Nitrogen to phosphorus ratio of plant biomass versus soil solution in a tropical pioneer tree, Ficus insipida

It is commonly assumed that the nitrogen to phosphorus (N:P) ratio of a terrestrial plant reflects the relative availability of N and P in the soil in which the plant grows. Here, this was assessed for a tropical pioneer tree, Ficus insipida. Seedlings were grown in sand and irrigated with nutrient solutions containing N:P ratios ranging from <1 to >100. The experimental design further allowed investigation of physiological responses to N and P availability. Homeostatic control over N:P ratios was stronger in leaves than in stems or roots, suggesting that N:P ratios of stems and roots are more sensitive indicators of the relative availability of N and P at a site than N:P ratios of leaves. The leaf N:P ratio at which the largest plant dry mass and highest photosynthetic rates were achieved was ∼11, whereas the corresponding whole-plant N:P ratio was ∼6. Plant P concentration varied as a function of transpiration rate at constant nutrient solution P concentration, possibly due to transpiration-induced variation in the mass flow of P to root surfaces. The transpiration rate varied in response to nutrient solution N concentration, but not to nutrient solution P concentration, demonstrating nutritional control over transpiration by N but not P. Water-use efficiency varied as a function of N availability, but not as a function of P availability

Transpiration efficiency of a tropical pioneer tree (Ficus insipida) in relation to soil fertility

The response of whole-plant water-use efficiency, termed transpiration efficiency (TE), to variation in soil fertility was assessed in a tropical pioneer tree, Ficus insipida Willd. Measurements of stable isotope ratios (d13C, d18O, d15N), elemental concentrations (C, N, P),plant growth, instantaneous leaf gas exchange, and whole-plant water use were used to analyse the mechanisms controlling TE. Plants were grown individually in 19 l pots with non-limiting soil moisture. Soil fertility was altered by mixing soil with varying proportions of rice husks, and applying a slow release fertilizer. A large variation was observed in leaf photosynthetic rate, mean relative growth rate (RGR), and TE in response to experimental treatments; these traits were well correlated with variation in leaf N concentration. Variation in TE showed a strong dependence on the ratio of intercellular to ambient CO2 mole fractions (ci/ca); both for instantaneous measurements of ci/ca (R2=0.69, P <0.0001, n=30), and integrated estimates based on C isotope discrimination (R2 =0.88, P<0.0001, n=30). On the other hand, variations in the leaf-to-air humidity gradient, unproductive water loss, and respiratory C use probably played only minor roles in modulating TE in the face of variable soil fertility. The pronounced variation in TE resulted from a combination of the strong response of ci/ca to leaf N, and inherently high values of ci/ca for this tropical tree species; these two factors conspired to cause a 4-fold variation among treatments in (1–ci/ca ), the term that actually modifies TE. Results suggest that variation in plant N status could have important implications for the coupling between C and water exchange in tropical forest trees

Why are tropical conifers disadvantaged in fertile soils? Comparison of Podocarpus guatemalensis with an angiosperm pioneer, Ficus insipida

Conifers are, for the most part, competitively excluded from tropical rainforests by angiosperms. Where they do occur, conifers often occupy sites that are relatively infertile. To gain insight into the physiological mechanisms by which angiosperms outcompete conifers in more productive sites, we grew seedlings of a tropical conifer (Podocarpus guatemalensis Standley) and an angiosperm pioneer (Ficus insipida Willd.) with and without added nutrients, supplied in the form of a slow-release fertilizer. At the conclusion of the experiment, the dry mass of P. guatemalensis seedlings in fertilized soil was approximately twofold larger than that of seedlings in unfertilized soil; on the other hand, the dry mass of F. insipida seedlings in fertilized soil was similar to 20-fold larger than seedlings in unfertilized soil. The higher relative growth rate of F. insipida was associated with a larger leaf area ratio and a higher photosynthetic rate per unit leaf area. Higher overall photosynthetic rates in F. insipida were associated with an approximately fivefold larger stomatal conductance than in P. guatemalensis. We surmise that a higher whole-plant hydraulic conductance in the vessel bearing angiosperm F. insipida enabled higher leaf area ratio and higher stomatal conductance per unit leaf area than in the tracheid bearing P. guatemalensis, which enabled F. insipida to capitalize on increased photosynthetic capacity driven by higher nitrogen availability in fertilized soil

Convergence in phosphorus constraints to photosynthesis in forests around the world

This is the final version. Available on open access from Nature Research via the DOI in this recordData availability: The photosynthesis and leaf nutrient data reported in the paper are available at https://doi.org/10.6084/m9.figshare.20010485.v1, and the model results are available on the European open-access repository Zenodo at https://doi.org/10.5281/zenodo.6619615. All other data reported in the paper are presented in the supplementary materials.Code availability: The R code used for analyses is at https://github.com/ellswor2/photo_p_repo2.git. The source code for ORCHIDEE is at https://doi.org/10.14768/20200407002.1.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

Changes in leaf functional traits with leaf age: when do leaves decrease their photosynthetic capacity in Amazonian trees?

