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2. The Central Amazon Biomass Sink Under Current and Future Atmospheric CO2: Predictions From Big-Leaf and Demographic Vegetation Models

Author

Holm, JA, Knox, RG, Zhu, Q, Fisher, RA, Koven, CD, Nogueira Lima, AJ, Riley, WJ, Longo, M, Negrón-Juárez, RI, de Araujo, AC, Kueppers, LM, Moorcroft, PR, Higuchi, N, and Chambers, JQ

Subjects
biomass storage, plant mortality, Brazil, carbon allocation, plant growth, climate change, Geophysics
Abstract

There is large uncertainty whether Amazon forests will remain a carbon sink as atmospheric CO2 increases. Hence, we simulated an old-growth tropical forest using six versions of four terrestrial models differing in scale of vegetation structure and representation of biogeochemical (BGC) cycling, all driven with CO2 forcing from the preindustrial period to 2100. The models were benchmarked against tree inventory and eddy covariance data from a Brazilian site for present-day predictions. All models predicted positive vegetation growth that outpaced mortality, leading to continual increases in present-day biomass accumulation. Notably, the two vegetation demographic models (VDMs) (ED2 and ELM-FATES) always predicted positive stem diameter growth in all size classes. The field data, however, indicated that a quarter of canopy trees didn't grow over the 15-year period, and while high interannual variation existed, biomass change was near neutral. With a doubling of CO2, three of the four models predicted an appreciable biomass sink (0.77 to 1.24 Mg ha−1 year−1). ELMv1-ECA, the only model used here that includes phosphorus constraints, predicted the lowest biomass sink relative to initial biomass stocks (+21%), lower than the other BGC model, CLM5 (+48%). Models projections differed primarily through variations in nutrient constraints, then carbon allocation, initial biomass, and density-dependent mortality. The VDM's performance was similar or better than the BGC models run in carbon-only mode, suggesting that nutrient competition in VDMs will improve predictions. We demonstrate that VDMs are comparable to nondemographic (i.e., “big-leaf”) models but also include finer scale demography and competition that can be evaluated against field observations.

Published
2020

3. Calibration, measurement, and characterization of soil moisture dynamics in a central Amazonian tropical forest

Author

Negrón-Juárez, R, Ferreira, SJF, Mota, MC, Faybishenko, B, Monteiro, MTF, Candido, LA, Ribeiro, RP, de Oliveira, RC, de Araujo, AC, Warren, JM, Newman, BD, Gimenez, BO, Varadharajan, C, Agarwal, D, Borma, L, Tomasella, J, Higuchi, N, and Chambers, JQ

Subjects
Environmental Engineering, Physical Geography and Environmental Geoscience, Soil Sciences, Crop and Pasture Production
Abstract

Soil moisture plays a key role in hydrological, biogeochemical, and energy budgets of terrestrial ecosystems. Accurate soil moisture measurements in remote ecosystems such as the Amazon are difficult and limited because of logistical constraints. Time domain reflectometry (TDR) sensors are widely used to monitor soil moisture and require calibration to convert the TDR's dielectric permittivity measurement (Ka) to volumetric water content (θv). In this study, our objectives were to develop a field-based calibration of TDR sensors in an old-growth upland forest in the central Amazon, to evaluate the performance of the calibration, and then to apply the calibration to determine the dynamics of soil moisture content within a 14.2-m-deep vertical soil profile. Depth-specific TDR calibration using local soils in a controlled laboratory setting yielded a novel Ka–θv third-degree polynomial calibration. The sensors were later installed to their specific calibration depth in a 14.2-m pit. The widely used Ka–θv relationship (Topp model) underestimated the site-specific θv by 22–42%, indicating significant error in the model when applied to these well-structured, clay-rich tropical forest soils. The calibrated wet- and dry-season θv data showed a variety of depth and temporal variations highlighting the importance of soil textural differentiation, root uptake depths, as well as event to seasonal precipitation effects. Data such as these are greatly needed for improving our understanding of ecohydrological processes within tropical forests and for improving models of these systems in the face of changing environmental conditions.

Published
2020

4. Recognizing Amazonian tree species in the field using bark tissues spectra

Author

Hadlich, HL, Durgante, FM, dos Santos, J, Higuchi, N, Chambers, JQ, and Vicentini, A

Subjects
Environmental Sciences, Biological Sciences, Agricultural and Veterinary Sciences, Forestry
Abstract

The identification of tree species in the field is often a subjective process and misidentifications cause many problems for forest management in the Amazon Forest. Near infrared spectra from dried leaves of herbarium specimens are able to distinguish species in tropical forests. However, toolsto improve species identification directly in the field are needed. In this study, we tested whether spectral reflectance of bark tissues (rhytidome and phloem) collected with a portable spectrometer in the field can be used for the discrimination of tree species. Spectral data was collected for 254 trees of 8 families, 10 genera and 11 species from terra firme forests in Central Amazon with an ASD field spectrometer. Data consisted of reflectance values within 350–2500 nm wavelengths. We compared the rate of correct species recognition for different datasets using linear discriminant models. The rate of correct species assignment using this technique was 98% when using spectra from the inner bark (phloem) and 94% with outer bark (rhytidome) spectra. We suggest that the application of this technique can improve the quality of species identification directly during field inventories, fostering better forest management practices.

Published
2018

5. Revealing the causes and temporal distribution of tree mortality in Central Amazonia

Author

Fontes, CG, Chambers, JQ, and Higuchi, N

Subjects
Environmental Sciences, Biological Sciences, Agricultural and Veterinary Sciences, Forestry
Abstract

Tree mortality is a critical process in forest ecosystems, as it influences floristic composition, structure, dynamics, carbon storage, and forest nutrient cycling. However, the mechanisms behind tree death in tropical regions are still poorly characterized. This lack of information is mainly because tree mortality data come from long-interval inventories and studies that measured tree death seasonally are scarce. Here we offer novel insights into the power of fine temporal scale observations and we use a natural history approach to understand the processes and mechanism of tree mortality. We monitored tree mortality every month during one year, in 10 ha of terra-firme forest. To determine the pathways of mortality, we considered the state of the tree at the start of the investigation and the pre and post-mortem characteristics. From November 2010 to October 2011, 67 out of 5808 trees died. Despite the 2010 drought, mortality was highly correlated with monthly rainfall (r = 0.85). In total, six pathways of mortality were assessed. Storms were the main cause of mortality, killing 45% of all dead trees, followed by Biotic/abiotic factors accounting for 30% of tree mortality. The high mortality registered in the rainy season was mostly (78%) due to healthy trees dying uprooted or snapped. Finally, we would benefit from studies that assess mortality on a monthly basis and in combination with quantitative long-term data, we can substantially improve our understanding of the mechanisms behind tree death in the tropics.

Published
2018

6. Vulnerability of Amazon forests to storm-driven tree mortality

Author

Negrón-Juárez, RI, Holm, JA, Marra, DM, Rifai, SW, Riley, WJ, Chambers, JQ, Koven, CD, Knox, RG, McGroddy, ME, Di Vittorio, AV, Urquiza-Muñoz, J, Tello-Espinoza, R, Muñoz, WA, Ribeiro, GHPM, and Higuchi, N

Subjects
severe convective systems, winds, demography model, Meteorology & Atmospheric Sciences
Abstract

Tree mortality is a key driver of forest community composition and carbon dynamics. Strong winds associated with severe convective storms are dominant natural drivers of tree mortality in the Amazon. Why forests vary with respect to their vulnerability to wind events and how the predicted increase in storm events might affect forest ecosystems within the Amazon are not well understood. We found that windthrows are common in the Amazon region extending from northwest (Peru, Colombia, Venezuela, and west Brazil) to central Brazil, with the highest occurrence of windthrows in the northwest Amazon. More frequent winds, produced by more frequent severe convective systems, in combination with well-known processes that limit the anchoring of trees in the soil, help to explain the higher vulnerability of the northwest Amazon forests to winds. Projected increases in the frequency and intensity of convective storms in the Amazon have the potential to increase wind-related tree mortality. A forest demographic model calibrated for the northwestern and the central Amazon showed that northwestern forests are more resilient to increased wind-related tree mortality than forests in the central Amazon. Our study emphasizes the importance of including wind-related tree mortality in model simulations for reliable predictions of the future of tropical forests and their effects on the Earth' system.

Published
2018

7. Demonstration of a strict molecular oxygen requirement of yellow latex oxidation in the central Amazon canopy tree muiratinga (Maquira sclerophylla (Ducke) C.C. Berg)

Author

Piva, LRO, Jardine, KJ, Cobello, LO, Gimenez, BO, Durgante, FM, Higuchi, N, and Chambers, JQ

Abstract

Plant-derived latex is widely used in rubber production and plays important roles in ecological processes in the tropics. Although it is known that latex oxidation from the commercially important tree Hevea brasiliensis, results in latex browning, little is known about latex oxidation in highly diverse tropical ecosystems. Here we show that upon physical trunk damage, yellow latex released from the canopy tree Muiratinga (Maquira sclerophylla (Ducke) C.C. Berg) is rapidly and extensively oxidized to a black resin in the presence of air within 15-30 min. In a nitrogen atmosphere, latex oxidation was inhibited, but was immediately activated upon exposure to air. The results suggest the occurrence of O2-dependent oxidative enzymes including polyphenol oxidase (PPO) within the latex of Muiratinga and supports previous findings of a key role of oxidation during latex coagulation.

Published
2018

9. Windthrow variability in central Amazonia

Author

Negrón-Juárez, RI, Jenkins, HS, Raupp, CFM, Riley, WJ, Kueppers, LM, Marra, DM, Ribeiro, GHPM, Monteiro, MTF, Candido, LA, Chambers, JQ, and Higuchi, N

Subjects
windthrows, deep convection, squall lines, Central Amazonia, Atmospheric Sciences, Environmental Science and Management
Abstract

Windthrows are a recurrent disturbance in Amazonia and are an important driver of forest dynamics and carbon storage. In this study, we present for the first time the seasonal and interannual variability of windthrows, focusing on Central Amazonia, and discuss the potential meteorological factors associated with this variability. Landsat images over the 1998-2010 time period were used to detect the occurrence of windthrows, which were identified based on their spectral characteristics and shape. Here, we found that windthrows occurred every year but were more frequent between September and February. Organized convective activity associated with multicell storms embedded in mesoscale convective systems, such as northerly squall lines (that move from northeast to southwest) and southerly squall lines (that move from southwest to northeast) can cause windthrows. We also found that southerly squall lines occurred more frequently than their previously reported ~50 year interval. At the interannual scale, we did not find an association between El Niño-Southern Oscillation (ENSO) and windthrows.

Published
2017

10. Mechanical vulnerability and resistance to snapping and uprooting for Central Amazon tree species

Author

Ribeiro, GHPM, Chambers, JQ, Peterson, CJ, Trumbore, SE, Magnabosco Marra, D, Wirth, C, Cannon, JB, Négron-Juárez, RI, Lima, AJN, de Paula, EVCM, Santos, J, and Higuchi, N

Subjects
Tree static winching, Critical turning moment, Tree allometry, Functional traits, Blowdown, Wind-disturbance, Forestry, Environmental Sciences, Biological Sciences, Agricultural and Veterinary Sciences
Abstract

High descending winds generated by convective storms are a frequent and a major source of tree mortality disturbance events in the Amazon, affecting forest structure and diversity across a variety of scales, and more frequently observed in western and central portions of the basin. Soil texture in the Central Amazon also varies significantly with elevation along a topographic gradient, with decreasing clay content on plateaus, slopes and valleys respectively. In this study we investigated the critical turning moments (Mcrit - rotational force at the moment of tree failure, an indicator of tree stability or wind resistance) of 60 trees, ranging from 19.0 to 41.1 cm in diameter at breast height (DBH) and located in different topographic positions, and for different species, using a cable-winch load-cell system. Our approach used torque as a measure of tree failure to the point of snapping or uprooting. This approach provides a better understanding of the mechanical forces required to topple trees in tropical forests, and will inform models of wind throw disturbance. Across the topographic positions, size controlled variation in Mcrit was quantified for cardeiro (Scleronema mincranthum (Ducke) Ducke), mata-matá (Eschweilera spp.), and a random selection of trees from 19 other species. Our analysis of Mcrit revealed that tree resistance to failure increased with size (DBH and ABG) and differed among species. No effects of topography or failure mode were found for the species either separately or pooled. For the random species, total variance in Mcrit explained by tree size metrics increased from an R2 of 0.49 for DBH alone, to 0.68 when both DBH and stem fresh wood density (SWD) were included in a multiple regression model. This mechanistic approach allows the comparison of tree vulnerability induced by wind damage across ecosystems, and facilitates the use of forest structural information in ecosystem models that include variable resistance of trees to mortality inducing factors. Our results indicate that observed topographic differences in windthrow vulnerability are likely due to elevational differences in wind velocities, rather than by differences in soil-related factors that might effect Mcrit.

Published
2016

11. Diurnal pattern of leaf, flower and fruit specific ambient volatiles above an Oil Palm Plantation in Pará State, Brazil

Author

Jardine, KJ, Gimenez, BO, Araüjo, AC, Cunha, RL, Felizzola, JF, Piva, LR, Chambers, JQ, and Higuchi, N

Subjects
bioactive compounds, biomarkers, chemical ecology, essential oils, environmental analysis/quality, mass spectrometry, Analytical Chemistry, Chemical Sciences
Abstract

Oil palm plantations are rapidly expanding in the tropics because of insatiable global demand for fruit oil to be used in food, biofuels and cosmetics. Here we show that three tissue-specific volatiles can be quantified in ambient air above an African-American hybrid oil palm plantation in Brazil and linked photosynthesis (isoprene), floral scent (estragole), and for the first time, fruit oil processing (6-methyl-5-hepten-2-one, MHO). Plant enclosure techniques verified their tissue specific emission sources with ambient concentrations displaying distinct diurnal patterns above the canopy. Isoprene concentrations were near zero at night, but dramatically increased during the day while estragole showed elevated concentrations at night suggesting a light-independent, temperature-driven emission pattern from flowers. MHO also showed elevated concentrations at night and both estragole and MHO increased during the day. Our observations demonstrate that the African-American oil palm hybrid is strong isoprene emitter and suggest that MHO is a specific oxidation product of lycopene released during the industrial processing of palm oil. This study highlights the potential value of quantifying volatile oil palm signals in the atmosphere as a novel, non-invasive method to better understand biological functioning and its interactions with the environment including carbon assimilation, floral-insect interactions, and fruit oil production/processing.

