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Phenology is the study of periodic biological occurrences and can provide important insights into the influence of climatic variability and change on ecosystems. Understanding Australia's vegetation phenology is a challenge due to its diverse range of ecosystems, from savannas and tropical rainforests to temperate eucalypt woodlands, semiarid scrublands, and alpine grasslands. These ecosystems exhibit marked differences in seasonal patterns of canopy development and plant life-cycle events, much of which deviates from the predictable seasonal phenological pulse of temperate deciduous and boreal biomes. Many Australian ecosystems are subject to irregular events (i.e. drought, flooding, cyclones, and fire) that can alter ecosystem composition, structure, and functioning just as much as seasonal change. We show how satellite remote sensing and ground-based digital repeat photography (i.e. phenocams) can be used to improve understanding of phenology in Australian ecosystems. First, we examine temporal variation in phenology on the continental scale using the enhanced vegetation index (EVI), calculated from MODerate resolution Imaging Spectroradiometer (MODIS) data. Spatial gradients are revealed, ranging from regions with pronounced seasonality in canopy development (i.e. tropical savannas) to regions where seasonal variation is minimal (i.e. tropical rainforests) or high but irregular (i.e. arid ecosystems). Next, we use time series colour information extracted from phenocam imagery to illustrate a range of phenological signals in four contrasting Australian ecosystems. These include greening and senescing events in tropical savannas and temperate eucalypt understorey, as well as strong seasonal dynamics of individual trees in a seemingly static evergreen rainforest. We also demonstrate how phenology links with ecosystem gross primary productivity (from eddy covariance) and discuss why these processes are linked in some ecosystems but not others. We conclude that phenocams have the potential to greatly improve the current understanding of Australian ecosystems. To facilitate the sharing of this information, we have formed the Australian Phenocam Network (http://phenocam.org.au/).

Key Message: Heat stress (HS) under well-watered conditions was not detrimental to leaf photosynthesis or yield but modified the elevated CO 2 response of photosynthesis and yield in two contrasting wheat cultivars. Climate change is increasing the frequency of extreme events such as heat waves, adversely affecting crop productivity. While positive impacts of elevated carbon dioxide (eCO 2 ) on crop productivity are evident, the interactive effects of eCO 2 and environmental stresses are still unclear. To investigate the interactive effects of elevated CO 2 and heat stress (HS), we grew two contrasting wheat cultivars, early-maturing Scout and high-tillering Yitpi, under non-limiting water and nutrients at ambient (aCO 2 , 450 ppm) or elevated (eCO 2 , 650 ppm) CO 2 and 22 °C in the glasshouse. Plants were exposed to two 3-day HS cycles at the vegetative (38.1 °C) and/or flowering (33.5 °C) stage. At aCO 2 , both wheat cultivars showed similar responses of photosynthesis and mesophyll conductance to temperature and produced similar grain yield. Relative to aCO 2 , eCO 2 enhanced photosynthesis rate and reduced stomatal conductance and maximal carboxylation rate (V cmax ). During HS, high temperature stimulated photosynthesis at eCO 2 in both cultivars, while eCO 2 stimulated photosynthesis in Scout. Electron transport rate (J max ) was unaffected by any treatment. eCO 2 equally enhanced biomass and grain yield of both cultivars in control, but not HS, plants. HS reduced biomass and yield of Scout at eCO 2 . Yitpi, the cultivar with higher grain nitrogen, underwent a trade-off between grain yield and nitrogen. In conclusion, eCO 2 improved photosynthesis of control and HS wheat, and improved biomass and grain yield of control plants only. Under well-watered conditions, HS was not detrimental to photosynthesis or growth but precluded a yield response to eCO 2 ., (© 2022. The Author(s).)

