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In order to quantify the effects of forests to oil palm conversion occurring in the tropics on land–atmosphere carbon, water and energy fluxes, we develop a new perennial crop sub-model CLM-Palm for simulating a palm plant functional type (PFT) within the framework of the Community Land Model (CLM4.5). CLM-Palm is tested here on oil palm only but is meant of generic interest for other palm crops (e.g., coconut). The oil palm has monopodial morphology and sequential phenology of around 40 stacked phytomers, each carrying a large leaf and a fruit bunch, forming a multilayer canopy. A sub-canopy phenological and physiological parameterization is thus introduced so that each phytomer has its own prognostic leaf growth and fruit yield capacity but with shared stem and root components. Phenology and carbon and nitrogen allocation operate on the different phytomers in parallel but at unsynchronized steps, separated by a thermal period. An important phenological phase is identified for the oil palm – the storage growth period of bud and "spear" leaves which are photosynthetically inactive before expansion. Agricultural practices such as transplanting, fertilization and leaf pruning are represented. Parameters introduced for the oil palm were calibrated and validated with field measurements of leaf area index (LAI), yield and net primary production (NPP) from Sumatra, Indonesia. In calibration with a mature oil palm plantation, the cumulative yields from 2005 to 2014 matched notably well between simulation and observation (mean percentage error = 3 %). Simulated inter-annual dynamics of PFT-level and phytomer-level LAI were both within the range of field measurements. Validation from eight independent oil palm sites shows the ability of the model to adequately predict the average leaf growth and fruit yield across sites and sufficiently represent the significant nitrogen- and age-related site-to-site variability in NPP and yield. Results also indicate that seasonal dynamics of yield and remaining small-scale site-to-site variability of NPP are driven by processes not yet implemented in the model or reflected in the input data. The new sub-canopy structure and phenology and allocation functions in CLM-Palm allow exploring the effects of tropical land-use change, from natural ecosystems to oil palm plantations, on carbon, water and energy cycles and regional climate.

A Landsat TM image, part of a rainforest located in the state of Rondonia, Brazil, was analysed using Fourier transform techniques in order to extract textural features. The lattice-like texture pattern recorded was then compared with the geological structural trends of the area, and measures of the soil surface moisture treated by geostatistical analysis. The consistency between the different sets of analysis suggests that the spatial distribution of moisture at the soil surface is related to the structure of the bedrock. The near infrared channel of the Landsat TM may be used to record trends in water content distributions at the soil surface under homogeneous forest canopies. [ABSTRACT FROM AUTHOR]

Plant Toll/interleukin-1 receptor/resistance protein (TIR) type nucleotide-binding and leucine-rich repeat immune receptors (NLRs) require enhanced disease susceptibility 1 (EDS1) family proteins and the helper NLRs NRG1 and ADR1 for immune activation. We show that the Nb EDS1- Nb SAG101b- Nb NRG1 signaling pathway in N. benthamiana is necessary for cell death signaling by TIR-NLRs from a range of plant species, suggesting a universal requirement for this module in TIR-NLR-mediated cell death in N. benthamiana. We also find that TIR domains physically associate with Nb EDS1, Nb PAD4, and Nb SAG101 in planta , independently of each other. Furthermore, Nb NRG1 associates with Nb SAG101b, but not with other EDS1 family members, via its C-terminal EP domain. Physical interaction between activated TIRs and EDS1 signaling complexes may facilitate the transfer of low abundance products of TIR catalytic activity or alter TIR catalytic activity to favor the production of EDS1 heterodimer ligands., Competing Interests: The authors declare no competing interests., (© 2024 The Author(s).)

Recent work shed light on how plant intracellular immune receptors of the nucleotide-binding leucine-rich repeat (NLR) family are activated upon pathogen effector recognition to trigger immune responses. Activation of Toll-interleukin-1 receptor (TIR) domain-containing NLRs (TNLs) induces receptor oligomerization and close proximity of the TIR domain, which is required for TIR enzymatic activity. TIR-catalyzed small signaling molecules bind to EDS1 family heterodimers and subsequently activate downstream helper NLRs, which function as Ca 2+ permeable channel to activate immune responses eventually leading to cell death. Subcellular localization requirements of TNLs and signaling partners are not well understood, although they are required to understand fully the mechanisms underlying NLR early signaling. TNLs show diverse subcellular localization while EDS1 shows nucleocytosolic localization. Here, we studied the impact of TIR and EDS1 mislocalization on the signaling activation of different TNLs. In Nicotiana benthamiana, our results suggest that close proximity of TIR domains isolated from flax L6 and Arabidopsis RPS4 and SNC1 TNLs drives signaling activation from different cell compartments. Nevertheless, both Golgi-membrane anchored L6 and nucleocytosolic RPS4 have the same requirements for EDS1 subcellular localization in Arabidopsis thaliana. By using mislocalized variants of EDS1, we found that autoimmune L6 and RPS4 TIR domain can induce seedling cell death when EDS1 is present in the cytosol. However, when EDS1 is restricted to the nucleus, both induce a stunting phenotype but no cell death. Our data point out the importance of thoroughly investigating the dynamics of TNLs and signaling partners subcellular localization to understand TNL signaling fully., (© 2023 Society for Experimental Biology and John Wiley & Sons Ltd.)