Published
2016

12. Methanol and isoprene emissions from the fast growing tropical pioneer species Vismia guianensis (Aubl.) Pers. (Hypericaceae) in the central Amazon forest

Author

Jardine, KJ, Jardine, AB, Souza, VF, Carneiro, V, Ceron, JV, Gimenez, BO, Soares, CP, Durgante, FM, Higuchi, N, Manzi, AO, Gonçalves, JFC, Garcia, S, Martin, ST, Zorzanelli, RF, Piva, LR, and Chambers, JQ

Subjects
Meteorology & Atmospheric Sciences, Atmospheric Sciences, Astronomical and Space Sciences
Abstract

Isoprene (Is) emissions by plants represent a loss of carbon and energy resources leading to the initial hypothesis that fast growing pioneer species in secondary tropical forests allocate carbon primarily to growth at the expense of isoprenoid defenses. In this study, we quantified leaf isoprene and methanol emissions from the abundant pantropical pioneer tree species Vismia guianensis and ambient isoprene concentrations above a diverse secondary forest in the central Amazon. As photosynthetically active radiation (PAR) was varied (0 to 3000μ2s-1) under standard leaf temperature (30°C), isoprene emissions from V. guianensis increased without saturation up to 80n2s-1. A nonlinear increase in isoprene emissions with respect to net photosynthesis (Pn) resulted in the fraction of Pn dedicated to isoprene emissions increasing with light intensity (up to 2% of Pn). Emission responses to temperature under standard light conditions (PAR of 1000μ2s-1) resulted in the classic uncoupling of isoprene emissions (Topt, iso>40°C) from net photosynthesis (Topt, Pn Combining double low line 30.0-32.5°C) with up to 7% of Pn emitted as isoprene at 40°C. Under standard environmental conditions of PAR and leaf temperature, young V. guianensis leaves showed high methanol emissions, low Pn, and low isoprene emissions. In contrast, mature leaves showed high Pn, high isoprene emissions, and low methanol emissions, highlighting the differential control of leaf phenology over methanol and isoprene emissions. High daytime ambient isoprene concentrations (11ppbv) were observed above a secondary Amazon rainforest, suggesting that isoprene emissions are common among neotropical pioneer species. The results are not consistent with the initial hypothesis and support a functional role of methanol during leaf expansion and the establishment of photosynthetic machinery and a protective role of isoprene for photosynthesis during high temperature extremes regularly experienced in secondary rainforest ecosystems.

Published
2016

13. Predicting biomass of hyperdiverse and structurally complex central Amazonian forests - A virtual approach using extensive field data

Author

Magnabosco Marra, D, Higuchi, N, Trumbore, SE, Ribeiro, GHPM, Dos Santos, J, Carneiro, VMC, Lima, AJN, Chambers, JQ, Negrón-Juárez, RI, Holzwarth, F, Reu, B, and Wirth, C

Subjects
Meteorology & Atmospheric Sciences, Earth Sciences, Environmental Sciences, Biological Sciences
Abstract

Old-growth forests are subject to substantial changes in structure and species composition due to the intensification of human activities, gradual climate change and extreme weather events. Trees store ca. 90 % of the total aboveground biomass (AGB) in tropical forests and precise tree biomass estimation models are crucial for management and conservation. In the central Amazon, predicting AGB at large spatial scales is a challenging task due to the heterogeneity of successional stages, high tree species diversity and inherent variations in tree allometry and architecture. We parameterized generic AGB estimation models applicable across species and a wide range of structural and compositional variation related to species sorting into height layers as well as frequent natural disturbances. We used 727 trees (diameter at breast height ≥ 5 cm) from 101 genera and at least 135 species harvested in a contiguous forest near Manaus, Brazil. Sampling from this data set we assembled six scenarios designed to span existing gradients in floristic composition and size distribution in order to select models that best predict AGB at the landscape level across successional gradients. We found that good individual tree model fits do not necessarily translate into reliable predictions of AGB at the landscape level. When predicting AGB (dry mass) over scenarios using our different models and an available pantropical model, we observed systematic biases ranging from -31 % (pantropical) to +39 %, with root-mean-square error (RMSE) values of up to 130 Mg ha-1 (pantropical). Our first and second best models had both low mean biases (0.8 and 3.9 %, respectively) and RMSE (9.4 and 18.6 Mg ha-1) when applied over scenarios. Predicting biomass correctly at the landscape level in hyperdiverse and structurally complex tropical forests, especially allowing good performance at the margins of data availability for model construction/calibration, requires the inclusion of predictors that express inherent variations in species architecture. The model of interest should comprise the floristic composition and size-distribution variability of the target forest, implying that even generic global or pantropical biomass estimation models can lead to strong biases. Reliable biomass assessments for the Amazon basin (i.e., secondary forests) still depend on the collection of allometric data at the local/regional scale and forest inventories including species-specific attributes, which are often unavailable or estimated imprecisely in most regions.

Published
2016

14. Windthrows increase soil carbon stocks in a central Amazon forest

Author

Dos Santos, LT, Marra, DM, Trumbore, S, De Camargo, PB, Negrón-Juárez, RI, Lima, AJN, Ribeiro, GHPM, Dos Santos, J, and Higuchi, N

Subjects
Meteorology & Atmospheric Sciences, Earth Sciences, Environmental Sciences, Biological Sciences
Abstract

Windthrows change forest structure and species composition in central Amazon forests. However, the effects of widespread tree mortality associated with wind disturbances on soil properties have not yet been described in this vast region. We investigated short-term effects (7 years after disturbance) of widespread tree mortality caused by a squall line event from mid-January of 2005 on soil carbon stocks and concentrations in a central Amazon terra firme forest. The soil carbon stock (averaged over a 0-30 cm depth profile) in disturbed plots (61.4 ± 8.2 Mg ha-1, mean ±95 % confidence interval) was marginally higher (p = 0.09) than that from undisturbed plots (47.7 ± 13.6 Mg h-1). The soil organic carbon concentration in disturbed plots (2.0 ± 0.17 %) was significantly higher (p < 0.001) than that from undisturbed plots (1.36 ± 0.24 %). Moreover, soil carbon stocks were positively correlated with soil clay content (r2 = 0.332, r = 0.575 and p = 0.019) and with tree mortality intensity (r2 = 0.257, r = 0.506 and p = 0.045). Our results indicate that large inputs of plant litter associated with large windthrow events cause a short-term increase in soil carbon content, and the degree of increase is related to soil clay content and tree mortality intensity. The higher carbon content and potentially higher nutrient availability in soils from areas recovering from windthrows may favor forest regrowth and increase vegetation resilience.

Published
2016

15. Highly reactive light‐dependent monoterpenes in the Amazon

Author

Jardine, AB, Jardine, KJ, Fuentes, JD, Martin, ST, Martins, G, Durgante, F, Carneiro, V, Higuchi, N, Manzi, AO, and Chambers, JQ

Subjects
Earth Sciences, Atmospheric Sciences, secondary organic aerosols, vertical forest structure, ozonolysis, atmospheric oxidation, light-dependent monoterpenes, tropical VOC emissions, Meteorology & Atmospheric Sciences
Abstract

Despite orders of magnitude difference in atmospheric reactivity and great diversity in biological functioning, little is known about monoterpene speciation in tropical forests. Here we report vertically resolved ambient air mixing ratios for 12 monoterpenes in a central Amazon rainforest including observations of the highly reactive cis-β-ocimene (160 ppt), trans-β-ocimene (79 ppt), and terpinolene (32 ppt) which accounted for an estimated 21% of total monoterpene composition yet 55% of the upper canopy monoterpene ozonolysis rate. All 12 monoterpenes showed a mixing ratio peak in the upper canopy, with three demonstrating subcanopy peaks in 7 of 11 profiles. Leaf level emissions of highly reactive monoterpenes accounted for up to 1.9% of photosynthesis confirming light-dependent emissions across several Amazon tree genera. These results suggest that highly reactive monoterpenes play important antioxidant roles during photosynthesis in plants and serve as near-canopy sources of secondary organic aerosol precursors through atmospheric photooxidation via ozonolysis.

Published
2015

16. Dimethyl sulfide in the Amazon rain forest

Author

Jardine, K, Yañez‐Serrano, AM, Williams, J, Kunert, N, Jardine, A, Taylor, T, Abrell, L, Artaxo, P, Guenther, A, Hewitt, CN, House, E, Florentino, AP, Manzi, A, Higuchi, N, Kesselmeier, J, Behrendt, T, Veres, PR, Derstroff, B, Fuentes, JD, Martin, ST, and Andreae, MO

Subjects
Earth Sciences, Atmospheric Sciences, Climate Action, dimethyl sulfide, DMS, Amazon, Geochemistry, Oceanography, Meteorology & Atmospheric Sciences, Geoinformatics, Climate change impacts and adaptation
Abstract

Surface-to-atmosphere emissions of dimethyl sulfide (DMS) may impact global climate through the formation of gaseous sulfuric acid, which can yield secondary sulfate aerosols and contribute to new particle formation. While oceans are generally considered the dominant sources of DMS, a shortage of ecosystem observations prevents an accurate analysis of terrestrial DMS sources. Using mass spectrometry, we quantified ambient DMS mixing ratios within and above a primary rainforest ecosystem in the central Amazon Basin in real-time (2010-2011) and at high vertical resolution (2013-2014). Elevated but highly variable DMS mixing ratios were observed within the canopy, showing clear evidence of a net ecosystem source to the atmosphere during both day and night in both the dry and wet seasons. Periods of high DMS mixing ratios lasting up to 8 h (up to 160 parts per trillion (ppt)) often occurred within the canopy and near the surface during many evenings and nights. Daytime gradients showed mixing ratios (up to 80 ppt) peaking near the top of the canopy as well as near the ground following a rain event. The spatial and temporal distribution of DMS suggests that ambient levels and their potential climatic impacts are dominated by local soil and plant emissions. A soil source was confirmed by measurements of DMS emission fluxes from Amazon soils as a function of temperature and soil moisture. Furthermore, light- and temperature-dependent DMS emissions were measured from seven tropical tree species. Our study has important implications for understanding terrestrial DMS sources and their role in coupled land-atmosphere climate feedbacks.

Published
2015

18. Forest response to increased disturbance in the central Amazon and comparison to western Amazonian forests

Author

Holm, JA, Chambers, JQ, Collins, WD, and Higuchi, N

Subjects
Meteorology & Atmospheric Sciences, Earth Sciences, Environmental Sciences, Biological Sciences
Abstract

Uncertainties surrounding vegetation response to increased disturbance rates associated with climate change remains a major global change issue for Amazonian forests. Additionally, turnover rates computed as the average of mortality and recruitment rates in the western Amazon basin are doubled when compared to the central Amazon, and notable gradients currently exist in specific wood density and aboveground biomass (AGB) between these two regions. This study investigates the extent to which the variation in disturbance regimes contributes to these regional gradients. To address this issue, we evaluated disturbance-recovery processes in a central Amazonian forest under two scenarios of increased disturbance rates using first ZELIG-TROP, a dynamic vegetation gap model which we calibrated using long-term inventory data, and second using the Community Land Model (CLM), a global land surface model that is part of the Community Earth System Model (CESM). Upon doubling the mortality rate in the central Amazon to mirror the natural disturbance regime in the western Amazon of ∼2% mortality, the two regions continued to differ in multiple forest processes. With the inclusion of elevated natural disturbances, at steady state, AGB significantly decreased by 41.9% with no significant difference between modeled AGB and empirical AGB from the western Amazon data sets (104 vs. 107 Mg C ha1, respectively). However, different processes were responsible for the reductions in AGB between the models and empirical data set. The empirical data set suggests that a decrease in wood density is a driver leading to the reduction in AGB. While decreased stand basal area was the driver of AGB loss in ZELIG-TROP, a forest attribute that does not significantly vary across the Amazon Basin. Further comparisons found that stem density, specific wood density, and basal area growth rates differed between the two Amazonian regions. Last, to help quantify the impacts of increased disturbances on the climate and earth system, we evaluated the fidelity of tree mortality and disturbance in CLM. Similar to ZELIG-TROP, CLM predicted a net carbon loss of 49.9%, with an insignificant effect on aboveground net primary productivity (ANPP). Decreased leaf area index (LAI) was the driver of AGB loss in CLM, another forest attribute that does not significantly vary across the Amazon Basin, and the temporal variability in carbon stock and fluxes was not replicated in CLM. Our results suggest that (1) the variability between regions cannot be entirely explained by the variability in disturbance regime, but rather potentially sensitive to intrinsic environmental factors; or (2) the models are not accurately simulating all tropical forest characteristics in response to increased disturbances.

Published
2014

19. The steady-state mosaic of disturbance and succession across an old-growth Central Amazon forest landscape

Author

Chambers, J. Q, Negron-Juarez, R. I, Marra, D. M, Di Vittorio, A., Tews, J., Roberts, D., Ribeiro, G. H. P. M, Trumbore, S. E, and Higuchi, N.

Subjects
carbon dioxide, article, biodiversity, biomass, community succession, controlled study, ecosystem, fertilization, field study, forest, gap dynamics, landscape, mortality, mosaicism, nonhuman, plots and curves, priority journal, probability, recycling, remote sensing, scoring system, sensitivity analysis, simulation, steady state, stochastic model, time perception, time series analysis, tree, trend study, tropical rain forest, Biomass, Brazil, Carbon Cycle, Computer Simulation, Ecosystem, Models, Biological, Rivers, Trees, Tropical Climate
Abstract

Old-growth forest ecosystems comprise a mosaic of patches in different successional stages, with the fraction of the landscape in any particular state relatively constant over large temporal and spatial scales. The size distribution and return frequency of disturbance events, and subsequent recovery processes, determine to a large extent the spatial scale over which this old-growth steady state develops. Here, we characterize this mosaic for a Central Amazon forest by integrating field plot data, remote sensing disturbance probability distribution functions, and individual-based simulation modeling. Results demonstrate that a steady state of patches of varying successional age occurs over a relatively large spatial scale, with important implications for detecting temporal trends on plots that sample a small fraction of the landscape. Long highly significant stochastic runs averaging 1.0 Mg biomass⋅ha−1⋅y−1 were often punctuated by episodic disturbance events, resulting in a sawtooth time series of hectare-scale tree biomass. To maximize the detection of temporal trends for this Central Amazon site (e.g., driven by CO2 fertilization), plots larger than 10 ha would provide the greatest sensitivity. A model-based analysis of fractional mortality across all gap sizes demonstrated that 9.1–16.9% of tree mortality was missing from plot-based approaches, underscoring the need to combine plot and remote-sensing methods for estimating net landscape carbon balance. Old-growth tropical forests can exhibit complex large-scale structure driven by disturbance and recovery cycles, with ecosystem and community attributes of hectare-scale plots exhibiting continuous dynamic departures from a steady-state condition.

Published
2013

20. Basin-wide variations in Amazon forest structure and function are mediated by both soils and climate

Author

Quesada, C. A, Phillips, O. L, Schwarz, M., Czimczik, C. I, Baker, T. R, Patiño, S., Fyllas, N. M, Hodnett, M. G, Herrera, R., Almeida, S., Alvarez Dávila, E., Arneth, A., Arroyo, L., Chao, K. J, Dezzeo, N., Erwin, T., di Fiore, A., Higuchi, N., Honorio Coronado, E., Jimenez, E. M, Killeen, T., Lezama, A. T, Lloyd, G., López-González, G., Luizão, F. J, Malhi, Y., Monteagudo, A., Neill, D. A, Núñez Vargas, P., Paiva, R., Peacock, J., Peñuela, M. C, Peña Cruz, A., Pitman, N., Priante Filho, N., Prieto, A., Ramírez, H., Rudas, A., Salomão, R., Santos, A. J. B, Schmerler, J., Silva, N., Silveira, M., Vásquez, R., Vieira, I., Terborgh, J., and Lloyd, J.