Data capturing multiple axes of tree size and shape, such as a tree's stem diameter, height and crown size, underpin a wide range of ecological research-from developing and testing theory on forest structure and dynamics, to estimating forest carbon stocks and their uncertainties, and integrating remote sensing imagery into forest monitoring programmes. However, these data can be surprisingly hard to come by, particularly for certain regions of the world and for specific taxonomic groups, posing a real barrier to progress in these fields. To overcome this challenge, we developed the Tallo database, a collection of 498,838 georeferenced and taxonomically standardized records of individual trees for which stem diameter, height and/or crown radius have been measured. These data were collected at 61,856 globally distributed sites, spanning all major forested and non-forested biomes. The majority of trees in the database are identified to species (88%), and collectively Tallo includes data for 5163 species distributed across 1453 genera and 187 plant families. The database is publicly archived under a CC-BY 4.0 licence and can be access from: https://doi.org/10.5281/zenodo.6637599. To demonstrate its value, here we present three case studies that highlight how the Tallo database can be used to address a range of theoretical and applied questions in ecology-from testing the predictions of metabolic scaling theory, to exploring the limits of tree allometric plasticity along environmental gradients and modelling global variation in maximum attainable tree height. In doing so, we provide a key resource for field ecologists, remote sensing researchers and the modelling community working together to better understand the role that trees play in regulating the terrestrial carbon cycle., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Atmospheric carbon dioxide enrichment (eCO 2 ) can enhance plant carbon uptake and growth 1-5 , thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO 2 concentration 6 . Although evidence gathered from young aggrading forests has generally indicated a strong CO 2 fertilization effect on biomass growth 3-5 , it is unclear whether mature forests respond to eCO 2 in a similar way. In mature trees and forest stands 7-10 , photosynthetic uptake has been found to increase under eCO 2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO 2 unclear 4,5,7-11 . Here using data from the first ecosystem-scale Free-Air CO 2 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 eCO 2 exposure. We show that, although the eCO 2 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 eCO 2 , and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO 2 fertilization as a driver of increased carbon sinks in global forests.

The classical theory of stomatal optimization stipulates that stomata should act to maximize photosynthesis while minimizing transpiration. This theory, despite its remarkable success in reproducing empirical patterns, does not account for the fact that the available water to plants is dynamically regulated by plants themselves, and that plants compete for water in most locations. Here, we develop an alternative theory in which plants maximize the expected carbon gain under stochastic rainfall in a competitive environment. We further incorporate xylem hydraulic limitation as an additional constraint to transpiration and evaluate its impacts on stomatal optimization by incorporating the direct carbon cost of xylem recovery and the opportunity cost of reduced future photosynthesis as a result of irrecoverable xylem damage. We predict stomatal behaviour to be more conservative with a higher cost induced by xylem damage. By varying the unit carbon cost and extent of xylem recovery, characterizing the direct and opportunity cost of xylem damage, respectively, our model can reproduce several key patterns of stomatal-hydraulic trait covariations. By addressing the key elements of water limitation in plant gas exchange simultaneously, including plants' self-regulation of water availability, competition for water and hydraulic risk, our study provides a comprehensive theoretical basis for understanding stomatal behaviour., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Hot days are becoming hotter and more frequent, threatening wheat yields worldwide. Developing wheat varieties ready for future climates calls for improved understanding of how elevated CO2 (eCO2) and heat stress (HS) interactively impact wheat yields. We grew a modern, high-yielding wheat cultivar (Scout) at ambient CO2 (aCO2, 419 μl l -1) or eCO2 (654 μl l-1) in a glasshouse maintained at 22/15 °C (day/night). Half of the plants were exposed to HS (40/24 °C) for 5 d at anthesis. In non-HS plants, eCO2 enhanced (+36%) CO2 assimilation rates (Asat) measured at growth CO2 despite down-regulation of photosynthetic capacity. HS reduced Asat (-42%) in aCO2- but not in eCO2-grown plants because eCO2 protected photosynthesis by increasing ribulose bisphosphate regeneration capacity and reducing photochemical damage under HS. eCO2 stimulated biomass (+35%) of all plants and grain yield (+30%) of non-HS plants only. Plant biomass initially decreased following HS but recovered at maturity due to late tillering. HS equally reduced grain yield (-40%) in aCO2- and eCO2-grown plants due to grain abortion and reduced grain filling. While eCO2 mitigated the negative impacts of HS at anthesis on wheat photosynthesis and biomass, grain yield was reduced by HS in both CO2 treatments., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Catastrophic failure of the water transport pathway in trees is a principal mechanism of mortality during extreme drought. To be able to predict the probability of mortality at an individual and landscape scale we need knowledge of the time for plants to reach critical levels of hydraulic failure. We grew plants of eight species of Eucalyptus originating from contrasting climates before allowing a subset to dehydrate. We tested whether a trait-based model of time to plant desiccation t crit , from stomatal closure g s90 to a critical level of hydraulic dysfunction Ψ crit is consistent with observed dry-down times. Plant desiccation time varied among species, ranging from 96.2 to 332 h at a vapour-pressure deficit of 1 kPa, and was highly predictable using the t crit model in conjunction with a leaf shedding function. Plant desiccation time was longest in species with high cavitation resistance, strong vulnerability segmentation, wide stomatal-hydraulic safety, and a high ratio of total plant water content to leaf area. Knowledge of t crit in combination with water-use traits that influence stomatal closure could significantly increase our ability to predict the timing of drought-induced mortality at tree and forest scales., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Contents Summary 693 I. Introduction 693 II. Comparison of various definitions and measurement techniques of minimum conductance 694 III. Cuticular conductance 695 IV. Contribution of stomata 696 V. Environmental and ecological variation in minimum conductance 696 VI. Use of minimum conductance in models 698 VII. Conclusions 703 Acknowledgements 703 References 703 SUMMARY: When the rate of photosynthesis is greatly diminished, such as during severe drought, extreme temperature or low light, it seems advantageous for plants to close stomata and completely halt water loss. However, water loss continues through the cuticle and incompletely closed stomata, together constituting the leaf minimum conductance (g min ). In this review, we critically evaluate the sources of variation in g min , quantitatively compare various methods for its estimation, and illustrate the role of g min in models of leaf gas exchange. A literature compilation of g min as measured by the weight loss of detached leaves is presented, which shows much variation in this trait, which is not clearly related to species groups, climate of origin or leaf type. Much evidence points to the idea that g min is highly responsive to the growing conditions of the plant, including soil water availability, temperature and air humidity - as we further demonstrate with two case studies. We pay special attention to the role of the minimum conductance in the Ball-Berry model of stomatal conductance, and caution against the usual regression-based method for its estimation. The synthesis presented here provides guidelines for the use of g min in ecosystem models, and points to clear research gaps for this drought tolerance trait., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Centrifuge-based techniques to assess xylem vulnerability to embolism are increasingly being used, although we are yet to reach a consensus on the nature and extent of artefactual embolism observed in some angiosperm species. In particular, there is disagreement over whether these artefacts influence both the spin (Cavitron) and static versions of the centrifuge technique equally. We tested two methods for inducing embolism: bench dehydration and centrifugation. We used three methods to measure the resulting loss of conductivity: gravimetric flow measured in bench-dehydrated and centrifuged samples (static centrifuge), in situ flow measured under tension during spinning in the centrifuge (Cavitron) and direct imaging using X-ray computed microtomography (microCT) observations in stems of two species of Hakea that differ in vessel length. Both centrifuge techniques were prone to artefactual embolism in samples with maximum vessel length longer than, or similar to, the centrifuge rotor diameter. Observations with microCT indicated that this artefactual embolism occurred in the outermost portions of samples. The artefact was largely eliminated if flow was measured in an excised central part of the segment in the static centrifuge or starting measurements with the Cavitron at pressures lower than the threshold of embolism formation in open vessels. The simulations of loss of conductivity in centrifuged samples with a new model, CAVITOPEN, confirmed that the impact of open vessels on the vulnerability to embolism curve was higher when vessels were long, samples short and when embolism is formed in open vessels at less negative pressures. This model also offers a robust and quantitative tool to test and correct for artefactual embolism at low xylem tensions.