Ruminant livestock are an important source of anthropogenic methane (CH 4 ). Decreasing the emissions of enteric CH 4 from ruminant production is strategic to limit the global temperature increase to 1.5°C by 2050. Research in the area of enteric CH 4 mitigation has grown exponentially in the last 2 decades, with various strategies for enteric CH 4 abatement being investigated: production intensification, dietary manipulation (including supplementation and processing of concentrates and lipids, and management of forage and pastures), rumen manipulation (supplementation of ionophores, 3-nitrooxypropanol, macroalgae, alternative electron acceptors, and phytochemicals), and selection of low-CH 4 -producing animals. Other enteric CH 4 mitigation strategies are at earlier stages of research but rapidly developing. Herein, we discuss and analyze the current status of available enteric CH 4 mitigation strategies with an emphasis on opportunities and barriers to their implementation in confined and partial grazing production systems, and in extensive and fully grazing production systems. For each enteric CH 4 mitigation strategy, we discuss its effectiveness to decrease total CH 4 emissions and emissions on a per animal product basis, safety issues, impacts on the emissions of other greenhouse gases, as well as other economic, regulatory, and societal aspects that are key to implementation. Most research has been conducted with confined animals, and considerably more research is needed to develop, adapt, and evaluate antimethanogenic strategies for grazing systems. In general, few options are currently available for extensive production systems without feed supplementation. Continuous research and development are needed to develop enteric CH 4 mitigation strategies that are locally applicable. Information is needed to calculate carbon footprints of interventions on a regional basis to evaluate the impact of mitigation strategies on net greenhouse gas emissions. Economically affordable enteric CH 4 mitigation solutions are urgently needed. Successful implementation of safe and effective antimethanogenic strategies will also require delivery mechanisms and adequate technical support for producers, as well as consumer involvement and acceptance. The most appropriate metrics should be used in quantifying the overall climate outcomes associated with mitigation of enteric CH 4 emissions. A holistic approach is required, and buy-in is needed at all levels of the supply chain., (The Authors. Published by Elsevier Inc. and Fass Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).)

Plants can detect microbial molecules via surface-localized pattern-recognition receptors (PRRs) and intracellular immune receptors from the nucleotide-binding, leucine-rich repeat receptor (NLR) family. The corresponding pattern-triggered (PTI) and effector-triggered (ETI) immunity were long considered separate pathways, although they converge on largely similar cellular responses, such as calcium influx and overlapping gene reprogramming. A number of studies recently uncovered genetic and molecular interconnections between PTI and ETI, highlighting the complexity of the plant immune network. Notably, PRR- and NLR-mediated immune responses require and potentiate each other to reach an optimal immune output. How PTI and ETI connect to confer robust immunity in different plant species, including crops will be an exciting future research area., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022. Published by Elsevier Ltd.)

Most national GHG inventories estimating direct N 2 O emissions from managed soils rely on a default Tier 1 emission factor (EF 1 ) amounting to 1% of nitrogen inputs. Recent research has, however, demonstrated the potential for refining the EF 1 considering variables that are readily available at national scales. Building on existing reviews, we produced a large dataset (n = 848) enriched in dry and low latitude tropical climate observations as compared to former global efforts and disaggregated the EF 1 according to most meaningful controlling factors. Using spatially explicit N fertilizer and manure inputs, we also investigated the implications of using the EF 1 developed as part of this research and adopted by the 2019 IPCC refinement report. Our results demonstrated that climate is a major driver of emission, with an EF 1 three times higher in wet climates (0.014, 95% CI 0.011-0.017) than in dry climates (0.005, 95% CI 0.000-0.011). Likewise, the form of the fertilizer markedly modulated the EF 1 in wet climates, where the EF 1 for synthetic and mixed forms (0.016, 95% CI 0.013-0.019) was also almost three times larger than the EF 1 for organic forms (0.006; 95% CI 0.001-0.011). Other factors such as land cover and soil texture, C content, and pH were also important regulators of the EF 1 . The uncertainty associated with the disaggregated EF 1 was considerably reduced as compared to the range in the 2006 IPCC guidelines. Compared to estimates from the 2006 IPCC EF 1 , emissions based on the 2019 IPCC EF 1 range from 15% to 46% lower in countries dominated by dry climates to 7%-37% higher in countries with wet climates and high synthetic N fertilizer consumption. The adoption of the 2019 IPCC EF 1 will allow parties to improve the accuracy of emissions' inventories and to better target areas for implementing mitigation strategies., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

To respect the Paris agreement targeting a limitation of global warming below 2°C by 2100, and possibly below 1.5°C, drastic reductions of greenhouse gas emissions are mandatory but not sufficient. Large-scale deployment of other climate mitigation strategies is also necessary. Among these, increasing soil organic carbon (SOC) stocks is an important lever because carbon in soils can be stored for long periods and land management options to achieve this already exist and have been widely tested. However, agricultural soils are also an important source of nitrous oxide (N 2 O), a powerful greenhouse gas, and increasing SOC may influence N 2 O emissions, likely causing an increase in many cases, thus tending to offset the climate change benefit from increased SOC storage. Here we review the main agricultural management options for increasing SOC stocks. We evaluate the amount of SOC that can be stored as well as resulting changes in N 2 O emissions to better estimate the climate benefits of these management options. Based on quantitative data obtained from published meta-analyses and from our current level of understanding, we conclude that the climate mitigation induced by increased SOC storage is generally overestimated if associated N 2 O emissions are not considered but, with the exception of reduced tillage, is never fully offset. Some options (e.g. biochar or non-pyrogenic C amendment application) may even decrease N 2 O emissions., (© 2020 John Wiley & Sons Ltd.)