Subjects
tropical rain-forest, plant-growth responses, ecological field experiments, net primary productivity, wood specific-gravity, branch xylem density, stem water storage, long term plots, geographical ecology, use efficiency
Abstract

Forest structure and dynamics vary across the Amazon Basin in an east-west gradient coincident with variations in soil fertility and geology. This has resulted in the hypothesis that soil fertility may play an important role in explaining Basin-wide variations in forest biomass, growth and stem turnover rates. Soil samples were collected in a total of 59 different forest plots across the Amazon Basin and analysed for exchangeable cations, carbon, nitrogen and pH, with several phosphorus fractions of likely different plant availability also quantified. Physical properties were additionally examined and an index of soil physical quality developed. Bivariate relationships of soil and climatic properties with above-ground wood productivity, stand-level tree turnover rates, above-ground wood biomass and wood density were first examined with multivariate regression models then applied. Both forms of analysis were undertaken with and without considerations regarding the underlying spatial structure of the dataset. Despite the presence of autocorrelated spatial structures complicating many analyses, forest structure and dynamics were found to be strongly and quantitatively related to edaphic as well as climatic conditions. Basin-wide differences in stand-level turnover rates are mostly influenced by soil physical properties with variations in rates of coarse wood production mostly related to soil phosphorus status. Total soil P was a better predictor of wood production rates than any of the fractionated organic- or inorganic-P pools. This suggests that it is not only the immediately available P forms, but probably the entire soil phosphorus pool that is interacting with forest growth on longer timescales. A role for soil potassium in modulating Amazon forest dynamics through its effects on stand-level wood density was also detected. Taking this into account, otherwise enigmatic variations in stand-level biomass across the Basin were then accounted for through the interacting effects of soil physical and chemical properties with climate. A hypothesis of self-maintaining forest dynamic feedback mechanisms initiated by edaphic conditions is proposed. It is further suggested that this is a major factor determining endogenous disturbance levels, species composition, and forest productivity across the Amazon Basin.

Published
2012

21. Recovery of above-ground tree biomass after moderate selective logging in a central Amazonian forest

Author

Otani T, Lima AJN, Suwa R, Amaral MRM, Ohashi S, Pinto ACM, Dos Santos J, Kajimoto T, Higuchi N, and Ishizuka M

Subjects
Annual Increment, Dynamics, Logistic Growth, Recovery Period, Terra Firme Forest, Forestry, SD1-669.5
Abstract

We examined the recovery and dynamics of living tree above-ground biomass (AGB) after selective logging in an Amazonian terra firme forest managed by a private company. The forest consisted of 24 blocks (including one set aside for conservation) selectively logged in different years on a managed schedule. Trees ≥10 cm in diameter at breast height (dbh) were surveyed in 2006 in 192 0.25-ha plots, in 2010 in 119 plots, and in 2012-2013 in 54 plots. A logistic growth model factoring in logging dynamics and mean AGB of a block in these years was established. Referencing the mean AGB of the unlogged forest, the model indicated that the logged forest would take on average 14 years to regain its preharvest AGB after selective logging at 1.9 trees ha-1 (dbh > 50 cm). In 2010 and 2012-2013, the AGB increased significantly for small and large trees (10-20 cm and >60 cm dbh, respectively) in the logged forest. In contrast, it decreased significantly for medium-sized trees (30-50 cm dbh) in the unlogged forest. Comparisons with the previous studies mainly conducted in the other regions of Amazon suggested that the estimated AGB recovery period with moderate logging intensity was almost appropriate and likely acceptable to forest managers.

Published
2018
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23. Pattern and Process in Amazon Tree Turnover, 1976-2001

Author

Phillips, O. L., Baker, T. R., Arroyo, L., Higuchi, N., Killeen, T. J., Laurance, W. F., Lewis, S. L., Lloyd, J., Malhi, Y., Monteagudo, A., Vargas, P. Núñez, Terborgh, J., Martínez, R. Vásquez, Almeida, S., Brown, S., Comiskey, J. A., Patiño, S., Quesada, C. A., and Vinceti, B.

Published
2004

25. Branch xylem density variations across the Amazon Basin

Author

Patiño, S., Lloyd, J., Paiva, R., Baker, T. R, Quesada, C. A, Mercado, L. M, Schmerler, J., Schwarz, M., Santos, A. J. B, Aguilar, A., Czimczik, C. I, Gallo, J., Horna, V., Hoyos, E. J, Jimenez, E. M, Palomino, W., Peacock, J., Peña-Cruz, A., Sarmiento, C., Sota, A., Turriago, J. D, Villanueva, B., Vitzthum, P., Alvarez, E., Arroyo, L., Baraloto, C., Bonal, D., Chave, J., Costa, A. C. L, Herrera, R., Higuchi, N., Killeen, T., Leal, E., Luizão, F., Meir, P., Monteagudo, A., Neil, D., Núñez-Vargas, P., Peñuela, M. C, Pitman, N., Priante Filho, N., Prieto, A., Panfil, S. N, Rudas, A., Salomão, R., Silva, N., Silveira, M., Soares deAlmeida, S., Torres-Lezama, A., Vásquez-Martínez, R., Vieira, I., Malhi, Y., and Phillips, O. L

Subjects
wood specific-gravity, tropical rain-forest, long-term plots, hydraulic architecture, cavitation resistance, carbon gain, functional-significance, water transport, savanna trees, canopy trees
Abstract

Xylem density is a physical property of wood that varies between individuals, species and environments. It reflects the physiological strategies of trees that lead to growth, survival and reproduction. Measurements of branch xylem density, ρx, were made for 1653 trees representing 598 species, sampled from 87 sites across the Amazon basin. Measured values ranged from 218 kg m−3for a Cordia sagotii (Boraginaceae) from Mountagne de Tortue, French Guiana to 1130 kg m−3 for an Aiouea sp. (Lauraceae) from Caxiuana, Central Pará, Brazil. Analysis of variance showed significant differences in average ρx across regions and sampled plots as well as significant differences between families, genera and species. A partitioning of the total variance in the dataset showed that species identity (family, genera and species) accounted for 33% with environment (geographic location and plot) accounting for an additional 26%; the remaining "residual" variance accounted for 41% of the total variance. Variations in plot means, were, however, not only accountable by differences in species composition because xylem density of the most widely distributed species in our dataset varied systematically from plot to plot. Thus, as well as having a genetic component, branch xylem density is a plastic trait that, for any given species, varies according to where the tree is growing in a predictable manner. Within the analysed taxa, exceptions to this general rule seem to be pioneer species belonging for example to the Urticaceae whose branch xylem density is more constrained than most species sampled in this study. These patterns of variation of branch xylem density across Amazonia suggest a large functional diversity amongst Amazonian trees which is not well understood.

Published
2009

26. An International Network to Monitor the Structure, Composition and Dynamics of Amazonian Forests (RAINFOR)

Author

Malhi, Y., Phillips, O. L., Lloyd, J., Baker, T., Wright, J., Almeida, S., Arroyo, L., Frederiksen, T., Grace, J., Higuchi, N., Killeen, T., Laurance, W. F., Leaño, C., Lewis, S., Meir, P., Monteagudo, A., Neill, D., Vargas, P. Núñez, Panfil, S. N., Patiño, S., Pitman, N., Quesada, C. A., Rudas-Ll., A., Salomão, R., Saleska, S., Silva, N., Silveira, M., Sombroek, W. G., Valencia, R., Martínez, R. Vásquez, Vieira, I. C. G., and Vinceti, B.

Published
2002

28. Slow growth rates of Amazonian trees: Consequences for carbon cycling

Author

Vieira, S., Trumbore, S., Camargo, P. B, Selhorst, D., Chambers, J. Q, Higuchi, N., and Martinelli, L. A

Subjects
carbon 14, rain, article, biomass, Brazil, carbon cycling, carbon sequestration, forest, growth rate, priority journal, season, species composition, tree growth, Biomass, Brazil, Carbon, Time Factors, Trees, Tropical Climate
Abstract

Quantifying age structure and tree growth rate of Amazonian forests is essential for understanding their role in the carbon cycle. Here, we use radiocarbon dating and direct measurement of diameter increment to document unexpectedly slow growth rates for trees from three locations spanning the Brazilian Amazon basin. Central Amazon trees, averaging only ≈1mm/year diameter increment, grow half as fast as those from areas with more seasonal rainfall to the east and west. Slow growth rates mean that trees can attain great ages; across our sites we estimate 17-50% of trees with diameter >10 cm have ages exceeding 300 years. Whereas a few emergent trees that make up a large portion of the biomass grow faster, small trees that are more abundant grow slowly and attain ages of hundreds of years. The mean age of carbon in living trees (60-110 years) is within the range of or slightly longer than the mean residence time calculated from C inventory divided by annual C allocation to wood growth (40-100 years). Faster C turnover is observed in stands with overall higher rates of diameter increment and a larger fraction of the biomass in large, fast-growing trees. As a consequence, forests can recover biomass relatively quickly after disturbance, whereas recovering species composition may take many centuries. Carbon cycle models that apply a single turnover time for carbon in forest biomass do not account for variations in life strategy and therefore may overestimate the carbon sequestration potential of Amazon forests.

Published
2005

29. Respiration from a tropical forest ecosystem: Partitioning of sources and low carbon use efficiency

Author

Chambers, JQ, Tribuzy, ES, Toledo, LC, Crispim, BF, Higuchi, N, Dos Santos, J, Araújo, AC, Kruijt, B, Nobre, AD, and Trumbore, SE

Abstract

Understanding how tropical forest carbon balance will respond to global change requires knowledge of individual heterotrophic and autotrophic respiratory sources, together with factors that control respiratory variability. We measured leaf, live wood, and soil respiration, along with additional environmental factors over a 1-yr period in a Central Amazon terra firme forest. Scaling these fluxes to the ecosystem, and combining our data with results from other studies, we estimated an average total ecosystem respiration (R eco) of 7.8 μmol·m-2·s-1. Average estimates (per unit ground area) for leaf, wood, soil, total heterotrophic, and total autotrophic respiration were 2.6, 1.1, 3.2, 5.6, and 2.2 μmol·m-2·s-1, respectively. Comparing autotrophic respiration with net primary production (NPP) estimates indicated that only ∼30% of carbon assimilated in photosynthesis was used to construct new tissues, with the remaining 70% being respired back to the atmosphere as autotrophic respiration. This low ecosystem carbon use efficiency (CUE) differs considerably from the relatively constant CUE of ∼0.5 found for temperate forests. Our Reco estimate was comparable to the above-canopy flux (Fac) from eddy covariance during defined sustained high turbulence conditions (when presumably Fac = Reco) of 8.4 (95% CI = 7.5-9.4). Multiple regression analysis demonstrated that ∼50% of the nighttime variability in Fac was accounted for by friction velocity (u*, a measure of turbulence) variables. After accounting for u* variability, mean Fac varied significantly with seasonal and daily changes in precipitation. A seasonal increase in precipitation resulted in a decrease in Fac, similar to our soil respiration response to moisture. The effect of daily changes in precipitation was complex: precipitation after a dry period resulted in a large increase in Fac, whereas additional precipitation after a rainy period had little effect. This response was similar to that of surface litter (coarse and fine), where respiration is greatly reduced when moisture is limiting, but increases markedly and quickly saturates with an increase in moisture.

Published
2004

30. The above-ground coarse wood productivity of 104 Neotropical forest plots.

Author

Malhi, Y, Baker, TR, Phillips, OL, Almeida, S, Alvarez, E, Arroyo, L, Chave, J, Czimczik, CI, Di Fiore, A, and Higuchi, N

Subjects
Amazonia, carbon, coarse wood productivity, GPP, growth, NPP, soil fertility, tropical forests, Biological Sciences, Environmental Sciences, Ecology
Abstract

The net primary production of tropical forests and its partitioning between long-lived carbon pools (wood) and shorter-lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above-ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree-size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above-ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha-1a-1) across the Neotropical plots, with a mean value of 3.1 Mg C ha-a-1. There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above-ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation. © 2004 Blackwell Publishing Ltd.

Published
2004

31. Pattern and process in Amazon tree turnover, 19762001

Author

Phillips, OL, Baker, TR, Arroyo, L, Higuchi, N, Killeen, TJ, Laurance, WF, Lewis, SL, Lloyd, J, Malhi, Y, Monteagudo, A, Neill, DA, Vargas, P Nez, Silva, JNM, Terborgh, J, Martnez, R Vsquez, Alexiades, M, Almeida, S, Brown, S, Chave, J, Comiskey, JA, Czimczik, CI, Di Fiore, A, Erwin, T, Kuebler, C, Laurance, SG, Nascimento, HEM, Olivier, J, Palacios, W, Patio, S, Pitman, NCA, Quesada, CA, Saldias, M, Lezama, A Torres, and Vinceti, B

Subjects
Biological Sciences, Biodiversity, Biomass, Carbon, Environmental Monitoring, Geography, Longitudinal Studies, Mortality, Population Dynamics, Rain, Reproduction, Soil, South America, Trees, Tropical Climate, recruitment, mortality, tree turnover, dynamics, Amazonia, forest, Medical and Health Sciences, Evolutionary Biology, Biological sciences, Biomedical and clinical sciences
Abstract

Previous work has shown that tree turnover, tree biomass and large liana densities have increased in mature tropical forest plots in the late twentieth century. These results point to a concerted shift in forest ecological processes that may already be having significant impacts on terrestrial carbon stocks, fluxes and biodiversity. However, the findings have proved controversial, partly because a rather limited number of permanent plots have been monitored for rather short periods. The aim of this paper is to characterize regional-scale patterns of 'tree turnover' (the rate with which trees die and recruit into a population) by using improved datasets now available for Amazonia that span the past 25 years. Specifically, we assess whether concerted changes in turnover are occurring, and if so whether they are general throughout the Amazon or restricted to one region or environmental zone. In addition, we ask whether they are driven by changes in recruitment, mortality or both. We find that: (i) trees 10 cm or more in diameter recruit and die twice as fast on the richer soils of southern and western Amazonia than on the poorer soils of eastern and central Amazonia; (ii) turnover rates have increased throughout Amazonia over the past two decades; (iii) mortality and recruitment rates have both increased significantly in every region and environmental zone, with the exception of mortality in eastern Amazonia; (iv) recruitment rates have consistently exceeded mortality rates; (v) absolute increases in recruitment and mortality rates are greatest in western Amazonian sites; and (vi) mortality appears to be lagging recruitment at regional scales. These spatial patterns and temporal trends are not caused by obvious artefacts in the data or the analyses. The trends cannot be directly driven by a mortality driver (such as increased drought or fragmentation-related death) because the biomass in these forests has simultaneously increased. Our findings therefore indicate that long-acting and widespread environmental changes are stimulating the growth and productivity of Amazon forests.