Plant species differ in many functional traits that drive differences in rates of photosynthesis, biomass allocation, and tissue turnover. However, it remains unclear how-and even if-such traits influence whole-plant growth, with the simple linear relationships predicted by existing theory often lacking empirical support. Here, we present a theoretical framework for understanding the effect of diverse functional traits on plant growth and shade tolerance by extending a widely used model, linking growth rate in seedlings with a single leaf trait, to explicitly include influences of size, light environment, and five prominent traits: seed mass, height at maturation, leaf mass per unit leaf area, leaf nitrogen per unit leaf area, and wood density. Based on biomass growth and allocation, this framework explains why the influence of traits on growth rate and shade tolerance often varies with plant size and why the impact of size on growth varies among traits. Specifically, we demonstrate why for height growth the influence of: ( i ) leaf mass per unit leaf area is strong in small plants but weakens with size; ( ii ) leaf nitrogen per unit leaf area does not change with size; ( iii ) wood density is present across sizes; ( iv ) height at maturation strengthens with size; and ( v ) seed mass decreases with size. Moreover, we show how traits moderate plant responses to light environment and also determine shade tolerance, supporting diverse empirical results., Competing Interests: The authors declare no conflict of interest.

Severe droughts have caused widespread tree mortality across many forest biomes with profound effects on the function of ecosystems and carbon balance. Climate change is expected to intensify regional-scale droughts, focusing attention on the physiological basis of drought-induced tree mortality. Recent work has shown that catastrophic failure of the plant hydraulic system is a principal mechanism involved in extensive crown death and tree mortality during drought, but the multi-dimensional response of trees to desiccation is complex. Here we focus on the current understanding of tree hydraulic performance under drought, the identification of physiological thresholds that precipitate mortality and the mechanisms of recovery after drought. Building on this, we discuss the potential application of hydraulic thresholds to process-based models that predict mortality.

