Convergence in phosphorus constraints to photosynthesis in forests around the world.

Tropical forests take up more carbon (C) from the atmosphere per annum by photosynthesis than any other type of vegetation. Phosphorus (P) limitations to C uptake are paramount for tropical and subtropical forests around the globe. Yet the generality of photosynthesis-P relationships underlying these limitations are in question, and hence are not represented well in terrestrial biosphere models. Here we demonstrate the dependence of photosynthesis and underlying processes on both leaf N and P concentrations. The regulation of photosynthetic capacity by P was similar across four continents. Implementing P constraints in the ORCHIDEE-CNP model, gross photosynthesis was reduced by 36% across the tropics and subtropics relative to traditional N constraints and unlimiting leaf P. Our results provide a quantitative relationship for the P dependence for photosynthesis for the front-end of global terrestrial C models that is consistent with canopy leaf measurements.

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.

Elevated CO2 does not affect stem CO2 efflux nor stem respiration in a dry Eucalyptus woodland, but it shifts the vertical gradient in xylem [CO2 ].

To quantify stem respiration (RS ) under elevated CO2 (eCO2 ), stem CO2 efflux (EA ) and CO2 flux through the xylem (FT ) should be accounted for, because part of respired CO2 is transported upwards with the sap solution. However, previous studies have used EA as a proxy of RS , which could lead to equivocal conclusions. Here, to test the effect of eCO2 on RS , both EA and FT were measured in a free-air CO2 enrichment experiment located in a mature Eucalyptus native forest. Drought stress substantially reduced EA and RS , which were unaffected by eCO2 , likely as a consequence of its neutral effect on stem growth in this phosphorus-limited site. However, xylem CO2 concentration measured near the stem base was higher under eCO2 , and decreased along the stem resulting in a negative contribution of FT to RS , whereas the contribution of FT to RS under ambient CO2 was positive. Negative FT indicates net efflux of CO2 respired below the monitored stem segment, likely coming from the roots. Our results highlight the role of nutrient availability on the dependency of RS on eCO2 and suggest stimulated root respiration under eCO2 that may shift vertical gradients in xylem [CO2 ] confounding the interpretation of EA measurements.

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.

Model-data synthesis for the next generation of forest free-air CO2 enrichment (FACE) experiments.

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.

Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species.

Leaf photosynthetic CO2 responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO2 assimilation rates (Anet). However, triose-phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower Anet in high [CO2] and low [O2 ] than at ambient [O2 ]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of Anet at saturating [CO2] (i.e. Amax) to 2 kPa O2. Three species showed >20% reductions in Amax in low [O2], a phenomenon potentially explained by orthophosphate (Pi) savings during photorespiration. These species, with largest photosynthetic capacity and Pi > 2 mmol P m(-2), rely the most on additional Pi made available from photorespiration rather than species growing in P-impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting Amax and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered.

Lifetime return on investment increases with leaf lifespan among 10 Australian woodland species.

• Co-occurring species often differ in their leaf lifespan (LL) and it remains unclear how such variation is maintained in a competitive context. Here we test the hypothesis that leaves of long-LL species yield a greater return in carbon (C) fixed per unit C or nutrient invested by the plant than those of short-LL species. • For 10 sympatric woodland species, we assessed three-dimensional shoot architecture, canopy openness, leaf photosynthetic light response, leaf dark respiration and leaf construction costs across leaf age sequences. We then used the YPLANT model to estimate light interception and C revenue along the measured leaf age sequences. This was done under a series of simulations that incorporated the potential covariates of LL in an additive fashion. • Lifetime return in C fixed per unit C, N or P invested increased with LL in all simulations. • In contrast to other recent studies, our results show that extended LL confers a fundamental economic advantage by increasing a plant's return on investment in leaves. This suggests that time-discounting effects, that is, the compounding of income that arises from quick reinvestment of C revenue, are key in allowing short-LL species to succeed in the face of this economic handicap.