Published
2004

32. Carbon sink for a century.

Author

Chambers, JQ, Higuchi, N, Tribuzy, ES, and Trumbore, SE

Subjects
Trees, Carbon Dioxide, Carbon, Atmosphere, Tropical Climate, Models, Biological, Time, Wood, South America, Models, Biological, General Science & Technology
Published
2001

33. X Chromosome Contribution to the Genetic Architecture of Primary Biliary Cholangitis

Author

Asselta, R, Paraboschi, E, Gerussi, A, Cordell, H, Mells, G, Sandford, R, Jones, D, Nakamura, M, Ueno, K, Hitomi, Y, Kawashima, M, Nishida, N, Tokunaga, K, Nagasaki, M, Tanaka, A, Tang, R, Li, Z, Shi, Y, Liu, X, Xiong, M, Hirschfield, G, Siminovitch, K, Walker, E, Xie, G, Mason, A, Myers, R, Peltekian, K, Ghent, C, Atkinson, E, Juran, B, Lazaridis, K, Lu, Y, Gu, X, Jing, K, Amos, C, Affronti, A, Brunetto, M, Coco, B, Spinzi, G, Elia, G, Ferrari, C, Lleo, A, Muratori, L, Muratori, P, Portincasa, P, Colli, A, Bruno, S, Colloredo, G, Azzaroli, F, Andreone, P, Bragazzi, M, Alvaro, D, Cardinale, V, Cazzagon, N, Rigamonti, C, Floreani, A, Rosina, F, Ciaccio, A, Cristoferi, L, D'Amato, D, Malinverno, F, Mancuso, C, Massironi, S, Milani, C, O'Donnell, S, Ronca, V, Barisani, D, Lampertico, P, Donato, F, Fagiuoli, S, Almasio, P, Giannini, E, Cursaro, C, Colombo, M, Valenti, L, Miele, L, Andriulli, A, Niro, G, Grattagliano, I, Morini, L, Casella, G, Vinci, M, Battezzati, P, Crosignani, A, Zuin, M, Mattalia, A, Calvaruso, V, Colombo, S, Benedetti, A, Marzioni, M, Galli, A, Marra, F, Tarocchi, M, Picciotto, A, Morisco, F, Fabris, L, Croce, L, Tiribelli, C, Toniutto, P, Strazzabosco, M, Ch'Ng, C, Rahman, M, Yapp, T, Sturgess, R, Healey, C, Czajkowski, M, Gunasekera, A, Gyawali, P, Premchand, P, Kapur, K, Marley, R, Foster, G, Watson, A, Dias, A, Subhani, J, Harvey, R, Mccorry, R, Ramanaden, D, Gasem, J, Evans, R, Mathialahan, T, Shorrock, C, Lipscomb, G, Southern, P, Tibble, J, Gorard, D, Palegwala, A, Jones, S, Dawwas, M, Alexander, G, Dolwani, S, Prince, M, Foxton, M, Elphick, D, Mitchison, H, Gooding, I, Karmo, M, Saksena, S, Mendall, M, Patel, M, Ede, R, Austin, A, Sayer, J, Hankey, L, Hovell, C, Fisher, N, Carter, M, Koss, K, Piotrowicz, A, Grimley, C, Neal, D, Lim, G, Levi, S, Ala, A, Broad, A, Saeed, A, Wood, G, Brown, J, Wilkinson, M, Gordon, H, Ramage, J, Ridpath, J, Ngatchu, T, Grover, B, Shaukat, S, Shidrawi, R, Abouda, G, Ali, F, Rees, I, Salam, I, Narain, M, Brown, A, Taylor-Robinson, S, Williams, S, Grellier, L, Banim, P, Das, D, Chilton, A, Heneghan, M, Curtis, H, Gess, M, Drake, I, Aldersley, M, Davies, M, Jones, R, Mcnair, A, Srirajaskanthan, R, Pitcher, M, Sen, S, Bird, G, Barnardo, A, Kitchen, P, Yoong, K, Chirag, O, Sivaramakrishnan, N, Macfaul, G, Shah, A, Evans, C, Saha, S, Pollock, K, Bramley, P, Mukhopadhya, A, Fraser, A, Mills, P, Shallcross, C, Campbell, S, Bathgate, A, Shepherd, A, Dillon, J, Rushbrook, S, Przemioslo, R, Macdonald, C, Metcalf, J, Shmueli, U, Davis, A, Naqvi, A, Lee, T, Ryder, S, Collier, J, Klass, H, Ninkovic, M, Cramp, M, Sharer, N, Aspinall, R, Goggin, P, Ghosh, D, Douds, A, Hoeroldt, B, Booth, J, Williams, E, Hussaini, H, Stableforth, W, Ayres, R, Thorburn, D, Marshall, E, Burroughs, A, Mann, S, Lombard, M, Richardson, P, Patanwala, I, Maltby, J, Brookes, M, Mathew, R, Vyas, S, Singhal, S, Gleeson, D, Misra, S, Butterworth, J, George, K, Harding, T, Douglass, A, Panter, S, Shearman, J, Bray, G, Butcher, G, Forton, D, Mclindon, J, Cowan, M, Whatley, G, Mandal, A, Gupta, H, Sanghi, P, Jain, S, Pereira, S, Prasad, G, Watts, G, Wright, M, Neuberger, J, Gordon, F, Unitt, E, Grant, A, Delahooke, T, Higham, A, Brind, A, Cox, M, Ramakrishnan, S, King, A, Collins, C, Whalley, S, Li, A, Fraser, J, Bell, A, Wong, V, Singhal, A, Gee, I, Ang, Y, Ransford, R, Gotto, J, Millson, C, Bowles, J, Thomas, C, Harrison, M, Galaska, R, Kendall, J, Whiteman, J, Lawlor, C, Gray, C, Elliott, K, Mulvaney-Jones, C, Hobson, L, Van Duyvenvoorde, G, Loftus, A, Seward, K, Penn, R, Maiden, J, Damant, R, Hails, J, Cloudsdale, R, Silvestre, V, Glenn, S, Dungca, E, Wheatley, N, Doyle, H, Kent, M, Hamilton, C, Braim, D, Wooldridge, H, Abrahams, R, Paton, A, Lancaster, N, Gibbins, A, Hogben, K, Desousa, P, Muscariu, F, Musselwhite, J, Mckay, A, Tan, L, Foale, C, Brighton, J, Flahive, K, Nambela, E, Townshend, P, Ford, C, Holder, S, Palmer, C, Featherstone, J, Nasseri, M, Sadeghian, J, Williams, B, Rolls, S, Hynes, A, Duggan, C, Crossey, M, Stansfield, G, Macnicol, C, Wilkins, J, Wilhelmsen, E, Raymode, P, Lee, H, Durant, E, Bishop, R, Ncube, N, Tripoli, S, Casey, R, Cowley, C, Miller, R, Houghton, K, Ducker, S, Wright, F, Bird, B, Baxter, G, Keggans, J, Hughes, M, Grieve, E, Young, K, Williams, D, Ocker, K, Hines, F, Martin, K, Innes, C, Valliani, T, Fairlamb, H, Thornthwaite, S, Eastick, A, Tanqueray, E, Morrison, J, Holbrook, B, Browning, J, Walker, K, Congreave, S, Verheyden, J, Slininger, S, Stafford, L, O'Donnell, D, Ainsworth, M, Lord, S, Kent, L, March, L, Dickson, C, Simpson, D, Longhurst, B, Hayes, M, Shpuza, E, White, N, Besley, S, Pearson, S, Wright, A, Jones, L, Gunter, E, Dewhurst, H, Fouracres, A, Farrington, L, Graves, L, Marriott, S, Leoni, M, Tyrer, D, Dali-kemmery, L, Lambourne, V, Green, M, Sirdefield, D, Amor, K, Colley, J, Shinder, B, Jones, J, Mills, M, Carnahan, M, Taylor, N, Boulton, K, Tregonning, J, Brown, C, Clifford, G, Archer, E, Hamilton, M, Curtis, J, Shewan, T, Walsh, S, Warner, K, Netherton, K, Mupudzi, M, Gunson, B, Gitahi, J, Gocher, D, Batham, S, Pateman, H, Desmennu, S, Conder, J, Clement, D, Gallagher, S, Orpe, J, Chan, P, Currie, L, O'Donohoe, L, Oblak, M, Morgan, L, Quinn, M, Amey, I, Baird, Y, Cotterill, D, Cumlat, L, Winter, L, Greer, S, Spurdle, K, Allison, J, Dyer, S, Sweeting, H, Kordula, J, Aiba, Y, Nakamura, H, Abiru, S, Nagaoka, S, Komori, A, Yatsuhashi, H, Ishibashi, H, Ito, M, Kawai, Y, Kohn, S, Gervais, O, Migita, K, Katsushima, S, Naganuma, A, Sugi, K, Komatsu, T, Mannami, T, Matsushita, K, Yoshizawa, K, Makita, F, Nikami, T, Nishimura, H, Kouno, H, Ota, H, Komura, T, Nakamura, Y, Shimada, M, Hirashima, N, Komeda, T, Ario, K, Nakamuta, M, Yamashita, T, Furuta, K, Kikuchi, M, Naeshiro, N, Takahashi, H, Mano, Y, Tsunematsu, S, Yabuuchi, I, Shimada, Y, Yamauchi, K, Sugimoto, R, Sakai, H, Mita, E, Koda, M, Tsuruta, S, Kamitsukasa, H, Sato, T, Masaki, N, Kobata, T, Fukushima, N, Higuchi, N, Ohara, Y, Muro, T, Takesaki, E, Takaki, H, Yamamoto, T, Kato, M, Nagaoki, Y, Hayashi, S, Ishida, J, Watanabe, Y, Kobayashi, M, Koga, M, Saoshiro, T, Yagura, M, Hirata, K, Takikawa, H, Ohira, H, Zeniya, M, Abe, M, Onji, M, Kaneko, S, Honda, M, Arai, K, Arinaga-Hino, T, Hashimoto, E, Taniai, M, Umemura, T, Joshita, S, Nakao, K, Ichikawa, T, Shibata, H, Yamagiwa, S, Seike, M, Honda, K, Sakisaka, S, Takeyama, Y, Harada, M, Senju, M, Yokosuka, O, Kanda, T, Ueno, Y, Kikuchi, K, Ebinuma, H, Himoto, T, Yasunami, M, Murata, K, Mizokami, M, Shimoda, S, Miyake, Y, Takaki, A, Yamamoto, K, Hirano, K, Ichida, T, Ido, A, Tsubouchi, H, Chayama, K, Harada, K, Nakanuma, Y, Maehara, Y, Taketomi, A, Shirabe, K, Soejima, Y, Mori, A, Yagi, S, Uemoto, S, Tanaka, T, Yamashiki, N, Tamura, S, Sugawara, Y, Kokudo, N, Carbone, M, Cardamone, G, Duga, S, Gershwin, M, Seldin, M, Invernizzi, P, Asselta R., Paraboschi E. M., Gerussi A., Cordell H. J., Mells G. F., Sandford R. N., Jones D. E., Nakamura M., Ueno K., Hitomi Y., Kawashima M., Nishida N., Tokunaga K., Nagasaki M., Tanaka A., Tang R., Li Z., Shi Y., Liu X., Xiong M., Hirschfield G., Siminovitch K. A., Walker E., Xie G., Mason A., Myers R., Peltekian K., Ghent C., Atkinson E., Juran B., Lazaridis K., Lu Y., Gu X., Jing K., Amos C., Affronti A., Brunetto M., Coco B., Spinzi G., Elia G., Ferrari C., Lleo A., Muratori L., Muratori P., Portincasa P., Colli A., Bruno S., Colloredo G., Azzaroli F., Andreone P., Bragazzi M., Alvaro D., Cardinale V., Cazzagon N., Rigamonti C., Floreani A., Rosina F., Ciaccio A., Cristoferi L., D'Amato D., Malinverno F., Mancuso C., Massironi S., Milani C., O'Donnell S. E., Ronca V., Barisani D., Lampertico P., Donato F., Fagiuoli S., Almasio P. L., Giannini E., Cursaro C., Colombo M., Valenti L., Miele L., Andriulli A., Niro G. A., Grattagliano I., Morini L., Casella G., Vinci M., Battezzati P. M., Crosignani A., Zuin M., Mattalia A., Calvaruso V., Colombo S., Benedetti A., Marzioni M., Galli A., Marra F., Tarocchi M., Picciotto A., Morisco F., Fabris L., Croce L. S., Tiribelli C., Toniutto P., Strazzabosco M., Ch'ng C. L., Rahman M., Yapp T., Sturgess R., Healey C., Czajkowski M., Gunasekera A., Gyawali P., Premchand P., Kapur K., Marley R., Foster G., Watson A., Dias A., Subhani J., Harvey R., McCorry R., Ramanaden D., Gasem J., Evans R., Mathialahan T., Shorrock C., Lipscomb G., Southern P., Tibble J., Gorard D., Palegwala A., Jones S., Dawwas M., Alexander G., Dolwani S., Prince M., Foxton M., Elphick D., Mitchison H., Gooding I., Karmo M., Saksena S., Mendall M., Patel M., Ede R., Austin A., Sayer J., Hankey L., Hovell C., Fisher N., Carter M., Koss K., Piotrowicz A., Grimley C., Neal D., Lim G., Levi S., Ala A., Broad A., Saeed A., Wood G., Brown J., Wilkinson M., Gordon H., Ramage J., Ridpath J., Ngatchu T., Grover B., Shaukat S., Shidrawi R., Abouda G., Ali F., Rees I., Salam I., Narain M., Brown A., Taylor-Robinson S., Williams S., Grellier L., Banim P., Das D., Chilton A., Heneghan M., Curtis H., Gess M., Drake I., Aldersley M., Davies M., Jones R., McNair A., Srirajaskanthan R., Pitcher M., Sen S., Bird G., Barnardo A., Kitchen P., Yoong K., Chirag O., Sivaramakrishnan N., MacFaul G., Jones D., Shah A., Evans C., Saha S., Pollock K., Bramley P., Mukhopadhya A., Fraser A., Mills P., Shallcross C., Campbell S., Bathgate A., Shepherd A., Dillon J., Rushbrook S., Przemioslo R., Macdonald C., Metcalf J., Shmueli U., Davis A., Naqvi A., Lee T., Ryder S. D., Collier J., Klass H., Ninkovic M., Cramp M., Sharer N., Aspinall R., Goggin P., Ghosh D., Douds A., Hoeroldt B., Booth J., Williams E., Hussaini H., Stableforth W., Ayres R., Thorburn D., Marshall E., Burroughs A., Mann S., Lombard M., Richardson P., Patanwala I., Maltby J., Brookes M., Mathew R., Vyas S., Singhal S., Gleeson D., Misra S., Butterworth J., George K., Harding T., Douglass A., Panter S., Shearman J., Bray G., Butcher G., Forton D., Mclindon J., Cowan M., Whatley G., Mandal A., Gupta H., Sanghi P., Jain S., Pereira S., Prasad G., Watts G., Wright M., Neuberger J., Gordon F., Unitt E., Grant A., Delahooke T., Higham A., Brind A., Cox M., Ramakrishnan S., King A., Collins C., Whalley S., Li A., Fraser J., Bell A., Wong V. S., Singhal A., Gee I., Ang Y., Ransford R., Gotto J., Millson C., Bowles J., Thomas C., Harrison M., Galaska R., Kendall J., Whiteman J., Lawlor C., Gray C., Elliott K., Mulvaney-Jones C., Hobson L., Van Duyvenvoorde G., Loftus A., Seward K., Penn R., Maiden J., Damant R., Hails J., Cloudsdale R., Silvestre V., Glenn S., Dungca E., Wheatley N., Doyle H., Kent M., Hamilton C., Braim D., Wooldridge H., Abrahams R., Paton A., Lancaster N., Gibbins A., Hogben K., Desousa P., Muscariu F., Musselwhite J., McKay A., Tan L., Foale C., Brighton J., Flahive K., Nambela E., Townshend P., Ford C., Holder S., Palmer C., Featherstone J., Nasseri M., Sadeghian J., Williams B., Rolls S. -A., Hynes A., Duggan C., Crossey M., Stansfield G., MacNicol C., Wilkins J., Wilhelmsen E., Raymode P., Lee H. -J., Durant E., Bishop R., Ncube N., Tripoli S., Casey R., Cowley C., Miller R., Houghton K., Ducker S., Wright F., Bird B., Baxter G., Keggans J., Hughes M., Grieve E., Young K., Williams D., Ocker K., Hines F., Martin K., Innes C., Valliani T., Fairlamb H., Thornthwaite S., Eastick A., Tanqueray E., Morrison J., Holbrook B., Browning J., Walker K., Congreave S., Verheyden J., Slininger S., Stafford L., O'Donnell D., Ainsworth M., Lord S., Kent L., March L., Dickson C., Simpson D., Longhurst B., Hayes M., Shpuza E., White N., Besley S., Pearson S., Wright A., Jones L., Gunter E., Dewhurst H., Fouracres A., Farrington L., Graves L., Marriott S., Leoni M., Tyrer D., Dali-kemmery L., Lambourne V., Green M., Sirdefield D., Amor K., Colley J., Shinder B., Jones J., Mills M., Carnahan M., Taylor N., Boulton K., Tregonning J., Brown C., Clifford G., Archer E., Hamilton M., Curtis J., Shewan T., Walsh S., Warner K., Netherton K., Mupudzi M., Gunson B., Gitahi J., Gocher D., Batham S., Pateman H., Desmennu S., Conder J., Clement D., Gallagher S., Orpe J., Chan P., Currie L., O'Donohoe L., Oblak M., Morgan L., Quinn M., Amey I., Baird Y., Cotterill D., Cumlat L., Winter L., Greer S., Spurdle K., Allison J., Dyer S., Sweeting H., Kordula J., Aiba Y., Nakamura H., Abiru S., Nagaoka S., Komori A., Yatsuhashi H., Ishibashi H., Ito M., Kawai Y., Kohn S. -S., Gervais O., Migita K., Katsushima S., Naganuma A., Sugi K., Komatsu T., Mannami T., Matsushita K., Yoshizawa K., Makita F., Nikami T., Nishimura H., Kouno H., Ota H., Komura T., Nakamura Y., Shimada M., Hirashima N., Komeda T., Ario K., Nakamuta M., Yamashita T., Furuta K., Kikuchi M., Naeshiro N., Takahashi H., Mano Y., Tsunematsu S., Yabuuchi I., Shimada Y., Yamauchi K., Sugimoto R., Sakai H., Mita E., Koda M., Tsuruta S., Kamitsukasa H., Sato T., Masaki N., Kobata T., Fukushima N., Higuchi N., Ohara Y., Muro T., Takesaki E., Takaki H., Yamamoto T., Kato M., Nagaoki Y., Hayashi S., Ishida J., Watanabe Y., Kobayashi M., Koga M., Saoshiro T., Yagura M., Hirata K., Takikawa H., Ohira H., Zeniya M., Abe M., Onji M., Kaneko S., Honda M., Arai K., Arinaga-Hino T., Hashimoto E., Taniai M., Umemura T., Joshita S., Nakao K., Ichikawa T., Shibata H., Yamagiwa S., Seike M., Honda K., Sakisaka S., Takeyama Y., Harada M., Senju M., Yokosuka O., Kanda T., Ueno Y., Kikuchi K., Ebinuma H., Himoto T., Yasunami M., Murata K., Mizokami M., Shimoda S., Miyake Y., Takaki A., Yamamoto K., Hirano K., Ichida T., Ido A., Tsubouchi H., Chayama K., Harada K., Nakanuma Y., Maehara Y., Taketomi