Rising atmospheric [CO 2 ] and associated climate change are expected to modify primary productivity across a range of ecosystems globally. Increasing aridity is predicted to reduce grassland productivity, although rising [CO 2 ] and associated increases in plant water use efficiency may partially offset the effect of drying on growth. Difficulties arise in predicting the direction and magnitude of future changes in ecosystem productivity, due to limited field experimentation investigating climate and CO 2 interactions. We use repeat near-surface digital photography to quantify the effects of water availability and experimentally manipulated elevated [CO 2 ] (eCO 2 ) on understorey live foliage cover and biomass over three growing seasons in a temperate grassy woodland in south-eastern Australia. We hypothesised that (i) understorey herbaceous productivity is dependent upon soil water availability, and (ii) that eCO 2 will increase productivity, with greatest stimulation occurring under conditions of low water availability. Soil volumetric water content (VWC) determined foliage cover and growth rates over the length of the growing season (August to March), with low VWC (<0.1 m 3  m -3 ) reducing productivity. However, eCO 2 did not increase herbaceous cover and biomass over the duration of the experiment, or mitigate the effects of low water availability on understorey growth rates and cover. Our findings suggest that projected increases in aridity in temperate woodlands are likely to lead to reduced understorey productivity, with little scope for eCO 2 to offset these changes., (© 2018 John Wiley & Sons Ltd.)

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 CO 2 and H 2 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., (© 2018 John Wiley & Sons Ltd.)

Plant hydraulic traits capture the impacts of drought stress on plant function, yet vegetation models lack sufficient information regarding trait coordination and variation with climate-of-origin across species. Here, we investigated key hydraulic and carbon economy traits of 12 woody species in Australia from a broad climatic gradient, with the aim of identifying the coordination among these traits and the role of climate in shaping cross-species trait variation. The influence of environmental variation was minimized by a common garden approach, allowing us to factor out the influence of environment on phenotypic variation across species. We found that hydraulic traits (leaf turgor loss point, stomatal sensitivity to drought [P gs ], xylem vulnerability to cavitation [P x ], and branch capacitance [C branch ]) were highly coordinated across species and strongly related to rainfall and aridity in the species native distributional range. In addition, trade-offs between drought tolerance and plant growth rate were observed across species. Collectively, these results provide critical insight into the coordination among hydraulic traits in modulating drought adaptation and will significantly advance our ability to predict drought vulnerability in these dominant trees species., (© 2018 John Wiley & Sons Ltd.)

Changes in atmospheric [CO2], temperature and precipitation under projected climate change scenarios may have significant impacts on the physiology and yield of cotton. Understanding the implications of integrated environmental impacts on cotton is critical for developing cotton systems that are resilient to stresses induced by climate change. The objective of this study was to quantify the physiological and growth capacity of two cotton cultivars under current and future climate regimes. This experiment compared the early-season growth and physiological response of an older (DP16, released in the 1970s) and a modern (Sicot 71BRF, released in 2008) cotton cultivar grown in ambient and elevated atmospheric [CO2] (CA, 400µLL-1 and CE, 640µLL-1 respectively) and two temperature (TA, 28/17°C and TE, 32/21°C, day/night, respectively) treatments under well-watered conditions. CE increased biomass and photosynthetic rates compared with CA, and TE increased plant biomass. Although limited by the comparison of one older and one modern cultivar, our results suggest that substantial potential may exist to increase breeding selection of cotton cultivars that are responsive to both TE and CE.

The terrestrial carbon and water cycles are intimately linked: the carbon cycle is driven by photosynthesis, while the water balance is dominated by transpiration, and both fluxes are controlled by plant stomatal conductance. The ratio between these fluxes, the plant water-use efficiency (WUE), is a useful indicator of vegetation function. WUE can be estimated using several techniques, including leaf gas exchange, stable isotope discrimination, and eddy covariance. Here we compare global compilations of data for each of these three techniques. We show that patterns of variation in WUE across plant functional types (PFTs) are not consistent among the three datasets. Key discrepancies include the following: leaf-scale data indicate differences between needleleaf and broadleaf forests, but ecosystem-scale data do not; leaf-scale data indicate differences between C 3 and C 4 species, whereas at ecosystem scale there is a difference between C 3 and C 4 crops but not grasslands; and isotope-based estimates of WUE are higher than estimates based on gas exchange for most PFTs. Our study quantifies the uncertainty associated with different methods of measuring WUE, indicates potential for bias when using WUE measures to parameterize or validate models, and indicates key research directions needed to reconcile alternative measures of WUE., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees., (© 2017 John Wiley & Sons Ltd.)

