Carbon-phosphorus cycle models overestimate CO2 enrichment response in a mature Eucalyptus forest.

The importance of phosphorus (P) in regulating ecosystem responses to climate change has fostered P-cycle implementation in land surface models, but their CO2 effects predictions have not been evaluated against measurements. Here, we perform a data-driven model evaluation where simulations of eight widely used P-enabled models were confronted with observations from a long-term free-air CO2 enrichment experiment in a mature, P-limited Eucalyptus forest. We show that most models predicted the correct sign and magnitude of the CO2 effect on ecosystem carbon (C) sequestration, but they generally overestimated the effects on plant C uptake and growth. We identify leaf-to-canopy scaling of photosynthesis, plant tissue stoichiometry, plant belowground C allocation, and the subsequent consequences for plant-microbial interaction as key areas in which models of ecosystem C-P interaction can be improved. Together, this data-model intercomparison reveals data-driven insights into the performance and functionality of P-enabled models and adds to the existing evidence that the global CO2-driven carbon sink is overestimated by models.

Microbial competition for phosphorus limits the CO2 response of a mature forest.

The capacity for terrestrial ecosystems to sequester additional carbon (C) with rising CO2 concentrations depends on soil nutrient availability1,2. Previous evidence suggested that mature forests growing on phosphorus (P)-deprived soils had limited capacity to sequester extra biomass under elevated CO2 (refs. 3-6), but uncertainty about ecosystem P cycling and its CO2 response represents a crucial bottleneck for mechanistic prediction of the land C sink under climate change7. Here, by compiling the first comprehensive P budget for a P-limited mature forest exposed to elevated CO2, we show a high likelihood that P captured by soil microorganisms constrains ecosystem P recycling and availability for plant uptake. Trees used P efficiently, but microbial pre-emption of mineralized soil P seemed to limit the capacity of trees for increased P uptake and assimilation under elevated CO2 and, therefore, their capacity to sequester extra C. Plant strategies to stimulate microbial P cycling and plant P uptake, such as increasing rhizosphere C release to soil, will probably be necessary for P-limited forests to increase C capture into new biomass. Our results identify the key mechanisms by which P availability limits CO2 fertilization of tree growth and will guide the development of Earth system models to predict future long-term C storage.

[Short-term nitrogen deposition changes chemical composition of litter and soil organic matter in a Moso bamboo forest]. / 短期氮沉降改变毛竹林凋落物和土壤有机质化学组成.

To investigate the effects of short-term nitrogen (N) deposition on organic matter composition of litter and soil in Moso bamboo (Phyllostachys edulis) forests, we established a N-addition treatments (50 kg N·hm-2·a-1) to simulate the ambient and N deposition in a subtropical Moso bamboo forest from July 2020 to January 2022. We analyzed the organic matter composition of Moso bamboo leaf/root litter and soil by using pyrolysis-gas chromatography/mass spectrometry (Py-GC/MS) technique. The results showed that short-term N deposition significantly increased the relative content of soil phenols by 50.9%, while significantly decreased fatty acids by 26.3%. The rela-tive content of alkanes & alkenes and lignin in leaf litter was significantly increased by 51.9% and 33.5%, respectively, while that of phenols and polysaccharides significantly decreased by 52.2% and 56.3%. In root litter, eleva-ted N significantly decreased the relative content of polycyclic aromatic hydrocarbons by 16.6%. Moreover, the relative content of fatty acids in soil organic matter was significantly positively correlated with the relative content of poly-saccharides in leaf litter. The relative content of phenols in soil organic matter was significantly positively correlated with the relative content of lignin, and negatively correlated with the relative content of polysaccharides in leaf litter. Our results demonstrated that short-term N deposition did not change the concentration of total organic carbon, total nitrogen, and C/N of the soil, leaf litter, and root litter, but significantly altered the chemical composition of organic matter. In addition, the changes in chemical composition of organic matter in soil under short-term N deposition were affected by the composition of organic matter in leaf litter.