A., Shirabe K., Soejima Y., Mori A., Yagi S., Uemoto S., Tanaka T., Yamashiki N., Tamura S., Sugawara Y., Kokudo N., Carbone M., Cardamone G., Duga S., Gershwin M. E., Seldin M. F., Invernizzi P., Asselta, R, Paraboschi, E, Gerussi, A, Cordell, H, Mells, G, Sandford, R, Jones, D, Nakamura, M, Ueno, K, Hitomi, Y, Kawashima, M, Nishida, N, Tokunaga, K, Nagasaki, M, Tanaka, A, Tang, R, Li, Z, Shi, Y, Liu, X, Xiong, M, Hirschfield, G, Siminovitch, K, Walker, E, Xie, G, Mason, A, Myers, R, Peltekian, K, Ghent, C, Atkinson, E, Juran, B, Lazaridis, K, Lu, Y, Gu, X, Jing, K, Amos, C, Affronti, A, Brunetto, M, Coco, B, Spinzi, G, Elia, G, Ferrari, C, Lleo, A, Muratori, L, Muratori, P, Portincasa, P, Colli, A, Bruno, S, Colloredo, G, Azzaroli, F, Andreone, P, Bragazzi, M, Alvaro, D, Cardinale, V, Cazzagon, N, Rigamonti, C, Floreani, A, Rosina, F, Ciaccio, A, Cristoferi, L, D'Amato, D, Malinverno, F, Mancuso, C, Massironi, S, Milani, C, O'Donnell, S, Ronca, V, Barisani, D, Lampertico, P, Donato, F, Fagiuoli, S, Almasio, P, Giannini, E, Cursaro, C, Colombo, M, Valenti, L, Miele, L, Andriulli, A, Niro, G, Grattagliano, I, Morini, L, Casella, G, Vinci, M, Battezzati, P, Crosignani, A, Zuin, M, Mattalia, A, Calvaruso, V, Colombo, S, Benedetti, A, Marzioni, M, Galli, A, Marra, F, Tarocchi, M, Picciotto, A, Morisco, F, Fabris, L, Croce, L, Tiribelli, C, Toniutto, P, Strazzabosco, M, Ch'Ng, C, Rahman, M, Yapp, T, Sturgess, R, Healey, C, Czajkowski, M, Gunasekera, A, Gyawali, P, Premchand, P, Kapur, K, Marley, R, Foster, G, Watson, A, Dias, A, Subhani, J, Harvey, R, Mccorry, R, Ramanaden, D, Gasem, J, Evans, R, Mathialahan, T, Shorrock, C, Lipscomb, G, Southern, P, Tibble, J, Gorard, D, Palegwala, A, Jones, S, Dawwas, M, Alexander, G, Dolwani, S, Prince, M, Foxton, M, Elphick, D, Mitchison, H, Gooding, I, Karmo, M, Saksena, S, Mendall, M, Patel, M, Ede, R, Austin, A, Sayer, J, Hankey, L, Hovell, C, Fisher, N, Carter, M, Koss, K, Piotrowicz, A, Grimley, C, Neal, D, Lim, G, Levi, S, Ala, A, Broad, A, Saeed, A, Wood, G, Brown, J, Wilkinson, M, Gordon, H, Ramage, J, Ridpath, J, Ngatchu, T, Grover, B, Shaukat, S, Shidrawi, R, Abouda, G, Ali, F, Rees, I, Salam, I, Narain, M, Brown, A, Taylor-Robinson, S, Williams, S, Grellier, L, Banim, P, Das, D, Chilton, A, Heneghan, M, Curtis, H, Gess, M, Drake, I, Aldersley, M, Davies, M, Jones, R, Mcnair, A, Srirajaskanthan, R, Pitcher, M, Sen, S, Bird, G, Barnardo, A, Kitchen, P, Yoong, K, Chirag, O, Sivaramakrishnan, N, Macfaul, G, Shah, A, Evans, C, Saha, S, Pollock, K, Bramley, P, Mukhopadhya, A, Fraser, A, Mills, P, Shallcross, C, Campbell, S, Bathgate, A, Shepherd, A, Dillon, J, Rushbrook, S, Przemioslo, R, Macdonald, C, Metcalf, J, Shmueli, U, Davis, A, Naqvi, A, Lee, T, Ryder, S, Collier, J, Klass, H, Ninkovic, M, Cramp, M, Sharer, N, Aspinall, R, Goggin, P, Ghosh, D, Douds, A, Hoeroldt, B, Booth, J, Williams, E, Hussaini, H, Stableforth, W, Ayres, R, Thorburn, D, Marshall, E, Burroughs, A, Mann, S, Lombard, M, Richardson, P, Patanwala, I, Maltby, J, Brookes, M, Mathew, R, Vyas, S, Singhal, S, Gleeson, D, Misra, S, Butterworth, J, George, K, Harding, T, Douglass, A, Panter, S, Shearman, J, Bray, G, Butcher, G, Forton, D, Mclindon, J, Cowan, M, Whatley, G, Mandal, A, Gupta, H, Sanghi, P, Jain, S, Pereira, S, Prasad, G, Watts, G, Wright, M, Neuberger, J, Gordon, F, Unitt, E, Grant, A, Delahooke, T, Higham, A, Brind, A, Cox, M, Ramakrishnan, S, King, A, Collins, C, Whalley, S, Li, A, Fraser, J, Bell, A, Wong, V, Singhal, A, Gee, I, Ang, Y, Ransford, R, Gotto, J, Millson, C, Bowles, J, Thomas, C, Harrison, M, Galaska, R, Kendall, J, Whiteman, J, Lawlor, C, Gray, C, Elliott, K, Mulvaney-Jones, C, Hobson, L, Van Duyvenvoorde, G, Loftus, A, Seward, K, Penn, R, Maiden, J, Damant, R, Hails, J, Cloudsdale, R, Silvestre, V, Glenn, S, Dungca, E, Wheatley, N, Doyle, H, Kent, M, Hamilton, C, Braim, D, Wooldridge, H, Abrahams, R, Paton, A, Lancaster, N, Gibbins, A, Hogben, K, Desousa, P, Muscariu, F, Musselwhite, J, Mckay, A, Tan, L, Foale, C, Brighton, J, Flahive, K, Nambela, E, Townshend, P, Ford, C, Holder, S, Palmer, C, Featherstone, J, Nasseri, M, Sadeghian, J, Williams, B, Rolls, S, Hynes, A, Duggan, C, Crossey, M, Stansfield, G, Macnicol, C, Wilkins, J, Wilhelmsen, E, Raymode, P, Lee, H, Durant, E, Bishop, R, Ncube, N, Tripoli, S, Casey, R, Cowley, C, Miller, R, Houghton, K, Ducker, S, Wright, F, Bird, B, Baxter, G, Keggans, J, Hughes, M, Grieve, E, Young, K, Williams, D, Ocker, K, Hines, F, Martin, K, Innes, C, Valliani, T, Fairlamb, H, Thornthwaite, S, Eastick, A, Tanqueray, E, Morrison, J, Holbrook, B, Browning, J, Walker, K, Congreave, S, Verheyden, J, Slininger, S, Stafford, L, O'Donnell, D, Ainsworth, M, Lord, S, Kent, L, March, L, Dickson, C, Simpson, D, Longhurst, B, Hayes, M, Shpuza, E, White, N, Besley, S, Pearson, S, Wright, A, Jones, L, Gunter, E, Dewhurst, H, Fouracres, A, Farrington, L, Graves, L, Marriott, S, Leoni, M, Tyrer, D, Dali-kemmery, L, Lambourne, V, Green, M, Sirdefield, D, Amor, K, Colley, J, Shinder, B, Jones, J, Mills, M, Carnahan, M, Taylor, N, Boulton, K, Tregonning, J, Brown, C, Clifford, G, Archer, E, Hamilton, M, Curtis, J, Shewan, T, Walsh, S, Warner, K, Netherton, K, Mupudzi, M, Gunson, B, Gitahi, J, Gocher, D, Batham, S, Pateman, H, Desmennu, S, Conder, J, Clement, D, Gallagher, S, Orpe, J, Chan, P, Currie, L, O'Donohoe, L, Oblak, M, Morgan, L, Quinn, M, Amey, I, Baird, Y, Cotterill, D, Cumlat, L, Winter, L, Greer, S, Spurdle, K, Allison, J, Dyer, S, Sweeting, H, Kordula, J, Aiba, Y, Nakamura, H, Abiru, S, Nagaoka, S, Komori, A, Yatsuhashi, H, Ishibashi, H, Ito, M, Kawai, Y, Kohn, S, Gervais, O, Migita, K, Katsushima, S, Naganuma, A, Sugi, K, Komatsu, T, Mannami, T, Matsushita, K, Yoshizawa, K, Makita, F, Nikami, T, Nishimura, H, Kouno, H, Ota, H, Komura, T, Nakamura, Y, Shimada, M, Hirashima, N, Komeda, T, Ario, K, Nakamuta, M, Yamashita, T, Furuta, K, Kikuchi, M, Naeshiro, N, Takahashi, H, Mano, Y, Tsunematsu, S, Yabuuchi, I, Shimada, Y, Yamauchi, K, Sugimoto, R, Sakai, H, Mita, E, Koda, M, Tsuruta, S, Kamitsukasa, H, Sato, T, Masaki, N, Kobata, T, Fukushima, N, Higuchi, N, Ohara, Y, Muro, T, Takesaki, E, Takaki, H, Yamamoto, T, Kato, M, Nagaoki, Y, Hayashi, S, Ishida, J, Watanabe, Y, Kobayashi, M, Koga, M, Saoshiro, T, Yagura, M, Hirata, K, Takikawa, H, Ohira, H, Zeniya, M, Abe, M, Onji, M, Kaneko, S, Honda, M, Arai, K, Arinaga-Hino, T, Hashimoto, E, Taniai, M, Umemura, T, Joshita, S, Nakao, K, Ichikawa, T, Shibata, H, Yamagiwa, S, Seike, M, Honda, K, Sakisaka, S, Takeyama, Y, Harada, M, Senju, M, Yokosuka, O, Kanda, T, Ueno, Y, Kikuchi, K, Ebinuma, H, Himoto, T, Yasunami, M, Murata, K, Mizokami, M, Shimoda, S, Miyake, Y, Takaki, A, Yamamoto, K, Hirano, K, Ichida, T, Ido, A, Tsubouchi, H, Chayama, K, Harada, K, Nakanuma, Y, Maehara, Y, Taketomi, A, Shirabe, K, Soejima, Y, Mori, A, Yagi, S, Uemoto, S, Tanaka, T, Yamashiki, N, Tamura, S, Sugawara, Y, Kokudo, N, Carbone, M, Cardamone, G, Duga, S, Gershwin, M, Seldin, M, Invernizzi, P, Asselta R., Paraboschi E. M., Gerussi A., Cordell H. J., Mells G. F., Sandford R. N., Jones D. E., Nakamura M., Ueno K., Hitomi Y., Kawashima M., Nishida N., Tokunaga K., Nagasaki M., Tanaka A., Tang R., Li Z., Shi Y., Liu X., Xiong M., Hirschfield G., Siminovitch K. A., Walker E., Xie G., Mason A., Myers R., Peltekian K., Ghent C., Atkinson E., Juran B., Lazaridis K., Lu Y., Gu X., Jing K., Amos C., Affronti A., Brunetto M., Coco B., Spinzi G., Elia G., Ferrari C., Lleo A., Muratori L., Muratori P., Portincasa P., Colli A., Bruno S., Colloredo G., Azzaroli F., Andreone P., Bragazzi M., Alvaro D., Cardinale V., Cazzagon N., Rigamonti C., Floreani A., Rosina F., Ciaccio A., Cristoferi L., D'Amato D., Malinverno F., Mancuso C., Massironi S., Milani C., O'Donnell S. E., Ronca V., Barisani D., Lampertico P., Donato F., Fagiuoli S., Almasio P. L., Giannini E., Cursaro C., Colombo M., Valenti L., Miele L., Andriulli A., Niro G. A., Grattagliano I., Morini L., Casella G., Vinci M., Battezzati P. M., Crosignani A., Zuin M., Mattalia A., Calvaruso V., Colombo S., Benedetti A., Marzioni M., Galli A., Marra F., Tarocchi M., Picciotto A., Morisco F., Fabris L., Croce L. S., Tiribelli C., Toniutto P., Strazzabosco M., Ch'ng C. L., Rahman M., Yapp T., Sturgess R., Healey C., Czajkowski M., Gunasekera A., Gyawali P., Premchand P., Kapur K., Marley R., Foster G., Watson A., Dias A., Subhani J., Harvey R., McCorry R., Ramanaden D., Gasem J., Evans R., Mathialahan T., Shorrock C., Lipscomb G., Southern P., Tibble J., Gorard D., Palegwala A., Jones S., Dawwas M., Alexander G., Dolwani S., Prince M., Foxton M., Elphick D., Mitchison H., Gooding I., Karmo M., Saksena S., Mendall M., Patel M., Ede R., Austin A., Sayer J., Hankey L., Hovell C., Fisher N., Carter M., Koss K., Piotrowicz A., Grimley C., Neal D., Lim G., Levi S., Ala A., Broad A., Saeed A., Wood G., Brown J., Wilkinson M., Gordon H., Ramage J., Ridpath J., Ngatchu T., Grover B., Shaukat S., Shidrawi R., Abouda G., Ali F., Rees I., Salam I., Narain M., Brown A., Taylor-Robinson S., Williams S., Grellier L., Banim P., Das D., Chilton A., Heneghan M., Curtis H., Gess M., Drake I., Aldersley M., Davies M., Jones R., McNair A., Srirajaskanthan R., Pitcher M., Sen S., Bird G., Barnardo A., Kitchen P., Yoong K., Chirag O., Sivaramakrishnan N., MacFaul G., Jones D., Shah A., Evans C., Saha S., Pollock K., Bramley P., Mukhopadhya A., Fraser A., Mills P., Shallcross C., Campbell S., Bathgate A., Shepherd A., Dillon J., Rushbrook S., Przemioslo R., Macdonald C., Metcalf J., Shmueli U., Davis A., Naqvi A., Lee T., Ryder S. D., Collier J., Klass H., Ninkovic M., Cramp M., Sharer N., Aspinall R., Goggin P., Ghosh D., Douds A., Hoeroldt B., Booth J., Williams E., Hussaini H., Stableforth W., Ayres R., Thorburn D., Marshall E., Burroughs A., Mann S., Lombard M., Richardson P., Patanwala I., Maltby J., Brookes M., Mathew R., Vyas S., Singhal S., Gleeson D., Misra S., Butterworth J., George K., Harding T., Douglass A., Panter S., Shearman J., Bray G., Butcher G., Forton D., Mclindon J., Cowan M., Whatley G., Mandal A., Gupta H., Sanghi P., Jain S., Pereira S., Prasad G., Watts G., Wright M., Neuberger J., Gordon F., Unitt E., Grant A., Delahooke T., Higham A., Brind A., Cox M., Ramakrishnan S., King A., Collins C., Whalley S., Li A., Fraser J., Bell A., Wong V. S., Singhal A., Gee I., Ang Y., Ransford R., Gotto J., Millson C., Bowles J., Thomas C., Harrison M., Galaska R., Kendall J., Whiteman J., Lawlor C., Gray C., Elliott K., Mulvaney-Jones C., Hobson L., Van Duyvenvoorde G., Loftus A., Seward K., Penn R., Maiden J., Damant R., Hails J., Cloudsdale R., Silvestre V., Glenn S., Dungca E., Wheatley N., Doyle H., Kent M., Hamilton C., Braim D., Wooldridge H., Abrahams R., Paton A., Lancaster N., Gibbins A., Hogben K., Desousa P., Muscariu F., Musselwhite J., McKay A., Tan L., Foale C., Brighton J., Flahive K., Nambela E., Townshend P., Ford C., Holder S., Palmer C., Featherstone J., Nasseri M., Sadeghian J., Williams B., Rolls S. -A., Hynes A., Duggan C., Crossey M., Stansfield G., MacNicol C., Wilkins J., Wilhelmsen E., Raymode P., Lee H. -J., Durant E., Bishop R., Ncube N., Tripoli S., Casey R., Cowley C., Miller R., Houghton K., Ducker S., Wright F., Bird B., Baxter G., Keggans J., Hughes M., Grieve E., Young K., Williams D., Ocker K., Hines F., Martin K., Innes C., Valliani T., Fairlamb H., Thornthwaite S., Eastick A., Tanqueray E., Morrison J., Holbrook B., Browning J., Walker K., Congreave S., Verheyden J., Slininger S., Stafford L., O'Donnell D., Ainsworth M., Lord S., Kent L., March L., Dickson C., Simpson D., Longhurst B., Hayes M., Shpuza E., White N., Besley S., Pearson S., Wright A., Jones L., Gunter E., Dewhurst H., Fouracres A., Farrington L., Graves L., Marriott S., Leoni M., Tyrer D., Dali-kemmery L., Lambourne V., Green M., Sirdefield D., Amor K., Colley J., Shinder B., Jones J., Mills M., Carnahan M., Taylor N., Boulton K., Tregonning J., Brown C., Clifford G., Archer E., Hamilton M., Curtis J., Shewan T., Walsh S., Warner K., Netherton K., Mupudzi M., Gunson B., Gitahi J., Gocher D., Batham S., Pateman H., Desmennu S., Conder J., Clement D., Gallagher S., Orpe J., Chan P., Currie L., O'Donohoe L., Oblak M., Morgan L., Quinn M., Amey I., Baird Y., Cotterill D., Cumlat L., Winter L., Greer S., Spurdle K., Allison J., Dyer S., Sweeting H., Kordula J., Aiba Y., Nakamura H., Abiru S., Nagaoka S., Komori A., Yatsuhashi H., Ishibashi H., Ito M., Kawai Y., Kohn S. -S., Gervais O., Migita K., Katsushima S., Naganuma A., Sugi K., Komatsu T., Mannami T., Matsushita K., Yoshizawa K., Makita F., Nikami T., Nishimura H., Kouno H., Ota H., Komura T., Nakamura Y., Shimada M., Hirashima N., Komeda T., Ario K., Nakamuta M., Yamashita T., Furuta K., Kikuchi M., Naeshiro N., Takahashi H., Mano Y., Tsunematsu S., Yabuuchi I., Shimada Y., Yamauchi K., Sugimoto R., Sakai H., Mita E., Koda M., Tsuruta S., Kamitsukasa H., Sato T., Masaki N., Kobata T., Fukushima N., Higuchi N., Ohara Y., Muro T., Takesaki E., Takaki H., Yamamoto T., Kato M., Nagaoki Y., Hayashi S., Ishida J., Watanabe Y., Kobayashi M., Koga M., Saoshiro T., Yagura M., Hirata K., Takikawa H., Ohira H., Zeniya M., Abe M., Onji M., Kaneko S., Honda M., Arai K., Arinaga-Hino T., Hashimoto E., Taniai M., Umemura T., Joshita S., Nakao K., Ichikawa T., Shibata H., Yamagiwa S., Seike M., Honda K., Sakisaka S., Takeyama Y., Harada M., Senju M., Yokosuka O., Kanda T., Ueno Y., Kikuchi K., Ebinuma H., Himoto T., Yasunami M., Murata K., Mizokami M., Shimoda S., Miyake Y., Takaki A., Yamamoto K., Hirano K., Ichida T., Ido A., Tsubouchi H., Chayama K., Harada K., Nakanuma Y., Maehara Y., Taketomi A., Shirabe K., Soejima Y., Mori A., Yagi S., Uemoto S., Tanaka T., Yamashiki N., Tamura S., Sugawara Y., Kokudo N., Carbone M., Cardamone G., Duga S., Gershwin M. E., Seldin M. F., and Invernizzi P.