Sink limitation is known to reduce plant growth, but it is not known how plant carbon (C) balance is affected, limiting our ability to predict growth under sink-limited conditions. We manipulated soil volume to impose sink limitation of growth in Eucalyptus tereticornis Sm. seedlings. Seedlings were grown in the field in containers of different sizes and planted flush to the soil alongside freely rooted (Free) seedlings. Container volume negatively affected aboveground growth throughout the experiment, and light saturated rates of leaf photosynthesis were consistently lower in seedlings in containers (-26%) compared with Free seedlings. Significant reductions in photosynthetic capacity in containerized seedlings were related to both reduced leaf nitrogen content and starch accumulation, indicating direct effects of sink limitation on photosynthetic downregulation. After 120 days, harvested biomass of Free seedlings was on average 84% higher than seedlings in containers, but biomass distribution in leaves, stems and roots was not different. However, the reduction in net leaf photosynthesis over the growth period was insufficient to explain the reduction in growth, so that we also observed an apparent reduction in whole-plant C-use efficiency (CUE) between Free seedlings and seedlings in containers. Our results show that sink limitation affects plant growth through feedbacks to both photosynthesis and CUE. Mass balance approaches to predicting plant growth under sink-limited conditions need to incorporate both of these feedbacks., (© The Author 2017. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Global change models indicate that rainfall patterns are likely to shift towards more extreme events concurrent with increasing atmospheric carbon dioxide concentration ([CO 2 ]). Both changes in [CO 2 ] and rainfall regime are known to impact above- and belowground communities, but the interactive effects of these global change drivers have not been well explored, particularly belowground. In this experimental study, we examined the effects of elevated [CO 2 ] (ambient + 240 ppm; [eCO 2 ]) and changes in rainfall patterns (seasonal drought) on soil microbial communities associated with forest ecosystems. Our results show that bacterial and archaeal communities are highly resistant to seasonal drought under ambient [CO 2 ]. However, substantial taxa specific responses to seasonal drought were observed at [eCO 2 ], suggesting that [eCO 2 ] compromise the resistance of microbial communities to extreme events. Within the microbial community we were able to identify three types of taxa specific responses to drought: tolerance, resilience and sensitivity that contributed to this pattern. All taxa were tolerant to seasonal drought at [aCO 2 ], whereas resilience and sensitivity to seasonal drought were much greater in [eCO 2 ]. These results provide strong evidence that [eCO 2 ] moderates soil microbial community responses to drought in forests, with potential implications for their long-term persistence and ecosystem functioning., (© 2017 Society for Applied Microbiology and John Wiley & Sons Ltd.)

Alterations in climate factors such as rising CO2 concentration ([CO2]), warming and reduced precipitation may have significant impacts on plant physiology and growth. This research investigated the interactive effects of elevated [CO2], warming and soil water deficit on biomass production, leaf-level physiological responses and whole-plant water use efficiency (WUEP) in cotton (Gossypium hirsutum L.). Cotton was grown in the glasshouse under two [CO2] treatments (CA, 400µLL-1; CE, 640µLL-1) and two temperature treatments (TA, 28°C:17°C day:night; TE, 32°C:21°C day:night). Plants were subjected to two progressive water deficit cycles, with a 5-day recovery period between the water deficit periods. CE increased vegetative biomass and photosynthetic rates, and decreased stomatal conductance in TA; however, these responses to CE were not evident under TE. CE increased whole-plant water loss under TA, but increased WUEp, whereas increased whole-plant water loss in TE decreased WUEp regardless of atmospheric [CO2]. CE may provide some positive growth and physiological benefits to cotton at TA if sufficient water is available but CE will not mitigate the negative effects of rising temperature on cotton growth and physiology in future environments.

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO 2 diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A n ), stomatal conductance (g s ) and mesophyll conductance (g m ) in Eucalyptus tereticornis trees grown in climate controlled whole-tree chambers. Compared to sun leaves, shade leaves had lower A n , g m , leaf nitrogen and photosynthetic capacity (A max ) but g s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g s and g m increased, and A n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up-scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up-regulation of g m with increased light enables shade leaves to respond quickly to sunflecks., (© 2016 John Wiley & Sons Ltd.)