Most leaf functional trait studies in the Amazon basin do not consider ontogenetic variations (leaf age), which may influence ecosystem productivity throughout the year. When leaf age is taken into account, it is generally considered discontinuous, and leaves are classified into age categories based on qualitative observations. Here, we quantified age-dependent changes in leaf functional traits such as the maximum carboxylation rate of ribulose-1,5-biphosphate carboxylase/oxygenase (Rubisco) (Vcmax), stomatal control (Cgs%), leaf dry mass per area and leaf macronutrient concentrations for nine naturally growing Amazon tropical trees with variable phenological strategies. Leaf ages were assessed by monthly censuses of branch-level leaf demography; we also performed leaf trait measurements accounting for leaf chronological age based on days elapsed since the first inclusion in the leaf demography, not predetermined age classes. At the tree community scale, a nonlinear relationship between Vcmax and leaf age existed: young, developing leaves showed the lowest mean photosynthetic capacity, increasing to a maximum at 45 days and then decreasing gradually with age in both continuous and categorical age group analyses. Maturation times among species and phenological habits differed substantially, from 8 ± 30 to 238 ± 30 days, and the rate of decline of Vcmax varied from −0.003 to −0.065 μmol CO2 m−2 s−1 day−1. Stomatal control increased significantly in young leaves but remained constant after peaking. Mass-based phosphorus and potassium concentrations displayed negative relationships with leaf age, whereas nitrogen did not vary temporally. Differences in life strategies, leaf nutrient concentrations and phenological types, not the leaf age effect alone, may thus be important factors for understanding observed photosynthesis seasonality in Amazonian forests. Furthermore, assigning leaf age categories in diverse tree communities may not be the recommended method for studying carbon uptake seasonality in the Amazon, since the relationship between Vcmax and leaf age could not be confirmed for all trees

Do 2H and 18O in leaf water reflect environmental drivers differently?

We compiled hydrogen and oxygen stable isotope compositions (δ H and δ O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ H was more closely correlated with δ H of xylem water or atmospheric vapour, whereas leaf water δ O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ O. We next expressed leaf water as isotopic enrichment above xylem water (Δ H and Δ O) to remove the impact of xylem water isotopic variation. For Δ H, leaf water still correlated with atmospheric vapour, whereas Δ O showed no such correlation. This was explained by covariance between air relative humidity and the Δ O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that H and O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water

Optimal stomatal behaviour around the world

© 2015 Macmillan Publishers Limited. All rights reserved. Stomatal conductance (g s) is a key land-surface attribute as it links transpiration, the dominant component of global land evapotranspiration, and photosynthesis, the driving force of the global carbon cycle. Despite the pivotal role of g s in predictions of global water and carbon cycle changes, a global-scale database and an associated globally applicable model of g s that allow predictions of stomatal behaviour are lacking. Here, we present a database of globally distributed g s obtained in the field for a wide range of plant functional types (PFTs) and biomes. We find that stomatal behaviour differs among PFTs according to their marginal carbon cost of water use, as predicted by the theory underpinning the optimal stomatal model and the leaf and wood economics spectrum. We also demonstrate a global relationship with climate. These findings provide a robust theoretical framework for understanding and predicting the behaviour of g s across biomes and across PFTs that can be applied to regional, continental and global-scale modelling of ecosystem productivity, energy balance and ecohydrological processes in a future changing climate