Abstract

Background & Aims: Genome-wide association studies in primary biliary cholangitis (PBC) have failed to find X chromosome (chrX) variants associated with the disease. Here, we specifically explore the chrX contribution to PBC, a sexually dimorphic complex autoimmune disease. Methods: We performed a chrX-wide association study, including genotype data from 5 genome-wide association studies (from Italy, United Kingdom, Canada, China, and Japan; 5244 case patients and 11,875 control individuals). Results: Single-marker association analyses found approximately 100 loci displaying P < 5 × 10–4, with the most significant being a signal within the OTUD5 gene (rs3027490; P = 4.80 × 10–6; odds ratio [OR], 1.39; 95% confidence interval [CI], 1.028–1.88; Japanese cohort). Although the transethnic meta-analysis evidenced only a suggestive signal (rs2239452, mapping within the PIM2 gene; OR, 1.17; 95% CI, 1.09–1.26; P = 9.93 × 10–8), the population-specific meta-analysis showed a genome-wide significant locus in East Asian individuals pointing to the same region (rs7059064, mapping within the GRIPAP1 gene; P = 6.2 × 10–9; OR, 1.33; 95% CI, 1.21–1.46). Indeed, rs7059064 tags a unique linkage disequilibrium block including 7 genes: TIMM17B, PQBP1, PIM2, SLC35A2, OTUD5, KCND1, and GRIPAP1, as well as a superenhancer (GH0XJ048933 within OTUD5) targeting all these genes. GH0XJ048933 is also predicted to target FOXP3, the main T-regulatory cell lineage specification factor. Consistently, OTUD5 and FOXP3 RNA levels were up-regulated in PBC case patients (1.75- and 1.64-fold, respectively). Conclusions: This work represents the first comprehensive study, to our knowledge, of the chrX contribution to the genetics of an autoimmune liver disease and shows a novel PBC-related genome-wide significant locus.

Published
2021

35. What controls tropical forest architecture? Testing environmental, structural and floristic drivers

Author

Banin, L., Feldpausch, T. R., Phillips, O. L., Baker, T. R., Lloyd, J., Affum-Baffoe, K., Arets, E. J. M. M., Berry, N. J., Bradford, M., Brienen, R. J. W., Davies, S., Drescher, M., Higuchi, N., Hilbert, D. W., Hladik, A., Iida, Y., Salim, K. Abu, Kassim, A. R., King, D. A., Lopez-Gonzalez, G., Metcalfe, D., Nilus, R., Peh, K. S.-H., Reitsma, J. M., Sonké, B., Taedoumg, H., Tan, S., White, L., Wöll, H., and Lewis, S. L.

Published
2012
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36. Stem respiration and growth in a central Amazon rainforest

Author

Jardine, KJ, Jardine, KJ, Cobello, LO, Teixeira, LM, East, MMS, Levine, S, Gimenez, BO, Robles, E, Spanner, G, Koven, C, Xu, C, Warren, JM, Higuchi, N, McDowell, N, Pastorello, G, Chambers, JQ, Jardine, KJ, Jardine, KJ, Cobello, LO, Teixeira, LM, East, MMS, Levine, S, Gimenez, BO, Robles, E, Spanner, G, Koven, C, Xu, C, Warren, JM, Higuchi, N, McDowell, N, Pastorello, G, and Chambers, JQ

Abstract

Key message: Annual stem CO2 efflux increases with stem wood production rates and are inhibited by daily moisture stress. Abstract: Tropical forests cycle a large amount of CO2 between the land and atmosphere, with a substantial portion of the return flux due tree respiratory processes. However, in situ estimates of woody tissue respiratory fluxes and carbon use efficiencies (CUEW) and their dependencies on physiological processes including stem wood production (Pw) and transpiration in tropical forests remain scarce. Here, we synthesize monthly Pw and daytime stem CO2 efflux (ES) measurements over 1 year from 80 trees with variable biomass accumulation rates in the central Amazon. On average, carbon flux to woody tissues, expressed in the same stem area normalized units as ES, averaged 0.90 ± 1.2 µmol m−2 s−1 for Pw, and 0.55 ± 0.33 µmol m−2 s−1 for daytime ES. A positive linear correlation was found between stem growth rates and stem CO2 efflux, with respiratory carbon loss equivalent to 15 ± 3% of stem carbon accrual. CUEW of stems was non-linearly correlated with growth and was as high as 77–87% for a fast-growing tree. Diurnal measurements of stem CO2 efflux for three individuals showed a daytime reduction of ES by 15–50% during periods of high sap flow and transpiration. The results demonstrate that high daytime ES fluxes are associated with high CUEW during fast tree growth, reaching higher values than previously observed in the Amazon Basin (e.g., maximum CUEW up to 77–87%, versus 30–56%). The observations are consistent with the emerging view that diurnal dynamics of stem water status influences growth processes and associated respiratory metabolism.

Published
2022

37. Soil moisture thresholds explain a shift from light-limited to water-limited sap velocity in the Central Amazon during the 2015-16 El Niño drought

Author

Meng, L, Meng, L, Chambers, J, Koven, C, Pastorello, G, Gimenez, B, Jardine, K, Tang, Y, McDowell, N, Negron-Juarez, R, Longo, M, Araujo, A, Tomasella, J, Fontes, C, Mohan, M, Higuchi, N, Meng, L, Meng, L, Chambers, J, Koven, C, Pastorello, G, Gimenez, B, Jardine, K, Tang, Y, McDowell, N, Negron-Juarez, R, Longo, M, Araujo, A, Tomasella, J, Fontes, C, Mohan, M, and Higuchi, N

Abstract

Transpiration is often considered to be light- but not water-limited in humid tropical rainforests due to abundant soil water, even during the dry seasons. The record-breaking 2015-16 El Niño drought provided a unique opportunity to examine whether transpiration is constrained by water under severe lack of rainfall. We measured sap velocity, soil water content, and meteorological variables in an old-growth upland forest in the Central Amazon throughout the 2015-16 drought. We found a rapid decline in sap velocity (-38 ± 21%, mean ± SD.) and in its temporal variability (-88%) during the drought compared to the wet season. Such changes were accompanied by a marked decline in soil moisture and an increase in temperature and vapor pressure deficit. Sap velocity was largely limited by net radiation during the wet and normal dry seasons; however, it shifted to be primarily limited by soil moisture during the drought. The threshold in which sap velocity became dominated by soil moisture was at 0.33 m3 m-3 (around -150 kPa in soil matric potential), below which sap velocity dropped steeply. Our study provides evidence for a soil water threshold on transpiration in a moist tropical forest, suggesting a shift from light limitation to water limitation under future climate characterized by increased temperature and an increased frequency, intensity, duration and extent of extreme drought events.