Frequency and severity of insect outbreaks in forest ecosystems are predicted to increase with climate change. How this will impact canopy leaf area in future climates is rarely tested. Here, we document function of insect outbreaks that fortuitously and rapidly occurred in an ecosystem under free-air CO 2 enrichment. Over the first 2 years of CO 2 fumigation of a naturally established mature Eucalyptus woodland, we continuously assessed population responses of three sap-feeding insect species of the psyllid genera Cardiaspina, Glycaspis and Spondyliaspis for up to ten consecutive generations. Concurrently, we quantified changes in the canopy leaf area index (LAI). Large and rapid shifts in psyllid community composition were recorded between species with either flush (Glycaspis) or senescence-inducing (Cardiaspina, Spondyliaspis) feeding strategies. Within the second year, two psyllid species experienced significant and rapid population build-up resulting in two consecutive outbreaks: first, rainfall stimulated Eucalyptus leaf production increasing LAI, which supported population growth of flush-feeding Glycaspis without impacting LAI. Glycaspis numbers then crashed and were followed by the outbreak of senescence-feeding Cardiaspina fiscella that led to significant defoliation and reduction in LAI. For all three psyllid species, the abundance of lerps, protective coverings excreted by the sessile nymphs, decreased at e[CO 2 ]. Higher lerp weight at e[CO 2 ] for Glycaspis but not the other psyllid species provided evidence for compensatory feeding by the flush feeder but not the two senescence feeders. Our study demonstrates that rainfall drives leaf phenology, facilitating the rapid boom-and-bust succession of psyllid species, eventually leading to significant defoliation due to the second but not the first outbreaking psyllid species. In contrast, e[CO 2 ] may impact psyllid abundance and feeding behaviour, with psyllid species-specific outcomes for defoliation severity, nutrient transfer and trophic cascades. Psyllid populations feeding on Eucalyptus experience rapid boom-and-bust cycles depending on availability of suitable foliage driven by rainfall patterns and leaf phenology., (© 2016 John Wiley & Sons Ltd.)

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., (© 2016 John Wiley & Sons Ltd.)

Here, we aim to understand differences in biomass distribution between major woody plant functional types (PFTs) (deciduous vs evergreen and gymnosperm vs angiosperm) in terms of underlying traits, in particular the leaf mass per area (LMA) and leaf area per unit stem basal area. We used a large compilation of plant biomass and size observations, including observations of 21 084 individuals on 656 species. We used a combination of semiparametric methods and variance partitioning to test the influence of PFT, plant height, LMA, total leaf area, stem basal area and climate on above-ground biomass distribution. The ratio of leaf mass to above-ground woody mass (MF /MS ) varied strongly among PFTs. We found that MF /MS at a given plant height was proportional to LMA across PFTs. As a result, the PFTs did not differ in the amount of leaf area supported per unit above-ground biomass or per unit stem basal area. Climate consistently explained very little additional variation in biomass distribution at a given plant size. Combined, these results demonstrate consistent patterns in above-ground biomass distribution and leaf area relationships among major woody PFTs, which can be used to further constrain global vegetation models., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

Given the contrasting short-term temperature dependences of gross primary production (GPP) and autotrophic respiration, the fraction of GPP respired by trees is predicted to increase with warming, providing a positive feedback to climate change. However, physiological acclimation may dampen or eliminate this response. We measured the fluxes of aboveground respiration (Ra ), GPP and their ratio (Ra /GPP) in large, field-grown Eucalyptus tereticornis trees exposed to ambient or warmed air temperatures (+3°C). We report continuous measurements of whole-canopy CO2 exchange, direct temperature response curves of leaf and canopy respiration, leaf and branch wood respiration, and diurnal photosynthetic measurements. Warming reduced photosynthesis, whereas physiological acclimation prevented a coincident increase in Ra . Ambient and warmed trees had a common nonlinear relationship between the fraction of GPP that was respired above ground (Ra /GPP) and the mean daily temperature. Thus, warming significantly increased Ra /GPP by moving plants to higher positions on the shared Ra /GPP vs daily temperature relationship, but this effect was modest and only notable during hot conditions. Despite the physiological acclimation of autotrophic respiration to warming, increases in temperature and the frequency of heat waves may modestly increase tree Ra /GPP, contributing to a positive feedback between climate warming and atmospheric CO2 accumulation., (© 2016 The Authors. New Phytologist © 2016 New Phytologist Trust.)

The response of terrestrial ecosystems to rising atmospheric CO2 concentration (Ca ), particularly under nutrient-limited conditions, is a major uncertainty in Earth System models. The Eucalyptus Free-Air CO2 Enrichment (EucFACE) experiment, recently established in a nutrient- and water-limited woodland presents a unique opportunity to address this uncertainty, but can best do so if key model uncertainties have been identified in advance. We applied seven vegetation models, which have previously been comprehensively assessed against earlier forest FACE experiments, to simulate a priori possible outcomes from EucFACE. Our goals were to provide quantitative projections against which to evaluate data as they are collected, and to identify key measurements that should be made in the experiment to allow discrimination among alternative model assumptions in a postexperiment model intercomparison. Simulated responses of annual net primary productivity (NPP) to elevated Ca ranged from 0.5 to 25% across models. The simulated reduction of NPP during a low-rainfall year also varied widely, from 24 to 70%. Key processes where assumptions caused disagreement among models included nutrient limitations to growth; feedbacks to nutrient uptake; autotrophic respiration; and the impact of low soil moisture availability on plant processes. Knowledge of the causes of variation among models is now guiding data collection in the experiment, with the expectation that the experimental data can optimally inform future model improvements., (© 2016 John Wiley & Sons Ltd.)