Plant and animal intracellular nucleotide-binding, leucine-rich repeat (NLR) immune receptors detect pathogen-derived molecules and activate defense. Plant NLRs can be divided into several classes based upon their N-terminal signaling domains, including TIR (Toll-like, Interleukin-1 receptor, Resistance protein)- and CC (coiled-coil)-NLRs. Upon ligand detection, mammalian NAIP and NLRC4 NLRs oligomerize, forming an inflammasome that induces proximity of its N-terminal signaling domains. Recently, a plant CC-NLR was revealed to form an inflammasome-like hetero-oligomer. To further investigate plant NLR signaling mechanisms, we fused the N-terminal TIR domain of several plant NLRs to the N terminus of NLRC4. Inflammasome-dependent induced proximity of the TIR domain in planta initiated defense signaling. Thus, induced proximity of a plant TIR domain imposed by oligomerization of a mammalian inflammasome is sufficient to activate authentic plant defense. Ligand detection and inflammasome formation is maintained when the known components of the NLRC4 inflammasome is transferred across kingdoms, indicating that NLRC4 complex can robustly function without any additional mammalian proteins. Additionally, we found NADase activity of a plant TIR domain is necessary for plant defense activation, but NADase activity of a mammalian or a bacterial TIR is not sufficient to activate defense in plants., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Adoption of no-till management on croplands has become a controversial approach for storing carbon in soil due to conflicting findings. Yet, no-till is still promoted as a management practice to stabilize the global climate system from additional change due to anthropogenic greenhouse gas emissions, including the 4 per mille initiative promoted through the UN Framework Convention on Climate Change. We evaluated the body of literature surrounding this practice, and found that SOC storage can be higher under no-till management in some soil types and climatic conditions even with redistribution of SOC, and contribute to reducing net greenhouse gas emissions. However, uncertainties tend to be large, which may make this approach less attractive as a contributor to stabilize the climate system compared to other options. Consequently, no-till may be better viewed as a method for reducing soil erosion, adapting to climate change, and ensuring food security, while any increase in SOC storage is a co-benefit for society in terms of reducing greenhouse gas emissions.

Tropical rainforest soils harbor a considerable diversity of soil fauna that contributes to emissions of N 2 O. Despite their ecological dominance, there is limited information available about the contribution of epigeal ant mounds to N 2 O emissions in these tropical soils. This study aimed to determine whether ant mounds contribute to local soil N emissions in the tropical humid rainforest. N 2 O emission was determined in vitro from individual live ants, ant-processed mound soils, and surrounding reference soils for two trophically distinct and abundant ant species: the leaf-cutting Atta mexicana and omnivorous Solenopsis geminata. The abundance of total bacteria, nitrifiers (AOA and AOB), and denitrifiers (nirK, nirS, and nosZ) was estimated in these soils using quantitative PCR, and their respective mineral N contents determined. There was negligible N 2 O emission detected from live ant individuals. However, the mound soils of both species emitted significantly greater (3-fold) amount of N 2 O than their respective surrounding reference soils. This emission increased significantly up to 6-fold in the presence of acetylene, indicating that, in addition to N 2 O, dinitrogen (N 2 ) is also produced from these mound soils at an equivalent rate (N 2 O/N 2  = 0.57). Functional gene abundance (nitrifiers and denitrifiers) and mineral N pools (ammonium and nitrate) were significantly greater in mound soils than in their respective reference soils. Furthermore, in the light of the measured parameters and their correlation trends, nitrification and denitrification appeared to represent the major N 2 O-producing microbial processes in ant mound soils. The ant mounds were estimated to contribute from 0.1 to 3.7% of the total N 2 O emissions of tropical rainforest soils.

The effector protein AvrP is secreted by the flax rust fungal pathogen (Melampsora lini) and recognized specifically by the flax (Linum usitatissimum) P disease resistance protein, leading to effector-triggered immunity. To investigate the biological function of this effector and the mechanisms of specific recognition by the P resistance protein, we determined the crystal structure of AvrP. The structure reveals an elongated zinc-finger-like structure with a novel interleaved zinc-binding topology. The residues responsible for zinc binding are conserved in AvrP effector variants and mutations of these motifs result in a loss of P-mediated recognition. The first zinc-coordinating region of the structure displays a positively charged surface and shows some limited similarities to nucleic acid-binding and chromatin-associated proteins. We show that the majority of the AvrP protein accumulates in the plant nucleus when transiently expressed in Nicotiana benthamiana cells, suggesting a nuclear pathogenic function. Polymorphic residues in AvrP and its allelic variants map to the protein surface and could be associated with differences in recognition specificity. Several point mutations of residues on the non-conserved surface patch result in a loss of recognition by P, suggesting that these residues are required for recognition., (© 2017 BSPP AND JOHN WILEY & SONS LTD.)

Race-specific resistance genes protect the global wheat crop from stem rust disease caused by Puccinia graminis f. sp. tritici ( Pgt ) but are often overcome owing to evolution of new virulent races of the pathogen. To understand virulence evolution in Pgt , we identified the protein ligand (AvrSr50) recognized by the Sr50 resistance protein. A spontaneous mutant of Pgt virulent to Sr50 contained a 2.5 mega-base pair loss-of-heterozygosity event. A haustorial secreted protein from this region triggers Sr50 -dependent defense responses in planta and interacts directly with the Sr50 protein. Virulence alleles of AvrSr50 have arisen through DNA insertion and sequence divergence, and our data provide molecular evidence that in addition to sexual recombination, somatic exchange can play a role in the emergence of new virulence traits in Pgt ., (Copyright © 2017 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

The first plant disease resistance (R) genes were identified and cloned more than two decades ago. Since then, many more R genes have been identified and characterized in numerous plant pathosystems. Most of these encode members of the large family of intracellular NLRs (NOD-like receptors), which also includes animal immune receptors. New discoveries in this expanding field of research provide new elements for our understanding of plant NLR function. But what do we know about plant NLR function today? Genetic, structural, and functional analyses have uncovered a number of commonalities and differences in pathogen recognition strategies as well as how NLRs are regulated and activate defense signaling, but many unknowns remain. This review gives an update on the latest discoveries and breakthroughs in this field, with an emphasis on structural findings and some comparison to animal NLRs, which can provide additional insights and paradigms in plant NLR function.