Published
2022

43. Long-term decline of the Amazon carbon sink

Author

Brienen, R. J. W., Phillips, O. L., Feldpausch, T. R., Gloor, E., Baker, T. R., Lloyd, J., Lopez-Gonzalez, G., Monteagudo-Mendoza, A., Malhi, Y., Lewis, S. L., Vásquez Martinez, R., Alexiades, M., Álvarez Dávila, E., Alvarez-Loayza, P., Andrade, A., Aragão, L. E. O. C., Araujo-Murakami, A., Arets, E. J. M. M., Arroyo, L., Aymard, G. A. C., Bánki, O. S., Baraloto, C., Barroso, J., Bonal, D., Boot, R. G. A., Camargo, J. L. C., Castilho, C. V., Chama, V., Chao, K. J., Chave, J., Comiskey, J. A., Cornejo Valverde, F., da Costa, L., de Oliveira, E. A., Di Fiore, A., Erwin, T. L., Fauset, S., Forsthofer, M., Galbraith, D. R., Grahame, E. S., Groot, N., Hérault, B., Higuchi, N., Honorio Coronado, E. N., Keeling, H., Killeen, T. J., Laurance, W. F., Laurance, S., Licona, J., Magnussen, W. E., Marimon, B. S., Marimon-Junior, B. H., Mendoza, C., Neill, D. A., Nogueira, E. M., Núñez, P., Pallqui Camacho, N. C., Parada, A., Pardo-Molina, G., Peacock, J., Peña-Claros, M., Pickavance, G. C., Pitman, N. C. A., Poorter, L., Prieto, A., Quesada, C. A., Ramírez, F., Ramírez-Angulo, H., Restrepo, Z., Roopsind, A., Rudas, A., Salomão, R. P., Schwarz, M., Silva, N., Silva-Espejo, J. E., Silveira, M., Stropp, J., Talbot, J., ter Steege, H., Teran-Aguilar, J., Terborgh, J., Thomas-Caesar, R., Toledo, M., Torello-Raventos, M., Umetsu, R. K., van der Heijden, G. M. F., van der Hout, P., Guimarães Vieira, I. C., Vieira, S. A., Vilanova, E., Vos, V. A., and Zagt, R. J.

Published
2015
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45. Author Correction: Tree mode of death and mortality risk factors across Amazon forests (Nature Communications, (2020), 11, 1, (5515), 10.1038/s41467-020-18996-3)

Author

Esquivel-Muelbert, A, Phillips, OL, Brienen, RJW, Fauset, S, Sullivan, MJP, Baker, TR, Chao, KJ, Feldpausch, TR, Gloor, E, Higuchi, N, Houwing-Duistermaat, J, Lloyd, J, Liu, H, Malhi, Y, Marimon, B, Marimon Junior, BH, Monteagudo-Mendoza, A, Poorter, L, Silveira, M, Torre, EV, Dávila, EA, del Aguila Pasquel, J, Almeida, E, Loayza, PA, Andrade, A, Aragão, LEOC, Araujo-Murakami, A, Arets, E, Arroyo, L, Aymard C, GA, Baisie, M, Baraloto, C, Camargo, PB, Barroso, J, Blanc, L, Bonal, D, Bongers, F, Boot, R, Brown, F, Burban, B, Camargo, JL, Castro, W, Moscoso, VC, Chave, J, Comiskey, J, Valverde, FC, da Costa, AL, Cardozo, ND, Di Fiore, A, Dourdain, A, Erwin, T, Llampazo, GF, Vieira, ICG, Herrera, R, Honorio Coronado, E, Huamantupa-Chuquimaco, I, Jimenez-Rojas, E, Killeen, T, Laurance, S, Laurance, W, Levesley, A, Lewis, SL, Ladvocat, KLLM, Lopez-Gonzalez, G, Lovejoy, T, Meir, P, Mendoza, C, Morandi, P, Neill, D, Nogueira Lima, AJ, Vargas, PN, de Oliveira, EA, Camacho, NP, Pardo, G, Peacock, J, Peña-Claros, M, Peñuela-Mora, MC, Pickavance, G, Pipoly, J, Pitman, N, Prieto, A, Pugh, TAM, Quesada, C, Ramirez-Angulo, H, de Almeida Reis, SM, Rejou-Machain, M, Correa, ZR, Bayona, LR, Rudas, A, Salomão, R, Serrano, J, Espejo, JS, Silva, N, Singh, J, Stahl, C, Stropp, J, Swamy, V, Talbot, J, ter Steege, H, and Terborgh, J

Abstract

The original version of this Article contained an error in Table 2, where the number of individuals in the “All Amazonia” row was reported as 11,6431 instead of 116,431. Also, the original version of this Article contained an error in the Methods, where the R2 for the proportion of broken/uprooted dead trees increase per year was reported as 0.12, the correct value being 0.06. The original version of this Article contained errors in the author affiliations. The affiliation of Gerardo A. Aymard C. with UNELLEZGuanare, Herbario Universitario (PORT), Portuguesa, Venezuela Compensation International Progress S.A. Ciprogress–Greenlife.

Published
2021

46. Canopy Position Influences the Degree of Light Suppression of Leaf Respiration in Abundant Tree Genera in the Amazon Forest

Author

Souza, DC, Souza, DC, Jardine, KJ, Rodrigues, JVFC, Gimenez, BO, Rogers, A, McDowell, N, Walker, AP, Higuchi, N, Sampaio-Filho, IJ, Chambers, J, Souza, DC, Souza, DC, Jardine, KJ, Rodrigues, JVFC, Gimenez, BO, Rogers, A, McDowell, N, Walker, AP, Higuchi, N, Sampaio-Filho, IJ, and Chambers, J

Abstract

Leaf respiration in the dark (Rdark) and light (Rday) is poorly characterized in diverse tropical ecosystems, and little to no information exists on the degree of light suppression in common tree species within the Amazon basin, and their dependences upon plant functional traits and position within the canopy. We quantified Rdark and apparent Rday using the Kok method and measured key leaf traits in 26 tree individuals of different species distributed in three different canopy positions: canopy, lower canopy, and understory. To explore the relationships between the leaf traits we used the standardized major axis (SMA). We found that canopy trees had significantly higher rates of Rdark and Rday than trees in the understory. The difference between Rdark and Rday (the light suppression of respiration) was greatest in the understory (68 ± 9%, 95% CI) and lower canopy (49 ± 9%, 95% CI) when compared to the canopy (37 ± 10%, 95% CI). We also found that Rday was significantly and strongly correlated with Rdark (p < 0.001) for all the canopy positions. Also, leaf mass per area (LMA) and leaf Phosphorus concentration (P) had a significant relationship with Rdark (p < 0.001; p = 0.003), respectively. In addition, a significant relationship was found for LMA in the canopy and lower canopy positions (p = 0.009; p = 0.048) while P was only significant in the canopy (p = 0.044). Finally, no significant relationship was found between Rdark and nitrogen, sugars, and starch. Our results highlight the importance of including representation of the light suppression of leaf respiration in terrestrial biosphere models and also of accounting for vertical gradients within forest canopies and connections with functional traits.

Published
2021

47. Amazon tree dominance across forest strata

Author

Draper, F.C., Costa, F.R.C., Arellano, G., Phillips, O.L., Duque, A., Macía, M.J., ter Steege, H., Asner, G.P., Berenguer, E., Schietti, J., Socolar, J.B., de Souza, F.C., Dexter, K.G., Jørgensen, P.M., Tello, J.S., Magnusson, W.E., Baker, T.R., Castilho, C.V., Monteagudo-Mendoza, A., Fine, P.V.A., Ruokolainen, K., Coronado, E.N.H., Aymard, G., Dávila, N., Sáenz, M.S., Paredes, M.A.R., Engel, J., Fortunel, C., Paine, C.E.T., Goret, J.-Y., Dourdain, A., Petronelli, P., Allie, E., Andino, J.E.G., Brienen, R.J.W., Pérez, L.C., Manzatto, Â.G., Zambrana, N.Y.P., Molino, J.-F., Sabatier, D., Chave, J., Fauset, S., Villacorta, R.G., Réjou-Méchain, M., Berry, P.E., Melgaço, K., Feldpausch, T.R., Sandoval, E.V., Martinez, R.V., Mesones, I., Junqueira, A.B., Roucoux, K.H., de Toledo, J.J., Andrade, A.C., Camargo, J.L., del Aguila Pasquel, J., Santana, F.D., Laurance, W.F., Laurance, S.G., Lovejoy, T.E., Comiskey, J.A., Galbraith, D.R., Kalamandeen, M., Aguilar, G.E.N., Arenas, J.V., Guerra, C.A.A., Flores, M., Llampazo, G.F., Montenegro, L.A.T., Gomez, R.Z., Pansonato, M.P., Moscoso, V.C., Vleminckx, J., Barrantes, O.J.V., Duivenvoorden, J.F., de Sousa, S.A., Arroyo, L., Perdiz, R.O., Cravo, J.S., Marimon, B.S., Junior, B.H.M., Carvalho, F.A., Damasco, G., Disney, M., Vital, M.S., Diaz, P.R.S., Vicentini, A., Nascimento, H., Higuchi, N., Van Andel, T., Malhi, Y., Ribeiro, S.C., Terborgh, J.W., Thomas, R.S., Dallmeier, F., Prieto, A., Hilário, R.R., Salomão, R.P., Silva, R.C., Casas, L.F., Vieira, I.C.G., Araujo-Murakami, A., Arevalo, F.R., Ramírez-Angulo, H., Torre, E.V., Peñuela, M.C., Killeen, T.J., Pardo, G., Jimenez-Rojas, E., Castro, W., Cabrera, D.G., Pipoly, J., de Sousa, T.R., Silvera, M., Vos, V., Neill, D., Vargas, P.N., Vela, D.M., Aragão, L.E.O.C., Umetsu, R.K., Sierra, R., Wang, O., Young, K.R., Prestes, N.C.C.S., Massi, K.G., Huaymacari, J.R., Gutierrez, G.A.P., Aldana, A.M., Alexiades, M.N., Baccaro, F., Céron, C., Muelbert, A.E., Rios, J.M.G., Lima, A.S., Lloyd, J.L., Pitman, N.C.A., Gamarra, L.V., Oroche, C.J.C., Fuentes, A.F., Palacios, W., Patiño, S., Torres-Lezama, A., Baraloto, C., Draper, F.C., Costa, F.R.C., Arellano, G., Phillips, O.L., Duque, A., Macía, M.J., ter Steege, H., Asner, G.P., Berenguer, E., Schietti, J., Socolar, J.B., de Souza, F.C., Dexter, K.G., Jørgensen, P.M., Tello, J.S., Magnusson, W.E., Baker, T.R., Castilho, C.V., Monteagudo-Mendoza, A., Fine, P.V.A., Ruokolainen, K., Coronado, E.N.H., Aymard, G., Dávila, N., Sáenz, M.S., Paredes, M.A.R., Engel, J., Fortunel, C., Paine, C.E.T., Goret, J.-Y., Dourdain, A., Petronelli, P., Allie, E., Andino, J.E.G., Brienen, R.J.W., Pérez, L.C., Manzatto, Â.G., Zambrana, N.Y.P., Molino, J.-F., Sabatier, D., Chave, J., Fauset, S., Villacorta, R.G., Réjou-Méchain, M., Berry, P.E., Melgaço, K., Feldpausch, T.R., Sandoval, E.V., Martinez, R.V., Mesones, I., Junqueira, A.B., Roucoux, K.H., de Toledo, J.J., Andrade, A.C., Camargo, J.L., del Aguila Pasquel, J., Santana, F.D., Laurance, W.F., Laurance, S.G., Lovejoy, T.E., Comiskey, J.A., Galbraith, D.R., Kalamandeen, M., Aguilar, G.E.N., Arenas, J.V., Guerra, C.A.A., Flores, M., Llampazo, G.F., Montenegro, L.A.T., Gomez, R.Z., Pansonato, M.P., Moscoso, V.C., Vleminckx, J., Barrantes, O.J.V., Duivenvoorden, J.F., de Sousa, S.A., Arroyo, L., Perdiz, R.O., Cravo, J.S., Marimon, B.S., Junior, B.H.M., Carvalho, F.A., Damasco, G., Disney, M., Vital, M.S., Diaz, P.R.S., Vicentini, A., Nascimento, H., Higuchi, N., Van Andel, T., Malhi, Y., Ribeiro, S.C., Terborgh, J.W., Thomas, R.S., Dallmeier, F., Prieto, A., Hilário, R.R., Salomão, R.P., Silva, R.C., Casas, L.F., Vieira, I.C.G., Araujo-Murakami, A., Arevalo, F.R., Ramírez-Angulo, H., Torre, E.V., Peñuela, M.C., Killeen, T.J., Pardo, G., Jimenez-Rojas, E., Castro, W., Cabrera, D.G., Pipoly, J., de Sousa, T.R., Silvera, M., Vos, V., Neill, D., Vargas, P.N., Vela, D.M., Aragão, L.E.O.C., Umetsu, R.K., Sierra, R., Wang, O., Young, K.R., Prestes, N.C.C.S., Massi, K.G., Huaymacari, J.R., Gutierrez, G.A.P., Aldana, A.M., Alexiades, M.N., Baccaro, F., Céron, C., Muelbert, A.E., Rios, J.M.G., Lima, A.S., Lloyd, J.L., Pitman, N.C.A., Gamarra, L.V., Oroche, C.J.C., Fuentes, A.F., Palacios, W., Patiño, S., Torres-Lezama, A., and Baraloto, C.

Abstract

The forests of Amazonia are among the most biodiverse plant communities on Earth. Given the immediate threats posed by climate and land-use change, an improved understanding of how this extraordinary biodiversity is spatially organized is urgently required to develop effective conservation strategies. Most Amazonian tree species are extremely rare but a few are common across the region. Indeed, just 227 ‘hyperdominant’ species account for >50% of all individuals >10 cm diameter at 1.3 m in height. Yet, the degree to which the phenomenon of hyperdominance is sensitive to tree size, the extent to which the composition of dominant species changes with size class and how evolutionary history constrains tree hyperdominance, all remain unknown. Here, we use a large floristic dataset to show that, while hyperdominance is a universal phenomenon across forest strata, different species dominate the forest understory, midstory and canopy. We further find that, although species belonging to a range of phylogenetically dispersed lineages have become hyperdominant in small size classes, hyperdominants in large size classes are restricted to a few lineages. Our results demonstrate that it is essential to consider all forest strata to understand regional patterns of dominance and composition in Amazonia. More generally, through the lens of 654 hyperdominant species, we outline a tractable pathway for understanding the functioning of half of Amazonian forests across vertical strata and geographical locations.