Convergence of carbon sink magnitude and water table depth in global wetlands.

Wetlands are strategic areas for carbon uptake, but accurate assessments of their sequestration ability are limited by the uncertainty and variability in their carbon balances. Based on 2385 observations of annual net ecosystem production from global wetlands, we show that the mean net carbon sinks of inland wetlands, peatlands and coastal wetlands are 0.57, 0.29 and 1.88 tons of carbon per hectare per year, respectively, with a mean value of 0.57 tons of carbon per hectare per year weighted by the distribution area of different wetland types. Carbon sinks are mainly in Asia and North America. Within and across wetland types, we find that water table depth (WTD) exerts greater control than climate- and ecosystem-related variables, and an increase in WTD results in a stronger carbon sink. Our results highlight an urgent need to sustain wetland hydrology under global change; otherwise, wetlands are at high risk of becoming carbon sources to the atmosphere.

Optimal stomatal theory predicts CO2 responses of stomatal conductance in both gymnosperm and angiosperm trees.

Optimal stomatal theory predicts that stomata operate to maximise photosynthesis (Anet ) and minimise transpirational water loss to achieve optimal intrinsic water-use efficiency (iWUE). We tested whether this theory can predict stomatal responses to elevated atmospheric CO2 (eCO2 ), and whether it can capture differences in responsiveness among woody plant functional types (PFTs). We conducted a meta-analysis of tree studies of the effect of eCO2 on iWUE and its components Anet and stomatal conductance (gs ). We compared three PFTs, using the unified stomatal optimisation (USO) model to account for confounding effects of leaf-air vapour pressure difference (D). We expected smaller gs , but greater Anet , responses to eCO2 in gymnosperms compared with angiosperm PFTs. We found that iWUE increased in proportion to increasing eCO2 in all PFTs, and that increases in Anet had stronger effects than reductions in gs . The USO model correctly captured stomatal behaviour with eCO2 across most datasets. The chief difference among PFTs was a lower stomatal slope parameter (g1 ) for the gymnosperm, compared with angiosperm, species. Land surface models can use the USO model to describe stomatal behaviour under changing atmospheric CO2 conditions.

Canopy dieback and recovery in Australian native forests following extreme drought.

In 2019, south-eastern Australia experienced its driest and hottest year on record, resulting in massive canopy dieback events in eucalypt dominated forests. A subsequent period of high precipitation in 2020 provided a rare opportunity to quantify the impacts of extreme drought and consequent recovery. We quantified canopy health and hydraulic impairment (native percent loss of hydraulic conductivity, PLC) of 18 native tree species growing at 15 sites that were heavily impacted by the drought both during and 8-10 months after the drought. Most species exhibited high PLC during drought (PLC:65.1 ± 3.3%), with no clear patterns across sites or species. Heavily impaired trees (PLC > 70%) showed extensive canopy browning. In the post-drought period, most surviving trees exhibited hydraulic recovery (PLC:26.1 ± 5.1%), although PLC remained high in some trees (50-70%). Regained hydraulic function (PLC < 50%) corresponded to decreased canopy browning indicating improved tree health. Similar drought (37.1 ± 4.2%) and post-drought (35.1 ± 4.4%) percentages of basal area with dead canopy suggested that trees with severely compromised canopies immediately after drought were not able to recover. This dataset provides insights into the impacts of severe natural drought on the health of mature trees, where hydraulic failure is a major contributor in canopy dieback and tree mortality during extreme drought events.

Unlocking Drought-Induced Tree Mortality: Physiological Mechanisms to Modeling.