The self-association of Toll/interleukin-1 receptor/resistance protein (TIR) domains has been implicated in signaling in plant and animal immunity receptors. Structure-based studies identified different TIR-domain dimerization interfaces required for signaling of the plant nucleotide-binding oligomerization domain-like receptors (NLRs) L6 from flax and disease resistance protein RPS4 from Arabidopsis Here we show that the crystal structure of the TIR domain from the Arabidopsis NLR suppressor of npr1-1, constitutive 1 (SNC1) contains both an L6-like interface involving helices αD and αE (DE interface) and an RPS4-like interface involving helices αA and αE (AE interface). Mutations in either the AE- or DE-interface region disrupt cell-death signaling activity of SNC1, L6, and RPS4 TIR domains and full-length L6 and RPS4. Self-association of L6 and RPS4 TIR domains is affected by mutations in either region, whereas only AE-interface mutations affect SNC1 TIR-domain self-association. We further show two similar interfaces in the crystal structure of the TIR domain from the Arabidopsis NLR recognition of Peronospora parasitica 1 (RPP1). These data demonstrate that both the AE and DE self-association interfaces are simultaneously required for self-association and cell-death signaling in diverse plant NLRs.

Plants possess intracellular immune receptors designated "nucleotide-binding domain and leucine-rich repeat" (NLR) proteins that translate pathogen-specific recognition into disease-resistance signaling. The wheat immune receptors Sr33 and Sr50 belong to the class of coiled-coil (CC) NLRs. They confer resistance against a broad spectrum of field isolates of Puccinia graminis f. sp. tritici, including the Ug99 lineage, and are homologs of the barley powdery mildew-resistance protein MLA10. Here, we show that, similarly to MLA10, the Sr33 and Sr50 CC domains are sufficient to induce cell death in Nicotiana benthamiana Autoactive CC domains and full-length Sr33 and Sr50 proteins self-associate in planta In contrast, truncated CC domains equivalent in size to an MLA10 fragment for which a crystal structure was previously determined fail to induce cell death and do not self-associate. Mutations in the truncated region also abolish self-association and cell-death signaling. Analysis of Sr33 and Sr50 CC domains fused to YFP and either nuclear localization or nuclear export signals in N benthamiana showed that cell-death induction occurs in the cytosol. In stable transgenic wheat plants, full-length Sr33 proteins targeted to the cytosol provided rust resistance, whereas nuclear-targeted Sr33 was not functional. These data are consistent with CC-mediated induction of both cell-death signaling and stem rust resistance in the cytosolic compartment, whereas previous research had suggested that MLA10-mediated cell-death and disease resistance signaling occur independently, in the cytosol and nucleus, respectively., Competing Interests: The authors declare no conflict of interest.

NOD-like receptors (NLRs) are central components of the plant immune system. L6 is a Toll/interleukin-1 receptor (TIR) domain-containing NLR from flax (Linum usitatissimum) conferring immunity to the flax rust fungus. Comparison of L6 to the weaker allele L7 identified two polymorphic regions in the TIR and the nucleotide binding (NB) domains that regulate both effector ligand-dependent and -independent cell death signaling as well as nucleotide binding to the receptor. This suggests that a negative functional interaction between the TIR and NB domains holds L7 in an inactive/ADP-bound state more tightly than L6, hence decreasing its capacity to adopt the active/ATP-bound state and explaining its weaker activity in planta. L6 and L7 variants with a more stable ADP-bound state failed to bind to AvrL567 in yeast two-hybrid assays, while binding was detected to the signaling active variants. This contrasts with current models predicting that effectors bind to inactive receptors to trigger activation. Based on the correlation between nucleotide binding, effector interaction, and immune signaling properties of L6/L7 variants, we propose that NLRs exist in an equilibrium between ON and OFF states and that effector binding to the ON state stabilizes this conformation, thereby shifting the equilibrium toward the active form of the receptor to trigger defense signaling., (© 2016 American Society of Plant Biologists. All rights reserved.)

The impact of deforestation on soil organic carbon (SOC) stocks is important in the context of climate change and agricultural soil use. Trends of SOC stock changes after agroecosystem establishment vary according to the spatial scale considered, and factors explaining these trends may differ sometimes according to meta-analyses. We have reviewed the knowledge about changes in SOC stocks in Amazonia after the establishment of pasture or cropland, sought relationships between observed changes and soil, climatic variables and management practices, and synthesized the δ 13 C measured in pastures. Our dataset consisted of 21 studies mostly synchronic, across 52 sites (Brazil, Colombia, French Guiana, Suriname), totalling 70 forest-agroecosystem comparisons. We found that pastures (n = 52, mean age = 17.6 years) had slightly higher SOC stocks than forest (+6.8 ± 3.1 %), whereas croplands (n = 18, mean age = 8.7 years) had lower SOC stocks than forest (-8.5 ± 2.9 %). Annual precipitation and SOC stocks under forest had no effect on the SOC changes in the agroecosystems. For croplands, we found a lower SOC loss than other meta-analyses, but the short time period after deforestation here could have reduced this loss. There was no clear effect of tillage on the SOC response. Management of pastures, whether they were degraded/nominal/improved, had no significant effect on SOC response. δ 13 C measurements on 16 pasture chronosequences showed that decay of forest-derived SOC was variable, whereas pasture-derived SOC was less so and was characterized by an accumulation plateau of 20 Mg SOC ha -1 after 20 years. The large uncertainties in SOC response observed could be derived from the chronosequence approach, sensitive to natural soil variability and to human management practices. This study emphasizes the need for diachronic and long-term studies, associated with better knowledge of agroecosystem management., (© 2015 John Wiley & Sons Ltd.)