The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling., (© 2015 John Wiley & Sons Ltd.)

Vegetation CO2 uptake is always accompanied by water loss. The balance in this gas exchange is controlled by the stomata, through which CO2 and water vapour diffuse between the leaf and the atmosphere. The optimal stomatal behaviour theory proposes that vegetation should optimise its stomatal behaviour such that, for given water availability, photosynthesis is maximised. In this paper, we optimise stomatal conductance as a function of soil water content for the maximum expected value of photosynthesis rate. This optimisation process is considered under stochastic rainfall. The optimal solution is largely shaped by two constraints: the risks of soil water exhaustion and surface runoff, which results in an inverse S-shaped curve of stomatal conductance along the soil water gradient. We derive the optimal functional relationship between stomatal conductance and soil water content under varying rainfall frequency, mean annual precipitation and atmospheric CO2 concentration. Comparisons with large-scale observational data show that the model is able to broadly capture responses of photosynthesis, transpiration, and water use efficiency along rainfall gradients, although notable discrepancies suggest additional factors are important in shaping these responses. Our work provides a theoretical framework for analysing the vegetation gas exchange under environmental change., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Canopy leaf area, quantified by the leaf area index (L), is a crucial driver of forest productivity, water use and energy balance. Because L responds to environmental drivers, it can represent an important feedback to climate change, but its responses to rising atmospheric [CO2] and water availability of forests have been poorly quantified. We studied canopy leaf area dynamics for 28 months in a native evergreen Eucalyptus woodland exposed to free-air CO2 enrichment (the EucFACE experiment), in a subtropical climate where water limitation is common. We hypothesized that, because of expected stimulation of productivity and water-use efficiency, L should increase with elevated [CO2]. We estimated L from diffuse canopy transmittance, and measured monthly leaf litter production. Contrary to expectation, L did not respond to elevated [CO2]. We found that L varied between 1.10 and 2.20 across the study period. The dynamics of L showed a quick increase after heavy rainfall and a steady decrease during periods of low rainfall. Leaf litter production was correlated to changes in L, both during periods of decreasing L (when no leaf growth occurred) and during periods of increasing L (active shedding of old foliage when new leaf growth occurred). Leaf lifespan, estimated from mean L and total annual litter production, was up to 2 months longer under elevated [CO2] (1.18 vs. 1.01 years; P = 0.05). Our main finding that L was not responsive to elevated CO2 is consistent with other forest FACE studies, but contrasts with the positive response of L commonly predicted by many ecosystem models., (© 2015 John Wiley & Sons Ltd.)

Models of tree responses to climate typically project that elevated atmospheric CO2 concentration (eCa ) will reduce drought impacts on forests. We tested one of the mechanisms underlying this interaction, the 'low Ci effect', in which stomatal closure in drought conditions reduces the intercellular CO2 concentration (Ci ), resulting in a larger relative enhancement of photosynthesis with eCa , and, consequently, a larger relative biomass response. We grew two Eucalyptus species of contrasting drought tolerance at ambient and elevated Ca for 6-9 months in large pots maintained at 50% (drought) and 100% field capacity. Droughted plants did not have significantly lower Ci than well-watered plants, which we attributed to long-term changes in leaf area. Hence, there should not have been an interaction between eCa and water availability on biomass, and we did not detect one. The xeric species did have higher Ci than the mesic species, indicating lower water-use efficiency, but both species exhibited similar responses of photosynthesis and biomass to eCa , owing to compensatory differences in the photosynthetic response to Ci . Our results demonstrate that long-term acclimation to drought, and coordination among species traits may be important for predicting plant responses to eCa under low water availability., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

The first generation of forest free-air CO2 enrichment (FACE) experiments has successfully provided deeper understanding about how forests respond to an increasing CO2 concentration in the atmosphere. Located in aggrading stands in the temperate zone, they have provided a strong foundation for testing critical assumptions in terrestrial biosphere models that are being used to project future interactions between forest productivity and the atmosphere, despite the limited inference space of these experiments with regards to the range of global ecosystems. Now, a new generation of FACE experiments in mature forests in different biomes and over a wide range of climate space and biodiversity will significantly expand the inference space. These new experiments are: EucFACE in a mature Eucalyptus stand on highly weathered soil in subtropical Australia; AmazonFACE in a highly diverse, primary rainforest in Brazil; BIFoR-FACE in a 150-yr-old deciduous woodland stand in central England; and SwedFACE proposed in a hemiboreal, Pinus sylvestris stand in Sweden. We now have a unique opportunity to initiate a model-data interaction as an integral part of experimental design and to address a set of cross-site science questions on topics including responses of mature forests; interactions with temperature, water stress, and phosphorus limitation; and the influence of biodiversity., (© UT-Battelle, LLC New Phytologist © 2015 New Phytologist Trust.)