Drought-related tree mortality has become a major concern worldwide due to its pronounced negative impacts on the functioning and sustainability of forest ecosystems. However, our ability to identify the species that are most vulnerable to drought, and to pinpoint the spatial and temporal patterns of mortality events, is still limited. Model is useful tools to capture the dynamics of vegetation at spatiotemporal scales, yet contemporary land surface models (LSMs) are often incapable of predicting the response of vegetation to environmental perturbations with sufficient accuracy, especially under stressful conditions such as drought. Significant progress has been made regarding the physiological mechanisms underpinning plant drought response in the past decade, and plant hydraulic dysfunction has emerged as a key determinant for tree death due to water shortage. The identification of pivotal physiological events and relevant plant traits may facilitate forecasting tree mortality through a mechanistic approach, with improved precision. In this review, we (1) summarize current understanding of physiological mechanisms leading to tree death, (2) describe the functionality of key hydraulic traits that are involved in the process of hydraulic dysfunction, and (3) outline their roles in improving the representation of hydraulic function in LSMs. We urge potential future research on detailed hydraulic processes under drought, pinpointing corresponding functional traits, as well as understanding traits variation across and within species, for a better representation of drought-induced tree mortality in models.

Bridge to the future: Important lessons from 20 years of ecosystem observations made by the OzFlux network.

In 2020, the Australian and New Zealand flux research and monitoring network, OzFlux, celebrated its 20th anniversary by reflecting on the lessons learned through two decades of ecosystem studies on global change biology. OzFlux is a network not only for ecosystem researchers, but also for those 'next users' of the knowledge, information and data that such networks provide. Here, we focus on eight lessons across topics of climate change and variability, disturbance and resilience, drought and heat stress and synergies with remote sensing and modelling. In distilling the key lessons learned, we also identify where further research is needed to fill knowledge gaps and improve the utility and relevance of the outputs from OzFlux. Extreme climate variability across Australia and New Zealand (droughts and flooding rains) provides a natural laboratory for a global understanding of ecosystems in this time of accelerating climate change. As evidence of worsening global fire risk emerges, the natural ability of these ecosystems to recover from disturbances, such as fire and cyclones, provides lessons on adaptation and resilience to disturbance. Drought and heatwaves are common occurrences across large parts of the region and can tip an ecosystem's carbon budget from a net CO2 sink to a net CO2 source. Despite such responses to stress, ecosystems at OzFlux sites show their resilience to climate variability by rapidly pivoting back to a strong carbon sink upon the return of favourable conditions. Located in under-represented areas, OzFlux data have the potential for reducing uncertainties in global remote sensing products, and these data provide several opportunities to develop new theories and improve our ecosystem models. The accumulated impacts of these lessons over the last 20 years highlights the value of long-term flux observations for natural and managed systems. A future vision for OzFlux includes ongoing and newly developed synergies with ecophysiologists, ecologists, geologists, remote sensors and modellers.

Corrigendum: Unlocking drought-induced tree mortality: Physiological mechanisms to modelling.

[This corrects the article DOI: 10.3389/fpls.2022.835921.].

Drought by CO2 interactions in trees: a test of the water savings mechanism.

Elevated atmospheric CO2 (eCa ) may benefit plants during drought by reducing stomatal conductance (gs ) but any 'water savings effect' could be neutralized by concurrent stimulation of leaf area. We investigated whether eCa enhanced water savings, thereby ameliorating the impact of drought on carbon and water relations in trees. We report leaf-level gas exchange and whole-plant and soil water relations during a short-term dry-down in two Eucalyptus species with contrasting drought tolerance. Plants had previously been established for 9 to 11 months in steady-state conditions of ambient atmospheric CO2 (aCa ) and eCa , with half of each treatment group exposed to sustained drought for 5 to 7 months. The lower stomatal conductance under eCa did not lead to soil moisture savings during the dry-down due to the counteractive effect of increased whole-plant leaf area. Nonetheless, eCa -grown plants maintained higher photosynthetic rates and leaf water potentials, making them less stressed during the dry-down, despite being larger. These effects were more pronounced in the xeric species than the mesic species, and in previously water-stressed plants. Our findings indicate that eCa may enhance plant performance during drought despite a lack of soil water savings, especially in species with more conservative growth and water-use strategies.

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2.