Plant immunity is often triggered by the specific recognition of pathogen effectors by intracellular nucleotide-binding, leucine-rich repeat receptors (NLR). Plant NLRs contain an N-terminal signaling domain that is mostly represented by either a Toll-interleukin1 receptor (TIR) domain or a coiled coil (CC) domain. In many cases, single NLR proteins are sufficient for both effector recognition and signaling activation. However, many paired NLRs have now been identified where both proteins are required to confer resistance to pathogens. Recent detailed studies on the Arabidopsis thaliana TIR-NLR pair RRS1 and RPS4 and on the rice CC-NLR pair RGA4 and RGA5 have revealed for the first time how such protein pairs function together. In both cases, the paired partners interact physically to form a hetero-complex receptor in which each partner plays distinct roles in effector recognition or signaling activation, highlighting a conserved mode of action of NLR pairs across both monocotyledonous and dicotyledonous plants. We also describe an "integrated decoy" model for the function of these receptor complexes. In this model, a plant protein targeted by an effector has been duplicated and fused to one member of the NLR pair, where it acts as a bait to trigger defense signaling by the second NLR upon effector binding. This mechanism may be common to many other plant NLR pairs.

Plant resistance proteins of the class of nucleotide-binding and leucine-rich repeat domain proteins (NB-LRRs) are immune sensors which recognize pathogen-derived molecules termed avirulence (AVR) proteins. We show that RGA4 and RGA5, two NB-LRRs from rice, interact functionally and physically to mediate resistance to the fungal pathogen Magnaporthe oryzae and accomplish different functions in AVR recognition. RGA4 triggers an AVR-independent cell death that is repressed in the presence of RGA5 in both rice protoplasts and Nicotiana benthamiana. Upon recognition of the pathogen effector AVR-Pia by direct binding to RGA5, repression is relieved and cell death occurs. RGA4 and RGA5 form homo- and hetero-complexes and interact through their coiled-coil domains. Localization studies in rice protoplast suggest that RGA4 and RGA5 localize to the cytosol. Upon recognition of AVR-Pia, neither RGA4 nor RGA5 is re-localized to the nucleus. These results establish a model for the interaction of hetero-pairs of NB-LRRs in plants: RGA4 mediates cell death activation, while RGA5 acts as a repressor of RGA4 and as an AVR receptor., (© 2014 The Authors.)

Cytoplasmic plant immune receptors recognize specific pathogen effector proteins and initiate effector-triggered immunity. In Arabidopsis, the immune receptors RPS4 and RRS1 are both required to activate defense to three different pathogens. We show that RPS4 and RRS1 physically associate. Crystal structures of the N-terminal Toll-interleukin-1 receptor/resistance (TIR) domains of RPS4 and RRS1, individually and as a heterodimeric complex (respectively at 2.05, 1.75, and 2.65 angstrom resolution), reveal a conserved TIR/TIR interaction interface. We show that TIR domain heterodimerization is required to form a functional RRS1/RPS4 effector recognition complex. The RPS4 TIR domain activates effector-independent defense, which is inhibited by the RRS1 TIR domain through the heterodimerization interface. Thus, RPS4 and RRS1 function as a receptor complex in which the two components play distinct roles in recognition and signaling.

The Toll/interleukin-1 receptor (TIR) domain is a protein-protein interaction domain that is found in both animal and plant immune receptors. The N-terminal TIR domain from the nucleotide-binding (NB)-leucine-rich repeat (LRR) class of plant disease-resistance (R) proteins has been shown to play an important role in defence signalling. Recently, the crystal structure of the TIR domain from flax R protein L6 was determined and this structure, combined with functional studies, demonstrated that TIR-domain homodimerization is a requirement for function of the R protein L6. To advance the molecular understanding of the function of TIR domains in R-protein signalling, the protein expression, purification, crystallization and X-ray diffraction analyses of the TIR domains of the Arabidopsis thaliana R proteins RPS4 (resistance to Pseudomonas syringae 4) and RRS1 (resistance to Ralstonia solanacearum 1) and the resistance-like protein SNC1 (suppressor of npr1-1, constitutive 1) are reported here. RPS4 and RRS1 function cooperatively as a dual resistance-protein system that prevents infection by three distinct pathogens. SNC1 is implicated in resistance pathways in Arabidopsis and is believed to be involved in transcriptional regulation through its interaction with the transcriptional corepressor TPR1 (Topless-related 1). The TIR domains of all three proteins have successfully been expressed and purified as soluble proteins in Escherichia coli. Plate-like crystals of the RPS4 TIR domain were obtained using PEG 3350 as a precipitant; they diffracted X-rays to 2.05 Å resolution, had the symmetry of space group P1 and analysis of the Matthews coefficient suggested that there were four molecules per asymmetric unit. Tetragonal crystals of the RRS1 TIR domain were obtained using ammonium sulfate as a precipitant; they diffracted X-rays to 1.75 Å resolution, had the symmetry of space group P4(1)2(1)2 or P4(3)2(1)2 and were most likely to contain one molecule per asymmetric unit. Crystals of the SNC1 TIR domain were obtained using PEG 3350 as a precipitant; they diffracted X-rays to 2.20 Å resolution and had the symmetry of space group P4(1)2(1)2 or P4(3)2(1)2, with two molecules predicted per asymmetric unit. These results provide a good foundation to advance the molecular and structural understanding of the function of the TIR domain in plant innate immunity.

