Comparing global models of terrestrial net primary productivity (NPP): the importance of water availability

Given that neither absolute measures nor direct model validations of global terrestrial net primary productivity (NPP) are feasible, intercomparison of global NPP models provides an effective tool to check model consistency. For this study, we tested the assumption that water availability is the primary limiting factor of NPP in global terrestrial biospheric models. We compared a water balance coefficient (WBC), calculated as the difference of mean annual precipitation and potential evapotranspiration to NPP for each grid cell (0.5° × 0.5° longitude/latitude) in each of 14 models. We also evaluated different approaches used for introducing water budget limitations on NPP: (1) direct physiological control on evapotranspiration through canopy conductance; (2) climatological computation of constraints from supply/demand for ecosystem productivity; and (3) water limitation inferred from satellite data alone. Plots of NPP vs. WBC showed comparable patterns for the models using the same method for water balance limitation on NPP. While correlation plots revealed similar patterns for most global models, other environmental controls on NPP introduced substantial variability.     

Theoretical basis for stabilizing messenger RNA through secondary structure design

RNA hydrolysis presents problems in manufacturing, long-term storage, world-wide delivery and in vivo stability of messenger RNA (mRNA)-based vaccines and therapeutics. A largely unexplored strategy to reduce mRNA hydrolysis is to redesign RNAs to form double-stranded regions, which are protected from in-line cleavage and enzymatic degradation, while coding for the same proteins. The amount of stabilization that this strategy can deliver and the most effective algorithmic approach to achieve stabilization remain poorly understood. Here, we present simple calculations for estimating RNA stability against hydrolysis, and a model that links the average unpaired probability of an mRNA, or AUP, to its overall hydrolysis rate. To characterize the stabilization achievable through structure design, we compare AUP optimization by conventional mRNA design methods to results from more computationally sophisticated algorithms and crowdsourcing through the OpenVaccine challenge on the Eterna platform. We find that rational design on Eterna and the more sophisticated algorithms lead to constructs with low AUP, which we term ‘superfolder’ mRNAs. These designs exhibit a wide diversity of sequence and structure features that may be desirable for translation, biophysical size, and immunogenicity. Furthermore, their folding is robust to temperature, computer modeling method, choice of flanking untranslated regions, and changes in target protein sequence, as illustrated by rapid redesign of superfolder mRNAs for B.1.351, P.1 and B.1.1.7 variants of the prefusion-stabilized SARS-CoV-2 spike protein. Increases in in vitro mRNA half-life by at least two-fold appear immediately achievable.     

Comparing global models of terrestrial net primary productivity (NPP): analysis of differences in light absorption and light-use efficiency

Twelve global net primary productivity (NPP) models were compared: BIOME3, CASA, CARAIB, FBM, GLO-PEM, HYBRID, KGBM, PLAI, SDBM, SIB2, SILVAN and TURC. These models all use solar radiation as an input, and compute either absorbed solar radiation directly, or the amount of leaves used to absorb solar radiation, represented by the leaf area index (LAI). For all models, we obtained or estimated photosynthetically active radiation absorbed by the canopy (APAR). We then computed the light use efficiency for NPP (LUE) on an annual basis as the ratio of NPP to APAR. We analysed the relative importance for NPP of APAR and LUE. The analyses consider the global values of these factors, their spatial patterns represented by latitudinal variations, and the overall grid cell by grid cell variability. Spatial variability in NPP within a model proved to be determined by APAR, and differences among models by LUE. There was a compensation between APAR and LUE, so that global NPP values fell within the range of ‘generally accepted values’. Overall, APAR was lower for satellite driven models than for the other models. Most computed values of LUE were within the range of published values, except for one model.     

Quantifying the response of photosynthesis to changes in leaf nitrogen content and leaf mass per area in plants grown under atmospheric CO2 enrichment

Previous modelling exercises and conceptual arguments have predicted that a reduction in biochemical capacity for photosynthesis (A_area) at elevated CO₂ may be compensated by an increase in mesophyll tissue growth if the total amount of photosynthetic machinery per unit leaf area is maintained (i.e. morphological upregulation). The model prediction was based on modelling photosynthesis as a function of leaf N per unit leaf area (N_area), where N_area = N_mass×LMA. Here, N_mass is percentage leaf N and is used to estimate biochemical capacity and LMA is leaf mass per unit leaf area and is an index of leaf morphology. To assess the relative importance of changes in biochemical capacity versus leaf morphology we need to control for multiple correlations that are known, or that are likely to exist between CO₂ concentration, N_area, N_mass, LMA and A_area. Although this is impractical experimentally, we can control for these correlations statistically using systems of linear multiple-regression equations. We developed a linear model to partition the response of A_area to elevated CO₂ into components representing the independent and interactive effects of changes in indexes of biochemical capacity, leaf morphology and CO₂ limitation of photosynthesis. The model was fitted to data from three pine and seven deciduous tree species grown in separate chamber-based field experiments. Photosynthetic enhancement at elevated CO₂ due to morphological upregulation was negligible for most species. The response of A_area in these species was dominated by the reduction in CO₂ limitation occurring at higher CO₂ concentration. However, some species displayed a significant reduction in potential photosynthesis at elevated CO₂ due to an increase in LMA that was independent of any changes in N_area. This morphologically based inhibition of A_area combined additively with a reduction in biochemical capacity to significantly offset the direct enhancement of A_area caused by reduced CO₂ limitation in two species. This offset was 100% for Acer rubrum, resulting in no net effect of elevated CO₂ on A_area for this species, and 44% for Betula pendula. This analysis shows that interactions between biochemical and morphological responses to elevated CO₂ can have important effects on photosynthesis.     

Comparing global models of terrestrial net primary productivity (NPP): analysis of the seasonal atmospheric CO2 signal

Eight terrestrial biospheric models (TBMs) calculating the monthly distributions of both net primary productivity (NPP) and soil heterotrophic respiration (R_H) in the Potsdam NPP Model Intercomparison workshop are used to simulate seasonal patterns of atmospheric CO₂ concentration. For each model, we used net ecosystem productivity (NEP = NPP – R_H) as the source function in the TM2 atmospheric transport model from the Max-Planck Institute for Meteorology. Comparing the simulated concentration fields with detrended measurements from 25 monitoring stations spread over the world, we found that the decreasing seasonal amplitude from north to south is rather well reproduced by all the models, though the amplitudes are slightly too low in the north. The agreement between the simulated and observed seasonality is good in the northern hemisphere, but poor in the southern hemisphere, even when the ocean is accounted for. Based on a Fourier analysis of the calculated zonal atmospheric signals, tropical NEP plays a key role in the seasonal cycle of the atmospheric CO₂ in the whole southern hemisphere. The relatively poor match between measured and predicted atmospheric CO₂ in this hemisphere suggests problems with all the models. The simulation of water relations, a dominant regulator of NEP in the tropics, is a leading candidate for the source of these problems.     

Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.