Atmospheric carbon dioxide concentration ([CO2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO2 ] increase and thus climate change. However, ecosystem CO2 responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO2 ]-driven terrestrial carbon sink can appear contradictory. Here we synthesize theory and broad, multidisciplinary evidence for the effects of increasing [CO2 ] (iCO2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industrial times. Established theory, supported by experiments, indicates that iCO2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO2 responses are high in comparison to experiments and predictions from theory. Plant mortality and soil carbon iCO2 responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

Low phosphorus supply constrains plant responses to elevated CO2 : A meta-analysis.

Phosphorus (P) is an essential macro-nutrient required for plant metabolism and growth. Low P availability could potentially limit plant responses to elevated carbon dioxide (eCO2 ), but consensus has yet to be reached on the extent of this limitation. Here, based on data from experiments that manipulated both CO2 and P for young individuals of woody and non-woody species, we present a meta-analysis of P limitation impacts on plant growth, physiological, and morphological response to eCO2 . We show that low P availability attenuated plant photosynthetic response to eCO2 by approximately one-quarter, leading to a reduced, but still positive photosynthetic response to eCO2 compared to those under high P availability. Furthermore, low P limited plant aboveground, belowground, and total biomass responses to eCO2 , by 14.7%, 14.3%, and 12.4%, respectively, equivalent to an approximate halving of the eCO2 responses observed under high P availability. In comparison, low P availability did not significantly alter the eCO2 -induced changes in plant tissue nutrient concentration, suggesting tissue nutrient flexibility is an important mechanism allowing biomass response to eCO2 under low P availability. Low P significantly reduced the eCO2 -induced increase in leaf area by 14.3%, mirroring the aboveground biomass response, but low P did not affect the eCO2 -induced increase in root length. Woody plants exhibited stronger attenuation effect of low P on aboveground biomass response to eCO2 than non-woody plants, while plants with different mycorrhizal associations showed similar responses to low P and eCO2 interaction. This meta-analysis highlights crucial data gaps in capturing plant responses to eCO2 and low P availability. Field-based experiments with longer-term exposure of both CO2 and P manipulations are critically needed to provide ecosystem-scale understanding. Taken together, our results provide a quantitative baseline to constrain model-based hypotheses of plant responses to eCO2 under P limitation, thereby improving projections of future global change impacts.

The fate of carbon in a mature forest under carbon dioxide enrichment.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1-5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3-5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7-10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7-11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests.

Nitrogen and Phosphorus Retranslocation of Leaves and Stemwood in a Mature Eucalyptus Forest Exposed to 5 Years of Elevated CO2.

Elevated CO2 affects C cycling processes which in turn can influence the nitrogen (N) and phosphorus (P) concentrations of plant tissues. Given differences in how N and P are used by plants, we asked if their stoichiometry in leaves and wood was maintained or altered in a long-term elevated CO2 experiment in a mature Eucalyptus forest on a low P soil (EucFACE). We measured N and P concentrations in green leaves at different ages at the top of mature trees across 6 years including 5 years in elevated CO2. N and P concentrations in green and senesced leaves and wood were determined to evaluate both spatial and temporal variation of leaf N and P concentrations, including the N and P retranslocation in leaves and wood. Leaf P concentrations were 32% lower in old mature leaves compared to newly flushed leaves with no effect of elevated CO2 on leaf P. By contrast, elevated CO2 significantly decreased leaf N concentrations in newly flushed leaves but this effect disappeared as leaves matured. As such, newly flushed leaves had 9% lower N:P ratios in elevated CO2 and N:P ratios were not different in mature green leaves (CO2 by Age effect, P = 0.02). Over time, leaf N and P concentrations in the upper canopy slightly declined in both CO2 treatments compared to before the start of the experiment. P retranslocation in leaves was 50%, almost double that of N retranslocation (29%), indicating that this site was P-limited and that P retranslocation was an important mechanism in this ecosystem to retain P in plants. As P-limited trees tend to store relatively more N than P, we found an increased N:P ratio in sapwood in response to elevated CO2 (P < 0.01), implying N accumulation in live wood. The flexible stoichiometric ratios we observed can have important implications for how plants adjust to variable environmental conditions including climate change. Hence, variable nutrient stoichiometry should be accounted for in large-scale Earth Systems models invoking biogeochemical processes.