Although highly weathered soils cover considerable areas in tropical regions, little is known about exploration by roots in deep soil layers. Intensively managed Eucalyptus plantations are simple forest ecosystems that can provide an insight into the belowground growth strategy of fast-growing tropical trees. Fast exploration of deep soil layers by eucalypt fine roots may contribute to achieving a gross primary production that is among the highest in the world for forests. Soil exploration by fine roots down to a depth of 10 m was studied throughout the complete cycle in Eucalyptus grandis plantations managed in short rotation. Intersects of fine roots, less than 1 mm in diameter, and medium-sized roots, 1-3 mm in diameter, were counted on trench walls in a chronosequence of 1-, 2-, 3.5-, and 6-year-old plantations on a sandy soil, as well as in an adjacent 6-year-old stand growing in a clayey soil. Two soil profiles were studied down to a depth of 10 m in each stand (down to 6 m at ages 1 and 2 years) and 4 soil profiles down to 1.5-3.0 m deep. The root intersects were counted on 224 m(2) of trench walls in 15 pits. Monitoring the soil water content showed that, after clear-cutting, almost all the available water stored down to a depth of 7 m was taken up by tree roots within 1.1 year of planting. The soil space was explored intensively by fine roots down to a depth of 3 m from 1 year after planting, with an increase in anisotropy in the upper layers throughout the rotation. About 60% of fine root intersects were found at a depth of more than 1 m, irrespective of stand age. The root distribution was isotropic in deep soil layers and kriged maps showed fine root clumping. A considerable volume of soil was explored by fine roots in eucalypt plantations on deep tropical soils, which might prevent water and nutrient losses by deep drainage after canopy closure and contribute to maximizing resource uses.

No information regarding the management of manure from beef cattle feedlots is available for Brazil. To fill this knowledge gap, a survey of 73 feedlots was conducted in 7 Brazilian states. In this survey, questions were asked regarding animal characteristics, their diets, and manure handling management from generation to disposal. These feedlots finished 831,450 animals in 2010. The predominant breed fed was Nellore, with average feeding periods of 60 to 135 d. Corn was the primary source of grain used in the feedlot diets (76% of surveyed animals) with concentrate inclusion levels ranging from 81 to 90% (38% of surveyed animals). The most representative manure management practice was the removal of manure from pens only at the end of the feeding period. Subsequently, the manure was stored in mounds before being applied to crop and pasture lands. Runoff, mainly from rainwater, was collected in retention ponds and used for agriculture. However, the quantity of runoff was not known. Manure was composted for only 20% of the animals in the survey and was treated in anaerobic digesters for only 1% of the animals. Manure from 59% of the cattle surveyed was used as fertilizer, providing a cost savings over the use of synthetic fertilizers. Overall, chemical analysis of the manure before application to fields was conducted for the manure of 56% of the surveyed animals, but the exact quantity applied (per hectare) was unknown for 48%. Feedlots representing 48% of the surveyed animals noted similar or greater crop and pasture yields when using manure, rather than synthetic fertilizers. In addition, 32% mentioned an increase in soil organic matter. Feedlots representing 88% of the surveyed cattle indicated that information concerning management practices that improve manure use efficiency is lacking. Feedlots representing 93% of the animals in the survey reported having basic information regarding the generation of energy and fertilizer with anaerobic digesters. However, only 1 feedlot implemented this technology. In conclusion, the manure management evaluated in this study represents an important indirect economic benefit that was represented by decreased use of synthetic fertilizers in crops. However, little attention was given to the specific treatments and environmental impacts of handling manure. This survey provides information that should assist in the development of better research practices and broader application of future models.

To investigate the role of N-terminal domains of plant disease resistance proteins in membrane targeting, the N termini of a number of Arabidopsis and flax disease resistance proteins were fused to green fluorescent protein (GFP) and the fusion proteins localized in planta using confocal microscopy. The N termini of the Arabidopsis RPP1-WsB and RPS5 resistance proteins and the PBS1 protein, which is required for RPS5 resistance, targeted GFP to the plasma membrane, and mutation of predicted myristoylation and potential palmitoylation sites resulted in a shift to nucleocytosolic localization. The N-terminal domain of the membrane-attached Arabidopsis RPS2 resistance protein was targeted incompletely to the plasma membrane. In contrast, the N-terminal domains of the Arabidopsis RPP1-WsA and flax L6 and M resistance proteins, which carry predicted signal anchors, were targeted to the endomembrane system, RPP1-WsA to the endoplasmic reticulum and the Golgi apparatus, L6 to the Golgi apparatus, and M to the tonoplast. Full-length L6 was also targeted to the Golgi apparatus. Site-directed mutagenesis of six nonconserved amino acid residues in the signal anchor domains of L6 and M was used to change the localization of the L6 N-terminal fusion protein to that of M and vice versa, showing that these residues control the targeting specificity of the signal anchor. Replacement of the signal anchor domain of L6 by that of M did not affect L6 protein accumulation or resistance against flax rust expressing AvrL567 but removal of the signal anchor domain reduced L6 protein accumulation and L6 resistance, suggesting that membrane attachment is required to stabilize the L6 protein.

