and Jmax (ToptJ ) under EO3 . Changes in both stomatal conductance (gs ) and photosynthetic capacity (Vcmax and Jmax ) contributed to the shift of ToptA by warming and EO3 . Neither Rdark measured at 25°C ( R dark 25 ) nor the temperature response of Rdark was affected by warming, EO3 , or their combination. The responses of Asat , Vcmax , and Jmax to warming and EO3 were closely correlated with changes in leaf nitrogen (N) content and N use efficiency. Overall, warming stimulated growth (leaf biomass and tree height), whereas EO3 reduced growth (leaf and woody biomass). The findings indicate that thermal acclimation of Asat may be overestimated if the impact of O3 pollution is not taken into account.

Modeling Light Response of Electron Transport Rate and Its Allocation for Ribulose Biphosphate Carboxylation and Oxygenation.

Accurately describing the light response curve of electron transport rate (J-I curve) and allocation of electron flow for ribulose biphosphate (RuBP) carboxylation (J C-I curve) and that for oxygenation (J O-I curve) is fundamental for modeling of light relations of electron flow at the whole-plant and ecosystem scales. The non-rectangular hyperbolic model (hereafter, NH model) has been widely used to characterize light response of net photosynthesis rate (A n; A n-I curve) and J-I curve. However, NH model has been reported to overestimate the maximum A n (A nmax) and the maximum J (J max), largely due to its asymptotic function. Meanwhile, few efforts have been delivered for describing J C-I and J O-I curves. The long-standing challenge on describing A n-I and J-I curves have been resolved by a recently developed A n-I and J-I models (hereafter, Ye model), which adopt a nonasymptotic function. To test whether Ye model can resolve the challenge of NH model in reproducing J-I, J C-I and J O-I curves over light-limited, light-saturated, and photoinhibitory I levels, we compared the performances of Ye model and NH model against measurements on two C3 crops (Triticum aestivum L. and Glycine max L.) grown in field. The results showed that NH model significantly overestimated the A nmax and J max for both species, which can be accurately obtained by Ye model. Furthermore, NH model significantly overestimated the maximum electron flow for carboxylation (J C-max) but not the maximum electron flow for oxygenation (J O-max) for both species, disclosing the reason underlying the long-standing problem of NH model-overestimation of J max and A nmax.

Quantifying Light Response of Leaf-Scale Water-Use Efficiency and Its Interrelationships With Photosynthesis and Stomatal Conductance in C3 and C4 Species.

Light intensity (I) is the most dynamic and significant environmental variable affecting photosynthesis (A n), stomatal conductance (g s), transpiration (T r), and water-use efficiency (WUE). Currently, studies characterizing leaf-scale WUE-I responses are rare and key questions have not been answered. In particular, (1) What shape does the response function take? (2) Are there maximum intrinsic (WUEi; WUEi-max) and instantaneous WUE (WUEinst; WUEinst-max) at the corresponding saturation irradiances (I i-sat and I inst-sat)? This study developed WUEi-I and WUEinst-I models sharing the same non-asymptotic function with previously published A n-I and g s-I models. Observation-modeling intercomparison was conducted for field-grown plants of soybean (C3) and grain amaranth (C4) to assess the robustness of our models versus the non-rectangular hyperbola models (NH models). Both types of models can reproduce WUE-I curves well over light-limited range. However, at light-saturated range, NH models overestimated WUEi-max and WUEinst-max and cannot return I i-sat and I inst-sat due to its asymptotic function. Moreover, NH models cannot describe the down-regulation of WUE induced by high light, on which our models described well. The results showed that WUEi and WUEinst increased rapidly within low range of I, driven by uncoupled photosynthesis and stomatal responsiveness. Initial response rapidity of WUEi was higher than WUEinst because the greatest increase of A n and T r occurred at low g s. C4 species showed higher WUEi-max and WUEinst-max than C3 species-at similar I i-sat and I inst-sat. Our intercomparison highlighted larger discrepancy between WUEi-I and WUEinst-I responses in C3 than C4 species, quantitatively characterizing an important advantage of C4 photosynthetic pathway-higher A n gain but lower T r cost per unit of g s change. Our models can accurately return the wealth of key quantities defining species-specific WUE-I responses-besides A n-I and g s-I responses. The key advantage is its robustness in characterizing these entangled responses over a wide I range from light-limited to light-inhibitory light intensities, through adopting the same analytical framework and the explicit and consistent definitions on these responses. Our models are of significance for physiologists and modelers-and also for breeders screening for genotypes concurrently achieving maximized photosynthesis and optimized WUE.

Effects of exogenous 3-indoleacetic acid and cadmium stress on the physiological and biochemical characteristics of Cinnamomum camphora.