Here I present the R package 'plantecophys', a toolkit to analyse and model leaf gas exchange data. Measurements of leaf photosynthesis and transpiration are routinely collected with portable gas exchange instruments, and analysed with a few key models. These models include the Farquhar-von Caemmerer-Berry (FvCB) model of leaf photosynthesis, the Ball-Berry models of stomatal conductance, and the coupled leaf gas exchange model which combines the supply and demand functions for CO2 in the leaf. The 'plantecophys' R package includes functions for fitting these models to measurements, as well as simulating from the fitted models to aid in interpreting experimental data. Here I describe the functionality and implementation of the new package, and give some examples of its use. I briefly describe functions for fitting the FvCB model of photosynthesis to measurements of photosynthesis-CO2 response curves ('A-Ci curves'), fitting Ball-Berry type models, modelling C3 photosynthesis with the coupled photosynthesis-stomatal conductance model, modelling C4 photosynthesis, numerical solution of optimal stomatal behaviour, and energy balance calculations using the Penman-Monteith equation. This open-source package makes technically challenging calculations easily accessible for many users and is freely available on CRAN.

Future climate regimes characterized by rising [CO2], rising temperatures and associated droughts may differentially affect tree growth and physiology. However, the interactive effects of these three factors are complex because elevated [CO2] and elevated temperature may generate differential physiological responses during drought. To date, the interactive effects of elevated [CO2] and elevated temperature on drought-induced tree mortality remain poorly understood in gymnosperm species that differ in stomatal regulation strategies. Water relations and carbon dynamics were examined in two species with contrasting stomatal regulation strategies: Pinus radiata D. Don (relatively isohydric gymnosperm; regulating stomata to maintain leaf water potential above critical thresholds) and Callitris rhomboidea R. Br (relatively anisohydric gymnosperm; allowing leaf water potential to decline as the soil dries), to assess response to drought as a function of [CO2] and temperature. Both species were grown in two [CO2] (C(a) (ambient, 400 μl l(-1)) and C(e) (elevated, 640 μl l(-1))) and two temperature (T(a) (ambient) and T(e) (ambient +4 °C)) treatments in a sun-lit glasshouse under well-watered conditions. Drought plants were then exposed to a progressive drought until mortality. Prior to mortality, extensive xylem cavitation occurred in both species, but significant depletion of non-structural carbohydrates was not observed in either species. Te resulted in faster mortality in P. radiata, but it did not modify the time-to-mortality in C. rhomboidea. C(e) did not delay the time-to-mortality in either species under drought or T(e) treatments. In summary, elevated temperature (+4 °C) had greater influence than elevated [CO2] (+240 μl l(-1)) on drought responses of the two studied gymnosperm species, while stomatal regulation strategies did not generally affect the relative contributions of hydraulic failure and carbohydrate depletion to mortality under severe drought., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

It has been reported that elevated temperature accelerates the time-to-mortality in plants exposed to prolonged drought, while elevated [CO(2)] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO(2)] and temperature on the inter-dependent carbon and hydraulic characteristics associated with drought-induced mortality in Eucalyptus radiata seedlings grown in two [CO(2)] (400 and 640 μL L(-1)) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO(2)] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO(2)], may be the primary contributors to drought-induced seedling mortality under future climates., (© 2013 John Wiley & Sons Ltd.)

Elevated [CO2] and temperature may alter the drought responses of tree seedling growth, photosynthesis, respiration and total non-structural carbohydrate (TNC) status depending on drought intensity and duration. Few studies have addressed these important climatic interactions or their consequences. We grew Eucalyptus globulus Labill. seedlings in two [CO2] concentrations (400 and 640 μl l(-1)) and two temperatures (28/17 and 32/21 °C) (day/night) in a sun-lit glasshouse, and grew them in well-watered conditions or exposed them to two drought treatments having undergone different previous water conditions (i.e., rewatered drought and sustained drought). Progressive drought in both drought treatments led to similar limitations in growth, photosynthesis and respiration, but reductions in TNC concentration were not observed. Elevated [CO2] ameliorated the impact of the drought during the moderate drought phase (i.e., Day 63 to Day 79) by increasing photosynthesis and enhancing leaf and whole-plant TNC content. In contrast, elevated temperature exacerbated the impact of the drought during the moderate drought phase by reducing photosynthesis, increasing leaf respiration and decreasing whole-plant TNC content. Extreme drought (i.e., Day 79 to Day 103) eliminated [CO2] and temperature effects on plant growth, photosynthesis and respiration. The combined effects of elevated [CO2] and elevated temperature on moderate drought stressed seedlings were reduced with progressive drought, with no sustained effects on growth despite greater whole-plant TNC content.