L locus resistance (R) proteins are nucleotide binding (NB-ARC) leucine-rich repeat (LRR) proteins from flax (Linum usitatissimum) that provide race-specific resistance to the causal agent of flax rust disease, Melampsora lini. L5 and L6 are two alleles of the L locus that directly recognize variants of the fungal effector AvrL567. In this study, we have investigated the molecular details of this recognition by site-directed mutagenesis of AvrL567 and construction of chimeric L proteins. Single, double and triple mutations of polymorphic residues in a variety of AvrL567 variants showed additive effects on recognition strength, suggesting that multiple contact points are involved in recognition. Domain-swap experiments between L5 and L6 show that specificity differences are determined by their corresponding LRR regions. Most positively selected amino acid sites occur in the N- and C-terminal LRR units, and polymorphisms in the first seven and last four LRR units contribute to recognition specificity of L5 and L6 respectively. This further confirms that multiple, additive contact points occur between AvrL567 variants and either L5 or L6. However, we also observed that recognition of AvrL567 is affected by co-operative polymorphisms between both adjacent and distant domains of the R protein, including the TIR, ARC and LRR domains, implying that these residues are involved in intramolecular interactions to optimize detection of the pathogen and defense signal activation. We suggest a model where Avr ligand interaction directly competes with intramolecular interactions to cause activation of the R protein.

Plant disease resistance can be triggered by specific recognition of microbial effectors by plant nucleotide binding-leucine rich repeat (NB-LRR) receptors. Over the last few years, many efforts have greatly improved the understanding of effector and NB-LRR function, but have left a lot of questions as to how effector perception activates NB-LRR induction of defense signaling. This review describes exciting new findings showing similarities and differences in function of diverse plant NB-LRR proteins in terms of pathogen recognition and where and how resistance proteins are activated. Localization studies have shown that some NB-LRRs can activate signaling from the cytosol while others act in the nucleus. Also, the structural determination of two NB-LRR signaling domains demonstrated that receptor oligomerization is fundamental for the activation of resistance signaling., (Crown Copyright © 2011. Published by Elsevier Ltd. All rights reserved.)

The Toll/interleukin-1 receptor (TIR) domain occurs in animal and plant immune receptors. In the animal Toll-like receptors, homodimerization of the intracellular TIR domain is required for initiation of signaling cascades leading to innate immunity. By contrast, the role of the TIR domain in cytoplasmic nucleotide-binding/leucine-rich repeat (NB-LRR) plant immune resistance proteins is poorly understood. L6 is a TIR-NB-LRR resistance protein from flax (Linum usitatissimum) that confers resistance to the flax rust phytopathogenic fungus (Melampsora lini). We determine the crystal structure of the L6 TIR domain and show that, although dispensable for pathogenic effector protein recognition, the TIR domain alone is both necessary and sufficient for L6 immune signaling. We demonstrate that the L6 TIR domain self-associates, most likely forming a homodimer. Analysis of the structure combined with site-directed mutagenesis suggests that self-association is a requirement for immune signaling and reveals distinct surface regions involved in self-association, signaling, and autoregulation., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Type III effector proteins from bacterial pathogens manipulate components of host immunity to suppress defence responses and promote pathogen development. In plants, host proteins targeted by some effectors called avirulence proteins are surveyed by plant disease resistance proteins referred to as "guards". The Ralstonia solanacearum effector protein PopP2 triggers immunity in Arabidopsis following its perception by the RRS1-R resistance protein. Here, we show that PopP2 interacts with RRS1-R in the nucleus of living plant cells. PopP2 belongs to the YopJ-like family of cysteine proteases, which share a conserved catalytic triad that includes a highly conserved cysteine residue. The catalytic cysteine mutant PopP2-C321A is impaired in its avirulence activity although it is still able to interact with RRS1-R. In addition, PopP2 prevents proteasomal degradation of RRS1-R, independent of the presence of an integral PopP2 catalytic core. A liquid chromatography/tandem mass spectrometry analysis showed that PopP2 displays acetyl-transferase activity leading to its autoacetylation on a particular lysine residue, which is well conserved among all members of the YopJ family. These data suggest that this lysine residue may correspond to a key binding site for acetyl-coenzyme A required for protein activity. Indeed, mutation of this lysine in PopP2 abolishes RRS1-R-mediated immunity. In agreement with the guard hypothesis, our results favour the idea that activation of the plant immune response by RRS1-R depends not only on the physical interaction between the two proteins but also on its perception of PopP2 enzymatic activity.

As in nearly every discipline of plant biology, new insights are constantly changing our understanding of plant immunity. It is now clear that plant immunity is controlled by two layers of inducible responses: basal responses triggered by conserved microbial features and specific responses triggered by gene-for-gene recognition of pathogen effector proteins by host resistance (R) proteins. The nucleotide-binding domain leucine-rich repeat (NB-LRR) class of R proteins plays a major role in the combat against a wide range of plant pathogens. The variation that has been generated and is maintained within these conserved proteins has diversified their specificity, subcellular localisations, activation and recognition mechanisms, allowing them to specifically adapt to different plant-pathogen interaction systems. This review addresses recent advances in the molecular role of NB-LRR proteins in pathogen recognition and activation of plant defence responses.