Towards a more physiological representation of vegetation phosphorus processes in land surface models.

Contents Summary 1223 I. Introduction 1223 II. Photosynthesis and respiration 1224 III. Biomass growth 1224 IV. Carbon allocation 1225 V. Plant internal P redistribution 1226 VI. Plant P uptake 1227 VII. Conclusion 1227 Acknowledgements 1228 References 1228 SUMMARY: Our ability to understand the effect of nutrient limitation on ecosystem productivity is key to the prediction of future terrestrial carbon storage. Significant progress has been made to include phosphorus (P) cycle processes in land surface models (LSMs), but these efforts are focused on the soil component of the P cycle. Incorporating the soil component is important to estimate plant-available P, but does not necessarily address the vegetation response to P limitation or plant-soil interactions. A more detailed representation of plant P processes is needed to link nutrient availability and ecosystem productivity. We review physiological and biochemical evidence for vegetation responses to P availability, and recommend ways to move towards a more physiological representation of vegetation P processes in LSMs.

Trees tolerate an extreme heatwave via sustained transpirational cooling and increased leaf thermal tolerance.

Heatwaves are likely to increase in frequency and intensity with climate change, which may impair tree function and forest C uptake. However, we have little information regarding the impact of extreme heatwaves on the physiological performance of large trees in the field. Here, we grew Eucalyptus parramattensis trees for 1 year with experimental warming (+3°C) in a field setting, until they were greater than 6 m tall. We withheld irrigation for 1 month to dry the surface soils and then implemented an extreme heatwave treatment of 4 consecutive days with air temperatures exceeding 43°C, while monitoring whole-canopy exchange of CO2 and H2 O, leaf temperatures, leaf thermal tolerance, and leaf and branch hydraulic status. The heatwave reduced midday canopy photosynthesis to near zero but transpiration persisted, maintaining canopy cooling. A standard photosynthetic model was unable to capture the observed decoupling between photosynthesis and transpiration at high temperatures, suggesting that climate models may underestimate a moderating feedback of vegetation on heatwave intensity. The heatwave also triggered a rapid increase in leaf thermal tolerance, such that leaf temperatures observed during the heatwave were maintained within the thermal limits of leaf function. All responses were equivalent for trees with a prior history of ambient and warmed (+3°C) temperatures, indicating that climate warming conferred no added tolerance of heatwaves expected in the future. This coordinated physiological response utilizing latent cooling and adjustment of thermal thresholds has implications for tree tolerance of future climate extremes as well as model predictions of future heatwave intensity at landscape and global scales.

Predictability of Precipitation Over the Conterminous U.S. Based on the CMIP5 Multi-Model Ensemble.

Characterizing precipitation seasonality and variability in the face of future uncertainty is important for a well-informed climate change adaptation strategy. Using the Colwell index of predictability and monthly normalized precipitation data from the Coupled Model Intercomparison Project Phase 5 (CMIP5) multi-model ensembles, this study identifies spatial hotspots of changes in precipitation predictability in the United States under various climate scenarios. Over the historic period (1950-2005), the recurrent pattern of precipitation is highly predictable in the East and along the coastal Northwest, and is less so in the arid Southwest. Comparing the future (2040-2095) to the historic period, larger changes in precipitation predictability are observed under Representative Concentration Pathways (RCP) 8.5 than those under RCP 4.5. Finally, there are region-specific hotspots of future changes in precipitation predictability, and these hotspots often coincide with regions of little projected change in total precipitation, with exceptions along the wetter East and parts of the drier central West. Therefore, decision-makers are advised to not rely on future total precipitation as an indicator of water resources. Changes in precipitation predictability and the subsequent changes on seasonality and variability are equally, if not more, important factors to be included in future regional environmental assessment.