Indoleacetic acid (IAA) is a plant growth regulator that plays an important role in plant growth and development, and participates in the regulation of abiotic stress. To explore the effect of IAA on cadmium toxicity in Cinnamomum camphora, an indoor potted experiment was conducted with one-year-old C. camphora seedlings. The influence of IAA on cadmium accumulation, net photosynthetic rates, respiration, photosynthetic pigments (chlorophyll a, chlorophyll b, total chlorophyll and carotenoids), osmoregulatory substances (proline, soluble sugar and soluble protein) and the malondialdehyde content in C. camphora leaves treated with 30 mg kg-1 cadmium was analysed with or without the addition of 10 mg kg-1 IAA. Cadmium accumulation in the leaves of C. camphora with the addition of exogenous IAA was significantly higher than accumulation during cadmium stress without additional IAA (ca 69.10% after 60 days' incubation). During the culture period, the net photosynthetic rate in C. camphora leaves subjected to cadmium stress without the addition of IAA was up to 24.31% lower than that of control plants. The net photosynthetic rate in C. camphora leaves subjected to cadmium stress and addition of IAA was up to 30.31% higher than that of leaves subjected to cadmium stress without the addition of IAA. Chlorophyll a, total chlorophyll and carotenoid contents in the cadmium-stressed leaves without the addition of IAA were lower than those in the control treatment. The presence of IAA increased the chlorophyll a, total chlorophyll and carotenoid contents relative to the cadmium stress without the addition of IAA. The respiration rate and concentrations of proline, soluble sugar, soluble protein and malondialdehyde in C. camphora leaves subjected to cadmium stress without the addition of IAA were higher than those in the control. The addition of IAA reduced the respiration rate, and the concentrations of proline, soluble sugar, soluble protein and malondialdehyde in C. camphora leaves when compared with the cadmium stress without the addition of IAA. These results indicate that exogenous IAA improves photosynthetic performance and the growth environment of C. camphora by enhancing the net photosynthetic rate, increasing concentrations of osmoregulatory substances, removing reactive oxygen radicals and eliminating potential damage, thereby reducing the toxic effects of cadmium on C. camphora.

Effects of elevated carbon dioxide and elevated temperature on morphological, physiological and anatomical responses of Eucalyptus tereticornis along a soil phosphorus gradient.

Eucalypts are likely to play a critical role in the response of Australian forests to rising atmospheric CO2 concentration ([CO2]) and temperature. Although eucalypts are frequently phosphorus (P) limited in native soils, few studies have examined the main and interactive effects of P availability, [CO2] and temperature on eucalypt morphology, physiology and anatomy. To address this issue, we grew seedlings of Eucalyptus tereticornis Smith across its P-responsive range (6-500 mg kg-1) for 120 days under two [CO2] (ambient: 400 µmol mol-1 (Ca) and elevated: 640 µmol mol-1 (Ce)) and two temperature (ambient: 24/16 °C (Ta) and elevated: 28/20 °C (Te) day/night) treatments in a sunlit glasshouse. Seedlings were well-watered and supplied with otherwise non-limiting macro- and micro-nutrients. Increasing soil P supply increased growth responses to Ce and Te. At the highest P supplies, Ce increased total dry mass, leaf number and total leaf area by ~50%, and Te increased leaf number by ~40%. By contrast, Ce and Te had limited effects on seedling growth at the lowest P supply. Soil P supply did not consistently modify photosynthetic responses to Ce or Te. Overall, effects of Ce and Te on growth, physiological and anatomical responses of E. tereticornis seedlings were generally neutral or negative at low soil P supply, suggesting that native tree responses to future climates may be relatively small in native low-P soils in Australian forests.

Comparing two measures of leaf photorespiration rate across a wide range of light intensities.

Suppression of photorespiration by low O2 concentrations (Method 1) and simultaneous measurements of gas exchange and chlorophyll fluorescence (Method 2) are often used to estimate leaf photorespiration rate (Rp) of C3 plants. However, it is largely unknown whether Method 1 and Method 2 can be used equivalently in estimating Rp. Using a field experiment on two wheat cultivars (T. aestivum JM22 and T. aestivum Z39-118) whose leaf gas exchange and chlorophyll fluorescence at low and normal O2 concentrations (2% versus 21% O2) were simultaneously measured across a wide range of light intensities (I), this study assessed the impacts of the two measures on Rp and its response under changing irradiance conditions. All the above quantities increased with the increasing I until reaching the cultivar-specific maximum values and the corresponding saturation light intensities. However, there were significant differences between Rp estimated by Method 1 and Method 2 at the I range from 150 to 2000 µmol m-2 s-1 for T. aestivum JM22 and from 150 to 1000 µmol m-2 s-1 for T. aestivum Z39-118. These findings demonstrated that the two methods cannot be used equivalently under changing irradiance conditions.

CO2 and temperature effects on morphological and physiological traits affecting risk of drought-induced mortality.

Despite a wealth of eco-physiological assessments of plant response to extreme drought, few studies have addressed the interactive effects of global change factors on traits driving mortality. To understand the interaction between hydraulic and carbon metabolic traits influencing tree mortality, which may be independently influenced by atmospheric [CO2] and temperature, we grew Eucalyptus sideroxylon A. Cunn. ex Woolls from seed in a full-factorial [CO2] (280, 400 and 640 µmol mol-1, Cp, Ca and Ce, respectively) and temperature (ambient and ambient +4 °C, Ta and Te, respectively) experiment. Prior to drought, growth across treatment combinations resulted in significant variation in physiological and morphological traits, including photosynthesis (Asat), respiration (Rd), stomatal conductance, carbohydrate storage, biomass and leaf area (LA). Ce increased Asat, LA and leaf carbohydrate concentration compared with Ca, while Cp generated the opposite response; Te reduced Rd. However, upon imposition of drought, Te hastened mortality (9 days sooner compared with Ta), while Ce significantly exacerbated drought stress when combined with Te. Across treatments, earlier time-to-mortality was mainly associated with lower (more negative) leaf water potential (Ψl) during the initial drought phase, along with higher water loss across the first 3 weeks of water limitation. Among many variables, Ψl was more important than carbon status in predicting time-to-mortality across treatments, yet leaf starch was associated with residual variation within treatments. These results highlight the need to carefully consider the integration, interaction and hierarchy of traits contributing to mortality, along with their responses to environmental drivers. Both morphological traits, which influence soil resource extraction, and physiological traits, which affect water-for-carbon exchange to the atmosphere, must be considered to adequately predict plant response to drought. Researchers have struggled with assessing the relative importance of hydraulic and carbon metabolic traits in determining mortality, yet an integrated trait, time-dependent framework provides considerable insight into the risk of death from drought for trees.

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.