Bacterial wilt, a disease impacting cultivated crops worldwide, is caused by the pathogenic bacterium Ralstonia solanacearum. PopP2 (for Pseudomonas outer protein P2) is an R. solanacearum type III effector that belongs to the YopJ/AvrRxv protein family and interacts with the Arabidopsis thaliana RESISTANT TO RALSTONIA SOLANACEARUM 1-R (RRS1-R) resistance protein. RRS1-R contains the Toll/Interleukin1 receptor-nucleotide binding site-Leu-rich repeat domains found in several cytoplasmic R proteins and a C-terminal WRKY DNA binding domain. In this study, we identified the Arabidopsis Cys protease RESPONSIVE TO DEHYDRATION19 (RD19) as being a PopP2-interacting protein whose expression is induced during infection by R. solanacearum. An Arabidopsis rd19 mutant in an RRS1-R genetic background is compromised in resistance to the bacterium, indicating that RD19 is required for RRS1-R-mediated resistance. RD19 normally localizes in mobile vacuole-associated compartments and, upon coexpression with PopP2, is specifically relocalized to the plant nucleus, where the two proteins physically interact. No direct physical interaction between RRS1-R and RD19 in the presence of PopP2 was detected in the nucleus as determined by Förster resonance energy transfer. We propose that RD19 associates with PopP2 to form a nuclear complex that is required for activation of the RRS1-R-mediated resistance response.

This paper serves two purposes: it provides a summarized scientific history of carbon sequestration in relation to the soil-plant system and gives a commentary on organic wastes and SOC sequestration. The concept of soil organic carbon (SOC) sequestration has its roots in: (i) the experimental work of Lundegårdh, particularly his in situ measurements of CO2 fluxes at the soil-plant interface (1924, 1927, 1930); (ii) the first estimates of SOC stocks at the global level made by Waksman [Waksman, S.A., 1938. Humus. Origin, Chemical Composition and Importance in Nature, second ed. revised. Williams and Wilkins, Baltimore, p. 526] and Rubey [Rubey, W.W., 1951. Geologic history of sea water. Bulletin of the Geological Society of America 62, 1111-1148]; (iii) the need for models dealing with soil organic matter (SOM) or SOC dynamics beginning with a conceptual SOM model by De Saussure (1780-1796) followed by the mathematical models of Jenny [Jenny, H., 1941. Factors of Soil Formation: a System of Quantitative Pedology. Dover Publications, New York, p. 288], Hénin and Dupuis [Hénin, S., Dupuis, M., 1945. Essai de bilan de la matière organique. Annales d'Agronomie 15, 17-29] and more recently the RothC [Jenkinson, D.S., Rayner, J.H., 1977. The turnover of soil organic matter in some of the Rothamsted classical experiments. Soil Science 123 (5), 298-305] and Century [Parton, W.J., Schimel, D.S., Cole, C.V., Ojima, D.S., 1987. Analysis of factors controlling soil organic matter levels in great plains grasslands. Soil Science Society of America Journal 51 (5), 1173-1179] models. The establishment of a soil C sequestration balance is not straightforward and depends greatly on the origin and the composition of organic matter that is to be returned to the system. Wastes, which are important sources of organic carbon for soils, are taken as an example. For these organic materials the following factors have to be considered: the presence or absence of fossil C, the potential of direct and indirect emissions of non-CO2 greenhouse gases (CH4 and N2O) following application and the agro-system which is being used as a comparative reference.

This paper shows the geographic distribution in Germany of iron (Fe), nickel (Ni) and lead (Pb) analyzed in mosses in 1995/96 and compares it with the results of the 1990/91 pilot study within a European moss-monitoring programme. Other elements (As, Cd, Cr, Cu, Ti, V, Zn) are compared on basis of the overall element medians for Germany of the 1990/91 and 1995/96 survey. Samples of Pleurozium schreberi, Scleropodium purum, Hypnum cupressiforme and Hylocomium splendens were taken at a total of 1026 sites. In the 1995/96 monitoring campaign, 95% of the original sites of the 1990/91 study were resampled. The results from 1995/96 display local elevated values and many cases of areas affected by known sources of heavy-metal emissions. The industrialized and urban regions of Germany are shown up clearly by the 1995/96 moss-monitoring results: the Ruhr area, parts of Saarland and Baden-Württemberg, as well as areas in eastern Germany. Relatively low values for many elements were found in large areas of Lower Saxony and Bavaria. A comparison of the results of the 1990/91 and 1995/96 moss-monitoring programmes shows a fall in the concentration of the presented elements (except cadmium, copper and zinc) over the relevant period. Especially in the former GDR, chromium (Cr), iron (Fe), titanium (Ti) and vanadium (V) decreased significantly. This is, firstly, a reflection of the closure of and/or technological improvements to large power plants; secondly it is due to the fact that lignite has given way to other fuels. Vanadium (V) and nickel (Ni), typical constituents of crude oil, also show a decrease in the western part and thus document changes in the type of fuel consumed. The significant fall in lead concentration in 1995/96 as compared to 1990/91 in what used to be East and West Germany probably results from the increasing use of lead-free petrol. A comparison of the median values for 1990/91 and 1995/96 in mosses to the rate of emission of heavy metals in Germany for 1990 and 1995 shows similar trends in the case of elements such as arsenic (As), chromium (Cr), nickel (Ni), and lead (Pb). The comparison of the medians of the elements analyzed for 19 European countries indicates for most of the elements a general tendency to lower values in 1995, except for Lithuania, Netherlands, Portugal, Italy and United Kingdom.