Optimal stomatal behaviour around the world

This is the author accepted manuscript. The final version is available from Springer Nature via the DOI in this recordStomatal conductance (g s) is a key land-surface attribute as it links transpiration, the dominant component of global land evapotranspiration, and photosynthesis, the driving force of the global carbon cycle. Despite the pivotal role of g s in predictions of global water and carbon cycle changes, a global-scale database and an associated globally applicable model of g s that allow predictions of stomatal behaviour are lacking. Here, we present a database of globally distributed g s obtained in the field for a wide range of plant functional types (PFTs) and biomes. We find that stomatal behaviour differs among PFTs according to their marginal carbon cost of water use, as predicted by the theory underpinning the optimal stomatal model and the leaf and wood economics spectrum. We also demonstrate a global relationship with climate. These findings provide a robust theoretical framework for understanding and predicting the behaviour of g s across biomes and across PFTs that can be applied to regional, continental and global-scale modelling of ecosystem productivity, energy balance and ecohydrological processes in a future changing climate.This research was supported by the Australian Research Council (ARC MIA Discovery Project 1433500-2012-14). A.R. was financially supported in part by The Next-Generation Ecosystem Experiments (NGEE-Arctic) project, which is supported by the Office of Biological and Environmental Research in the Department of Energy, Office of Science, and through the United States Department of Energy contract No. DE-AC02-98CH10886 to Brookhaven National Laboratory. M.O.d.B. acknowledges that the Brassica data were obtained within a research project financed by the Belgian Science Policy (OFFQ, contract number SD/AF/02) and coordinated by K. Vandermeiren at the Open-Top Chamber research facilities of CODA-CERVA (Tervuren, Belgium)

Pantropical modelling of canopy functional traits using Sentinel-2 remote sensing data

Funding Information: This work is a product of the Global Ecosystems Monitoring (GEM) network (gem.tropicalforests.ox.ac.uk). J.A.G. was funded by the Natural Environment Research Council (NERC; NE/T011084/1 and NE/S011811/1) and the Netherlands Organisation for Scientific Research (NWO) under the Rubicon programme with project number 019.162LW.010. The traits field campaign was funded by a grant to Y.M. from the European Research Council (Advanced Grant GEM-TRAIT: 321131) under the European Union‘s Seventh Framework Programme (FP7/2007-2013), with additional support from NERC Grant NE/D014174/1 and NE/J022616/1 for traits work in Peru, NERC Grant ECOFOR (NE/K016385/1) for traits work in Santarem, NERC Grant BALI (NE/K016369/1) for plot and traits work in Malaysia and ERC Advanced Grant T-FORCES (291585) to Phillips for traits work in Australia. Plot setup in Ghana and Gabon were funded by a NERC Grant NE/I014705/1 and by the Royal Society-Leverhulme Africa Capacity Building Programme. The Malaysia campaign was also funded by NERC GrantNE/K016253/1. Plot inventories in Peru were supported by funding from the US National Science Foundation Long-Term Research in Environmental Biology program (LTREB; DEB 1754647) and the Gordon and Betty Moore Foundation Andes-Amazon Program. Plots inventories in Nova Xavantina (Brazil) were supported by the National Council for Scientific and Technological Development (CNPq), Long Term Ecological Research Program (PELD), Proc. 441244/2016-5, and the Foundation of Research Support of Mato Grosso (FAPEMAT), Project ReFlor, Proc. 589267/2016. During data collection, I.O. was supported by a Marie Curie Fellowship (FP7-PEOPLE-2012-IEF-327990). GEM trait data in Gabon was collected under authorisation to Y.M. and supported by the Gabon National Parks Agency. D.B. was funded by the Fondation Wiener-Anspach. W.D.K. acknowledges support from the Faculty Research Cluster ‘Global Ecology’ of the University of Amsterdam. M.S. was funded by a grant from the Ministry of Education, Youth and Sports of the Czech Republic (INTER-TRANSFER LTT19018). Y.M. is supported by the Jackson Foundation. We thank the two anonymous reviewers and Associate Editor G. Henebry for their insightful comments that helped improved this manuscript.Peer reviewedPostprin

Effects of two contrasting canopy manipulations on growth and water use of London plane (Platanus x acerifolia) trees

Aims: Two contrasting canopy manipulations were compared to unpruned controls on London plane trees, to determine the effects on canopy regrowth, soil and leaf water relations. Methods: ‘Canopy reduction’, was achieved by removing the outer 30 % length of all major branches and ‘canopy thinning’, by removing 30 % of lateral branches arising from major branches. Results: Total canopy leaf areas recovered within two and three years of pruning for the canopy-thinned and reduced trees respectively. Canopy reduction increased mean leaf size, nitrogen concentration, canopy leaf area density and conserved soil moisture for up to 3 years, whereas canopy thinning had no effects. Another experiment compared more severe canopy reduction to unpruned trees. This produced a similar growth response to the previous experiment, but soil moisture was conserved nearer to the trunk. Analysis of 13C and 18O signals along with leaf water relations and soil moisture data suggested that lower boundary layer conductance within the canopy-reduced trees restricted tree water use, whereas for the canopy-thinned trees the opposite occurred. Conclusions: Only canopy reduction conserved soil moisture and this was due to a combination of reduced total canopy leaf area and structural changes in canopy architecture