Published
2021

48. X Chromosome Contribution to the Genetic Architecture of Primary Biliary Cholangitis

Author

Asselta, R., Paraboschi, E. M., Gerussi, A., Cordell, H. J., Mells, G. F., Sandford, R. N., Jones, D. E., Nakamura, M., Ueno, K., Hitomi, Y., Kawashima, M., Nishida, N., Tokunaga, K., Nagasaki, M., Tanaka, A., Tang, R., Li, Z., Shi, Y., Liu, X., Xiong, M., Hirschfield, G., Siminovitch, K. A., Walker, E., Xie, G., Mason, A., Myers, R., Peltekian, K., Ghent, C., Atkinson, E., Juran, B., Lazaridis, K., Lu, Y., Gu, X., Jing, K., Amos, C., Affronti, A., Brunetto, M., Coco, B., Spinzi, G., Elia, G., Ferrari, C., Lleo, A., Muratori, L., Muratori, P., Portincasa, P., Colli, A., Bruno, S., Colloredo, G., Azzaroli, F., Andreone, P., Bragazzi, M., Alvaro, D., Cardinale, V., Cazzagon, N., Rigamonti, C., Floreani, A., Rosina, F., Ciaccio, A., Cristoferi, L., D'Amato, D., Malinverno, F., Mancuso, C., Massironi, S., Milani, C., O'Donnell, S. E., Ronca, V., Barisani, D., Lampertico, P., Donato, Federica, Fagiuoli, Stefano, Almasio, P. L., Giannini, E., Cursaro, C., Colombo, M., Valenti, L., Miele, Luca, Andriulli, A., Niro, G. A., Grattagliano, I., Morini, L., Casella, G., Vinci, Maria Rosaria, Battezzati, P. M., Crosignani, A., Zuin, M., Mattalia, A., Calvaruso, V., Colombo, S., Benedetti, A., Marzioni, M., Galli, A., Marra, F., Tarocchi, M., Picciotto, A., Morisco, F., Fabris, L., Croce, L. S., Tiribelli, C., Toniutto, P., Strazzabosco, M., Ch'Ng, C. L., Rahman, M., Yapp, T., Sturgess, R., Healey, C., Czajkowski, M., Gunasekera, A., Gyawali, P., Premchand, P., Kapur, K., Marley, R., Foster, G., Watson, A., Dias, A., Subhani, J., Harvey, R., Mccorry, R., Ramanaden, D., Gasem, J., Evans, R., Mathialahan, T., Shorrock, C., Lipscomb, G., Southern, P., Tibble, J., Gorard, D., Palegwala, A., Jones, S., Dawwas, M., Alexander, G., Dolwani, S., Prince, M., Foxton, M., Elphick, D., Mitchison, H., Gooding, I., Karmo, M., Saksena, S., Mendall, M., Patel, M., Ede, R., Austin, A., Sayer, J., Hankey, L., Hovell, C., Fisher, N., Carter, M., Koss, K., Piotrowicz, A., Grimley, C., Neal, D., Lim, G., Levi, S., Ala, A., Broad, A., Saeed, A., Wood, G., Brown, J., Wilkinson, M., Gordon, H., Ramage, J., Ridpath, J., Ngatchu, T., Grover, B., Shaukat, S., Shidrawi, R., Abouda, G., Ali, F., Rees, I., Salam, I., Narain, M., Brown, A., Taylor-Robinson, S., Williams, S., Grellier, L., Banim, P., Das, D., Chilton, A., Heneghan, M., Curtis, H., Gess, M., Drake, I., Aldersley, M., Davies, M., Jones, R., Mcnair, A., Srirajaskanthan, R., Pitcher, M., Sen, S., Bird, G., Barnardo, A., Kitchen, P., Yoong, K., Chirag, O., Sivaramakrishnan, N., Macfaul, G., Jones, D., Shah, A., Evans, C., Saha, S., Pollock, K., Bramley, P., Mukhopadhya, A., Fraser, A., Mills, P., Shallcross, C., Campbell, S., Bathgate, A., Shepherd, A., Dillon, J., Rushbrook, S., Przemioslo, R., Macdonald, C., Metcalf, J., Shmueli, U., Davis, A., Naqvi, A., Lee, T., Ryder, S. D., Collier, J., Klass, H., Ninkovic, M., Cramp, M., Sharer, N., Aspinall, R., Goggin, P., Ghosh, D., Douds, A., Hoeroldt, B., Booth, J., Williams, E., Hussaini, H., Stableforth, W., Ayres, R., Thorburn, D., Marshall, E., Burroughs, A., Mann, S., Lombard, M., Richardson, P., Patanwala, I., Maltby, J., Brookes, M., Mathew, R., Vyas, S., Singhal, S., Gleeson, D., Misra, S., Butterworth, J., George, K., Harding, T., Douglass, A., Panter, S., Shearman, J., Bray, G., Butcher, G., Forton, D., Mclindon, J., Cowan, M., Whatley, G., Mandal, A., Gupta, H., Sanghi, P., Jain, S., Pereira, S., Prasad, G., Watts, G., Wright, M., Neuberger, J., Gordon, F., Unitt, E., Grant, A., Delahooke, T., Higham, A., Brind, A., Cox, M., Ramakrishnan, S., King, A., Collins, C., Whalley, S., Li, A., Fraser, J., Bell, A., Wong, V. 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E., Seldin, M. F., Invernizzi, P., Donato F., Fagiuoli S., Miele L. (ORCID:0000-0003-3464-0068), Vinci M., Asselta, R., Paraboschi, E. M., Gerussi, A., Cordell, H. J., Mells, G. F., Sandford, R. N., Jones, D. E., Nakamura, M., Ueno, K., Hitomi, Y., Kawashima, M., Nishida, N., Tokunaga, K., Nagasaki, M., Tanaka, A., Tang, R., Li, Z., Shi, Y., Liu, X., Xiong, M., Hirschfield, G., Siminovitch, K. A., Walker, E., Xie, G., Mason, A., Myers, R., Peltekian, K., Ghent, C., Atkinson, E., Juran, B., Lazaridis, K., Lu, Y., Gu, X., Jing, K., Amos, C., Affronti, A., Brunetto, M., Coco, B., Spinzi, G., Elia, G., Ferrari, C., Lleo, A., Muratori, L., Muratori, P., Portincasa, P., Colli, A., Bruno, S., Colloredo, G., Azzaroli, F., Andreone, P., Bragazzi, M., Alvaro, D., Cardinale, V., Cazzagon, N., Rigamonti, C., Floreani, A., Rosina, F., Ciaccio, A., Cristoferi, L., D'Amato, D., Malinverno, F., Mancuso, C., Massironi, S., Milani, C., O'Donnell, S. E., Ronca, V., Barisani, D., Lampertico, P., Donato, Federica, Fagiuoli, Stefano, Almasio, P. L., Giannini, E., Cursaro, C., Colombo, M., Valenti, L., Miele, Luca, Andriulli, A., Niro, G. A., Grattagliano, I., Morini, L., Casella, G., Vinci, Maria Rosaria, Battezzati, P. M., Crosignani, A., Zuin, M., Mattalia, A., Calvaruso, V., Colombo, S., Benedetti, A., Marzioni, M., Galli, A., Marra, F., Tarocchi, M., Picciotto, A., Morisco, F., Fabris, L., Croce, L. S., Tiribelli, C., Toniutto, P., Strazzabosco, M., Ch'Ng, C. L., Rahman, M., Yapp, T., Sturgess, R., Healey, C., Czajkowski, M., Gunasekera, A., Gyawali, P., Premchand, P., Kapur, K., Marley, R., Foster, G., Watson, A., Dias, A., Subhani, J., Harvey, R., Mccorry, R., Ramanaden, D., Gasem, J., Evans, R., Mathialahan, T., Shorrock, C., Lipscomb, G., Southern, P., Tibble, J., Gorard, D., Palegwala, A., Jones, S., Dawwas, M., Alexander, G., Dolwani, S., Prince, M., Foxton, M., Elphick, D., Mitchison, H., Gooding, I., Karmo, M., Saksena, S., Mendall, M., Patel, M., Ede, R., Austin, A., Sayer, J., Hankey, L., Hovell, C., Fisher, N., Carter, M., Koss, K., Piotrowicz, A., Grimley, C., Neal, D., Lim, G., Levi, S., Ala, A., Broad, A., Saeed, A., Wood, G., Brown, J., Wilkinson, M., Gordon, H., Ramage, J., Ridpath, J., Ngatchu, T., Grover, B., Shaukat, S., Shidrawi, R., Abouda, G., Ali, F., Rees, I., Salam, I., Narain, M., Brown, A., Taylor-Robinson, S., Williams, S., Grellier, L., Banim, P., Das, D., Chilton, A., Heneghan, M., Curtis, H., Gess, M., Drake, I., Aldersley, M., Davies, M., Jones, R., Mcnair, A., Srirajaskanthan, R., Pitcher, M., Sen, S., Bird, G., Barnardo, A., Kitchen, P., Yoong, K., Chirag, O., Sivaramakrishnan, N., Macfaul, G., Jones, D., Shah, A., Evans, C., Saha, S., Pollock, K., Bramley, P., Mukhopadhya, A., Fraser, A., Mills, P., Shallcross, C., Campbell, S., Bathgate, A., Shepherd, A., Dillon, J., Rushbrook, S., Przemioslo, R., Macdonald, C., Metcalf, J., Shmueli, U., Davis, A., Naqvi, A., Lee, T., Ryder, S. D., Collier, J., Klass, H., Ninkovic, M., Cramp, M., Sharer, N., Aspinall, R., Goggin, P., Ghosh, D., Douds, A., Hoeroldt, B., Booth, J., Williams, E., Hussaini, H., Stableforth, W., Ayres, R., Thorburn, D., Marshall, E., Burroughs, A., Mann, S., Lombard, M., Richardson, P., Patanwala, I., Maltby, J., Brookes, M., Mathew, R., Vyas, S., Singhal, S., Gleeson, D., Misra, S., Butterworth, J., George, K., Harding, T., Douglass, A., Panter, S., Shearman, J., Bray, G., Butcher, G., Forton, D., Mclindon, J., Cowan, M., Whatley, G., Mandal, A., Gupta, H., Sanghi, P., Jain, S., Pereira, S., Prasad, G., Watts, G., Wright, M., Neuberger, J., Gordon, F., Unitt, E., Grant, A., Delahooke, T., Higham, A., Brind, A., Cox, M., Ramakrishnan, S., King, A., Collins, C., Whalley, S., Li, A., Fraser, J., Bell, A., Wong, V. S., Singhal, A., Gee, I., Ang, Y., Ransford, R., Gotto, J., Millson, C., Bowles, J., Thomas, C., Harrison, M., Galaska, R., Kendall, J., Whiteman, J., Lawlor, C., Gray, C., Elliott, K., Mulvaney-Jones, C., Hobson, L., Van Duyvenvoorde, G., Loftus, A., Seward, K., Penn, R., Maiden, J., Damant, R., Hails, J., Cloudsdale, R., Silvestre, V., Glenn, S., Dungca, E., Wheatley, N., Doyle, H., Kent, M., Hamilton, C., Braim, D., Wooldridge, H., Abrahams, R., Paton, A., Lancaster, N., Gibbins, A., Hogben, K., Desousa, P., Muscariu, F., Musselwhite, J., Mckay, A., Tan, L., Foale, C., Brighton, J., Flahive, K., Nambela, E., Townshend, P., Ford, C., Holder, S., Palmer, C., Featherstone, J., Nasseri, M., Sadeghian, J., Williams, B., Rolls, S. -A., Hynes, A., Duggan, C., Crossey, M., Stansfield, G., Macnicol, C., Wilkins, J., Wilhelmsen, E., Raymode, P., Lee, H. -J., Durant, E., Bishop, R., Ncube, N., Tripoli, S., Casey, R., Cowley, C., Miller, R., Houghton, K., Ducker, S., Wright, F., Bird, B., Baxter, G., Keggans, J., Hughes, M., Grieve, E., Young, K., Williams, D., Ocker, K., Hines, F., Martin, K., Innes, C., Valliani, T., Fairlamb, H., Thornthwaite, S., Eastick, A., Tanqueray, E., Morrison, J., Holbrook, B., Browning, J., Walker, K., Congreave, S., Verheyden, J., Slininger, S., Stafford, L., O'Donnell, D., Ainsworth, M., Lord, S., Kent, L., March, L., Dickson, C., Simpson, D., Longhurst, B., Hayes, M., Shpuza, E., White, N., Besley, S., Pearson, S., Wright, A., Jones, L., Gunter, E., Dewhurst, H., Fouracres, A., Farrington, L., Graves, L., Marriott, S., Leoni, M., Tyrer, D., Dali-kemmery, L., Lambourne, V., Green, M., Sirdefield, D., Amor, K., Colley, J., Shinder, B., Jones, J., Mills, M., Carnahan, M., Taylor, N., Boulton, K., Tregonning, J., Brown, C., Clifford, G., Archer, E., Hamilton, M., Curtis, J., Shewan, T., Walsh, S., Warner, K., Netherton, K., Mupudzi, M., Gunson, B., Gitahi, J., Gocher, D., Batham, S., Pateman, H., Desmennu, S., Conder, J., Clement, D., Gallagher, S., Orpe, J., Chan, P., Currie, L., O'Donohoe, L., Oblak, M., Morgan, L., Quinn, M., Amey, I., Baird, Y., Cotterill, D., Cumlat, L., Winter, L., Greer, S., Spurdle, K., Allison, J., Dyer, S., Sweeting, H., Kordula, J., Aiba, Y., Nakamura, H., Abiru, S., Nagaoka, S., Komori, A., Yatsuhashi, H., Ishibashi, H., Ito, M., Kawai, Y., Kohn, S. -S., Gervais, O., Migita, K., Katsushima, S., Naganuma, A., Sugi, K., Komatsu, T., Mannami, T., Matsushita, K., Yoshizawa, K., Makita, F., Nikami, T., Nishimura, H., Kouno, H., Ota, H., Komura, T., Nakamura, Y., Shimada, M., Hirashima, N., Komeda, T., Ario, K., Nakamuta, M., Yamashita, T., Furuta, K., Kikuchi, M., Naeshiro, N., Takahashi, H., Mano, Y., Tsunematsu, S., Yabuuchi, I., Shimada, Y., Yamauchi, K., Sugimoto, R., Sakai, H., Mita, E., Koda, M., Tsuruta, S., Kamitsukasa, H., Sato, T., Masaki, N., Kobata, T., Fukushima, N., Higuchi, N., Ohara, Y., Muro, T., Takesaki, E., Takaki, H., Yamamoto, T., Kato, M., Nagaoki, Y., Hayashi, S., Ishida, J., Watanabe, Y., Kobayashi, M., Koga, M., Saoshiro, T., Yagura, M., Hirata, K., Takikawa, H., Ohira, H., Zeniya, M., Abe, M., Onji, M., Kaneko, S., Honda, M., Arai, K., Arinaga-Hino, T., Hashimoto, E., Taniai, M., Umemura, T., Joshita, S., Nakao, K., Ichikawa, T., Shibata, H., Yamagiwa, S., Seike, M., Honda, K., Sakisaka, S., Takeyama, Y., Harada, M., Senju, M., Yokosuka, O., Kanda, T., Ueno, Y., Kikuchi, K., Ebinuma, H., Himoto, T., Yasunami, M., Murata, K., Mizokami, M., Shimoda, S., Miyake, Y., Takaki, A., Yamamoto, K., Hirano, K., Ichida, T., Ido, A., Tsubouchi, H., Chayama, K., Harada, K., Nakanuma, Y., Maehara, Y., Taketomi, A., Shirabe, K., Soejima, Y., Mori, A., Yagi, S., Uemoto, S., Tanaka, T., Yamashiki, N., Tamura, S., Sugawara, Y., Kokudo, N., Carbone, M., Cardamone, G., Duga, S., Gershwin, M. E., Seldin, M. F., Invernizzi, P., Donato F., Fagiuoli S., Miele L. (ORCID:0000-0003-3464-0068), and Vinci M.

Abstract

Background & Aims: Genome-wide association studies in primary biliary cholangitis (PBC) have failed to find X chromosome (chrX) variants associated with the disease. Here, we specifically explore the chrX contribution to PBC, a sexually dimorphic complex autoimmune disease. Methods: We performed a chrX-wide association study, including genotype data from 5 genome-wide association studies (from Italy, United Kingdom, Canada, China, and Japan; 5244 case patients and 11,875 control individuals). Results: Single-marker association analyses found approximately 100 loci displaying P < 5 × 10–4, with the most significant being a signal within the OTUD5 gene (rs3027490; P = 4.80 × 10–6; odds ratio [OR], 1.39; 95% confidence interval [CI], 1.028–1.88; Japanese cohort). Although the transethnic meta-analysis evidenced only a suggestive signal (rs2239452, mapping within the PIM2 gene; OR, 1.17; 95% CI, 1.09–1.26; P = 9.93 × 10–8), the population-specific meta-analysis showed a genome-wide significant locus in East Asian individuals pointing to the same region (rs7059064, mapping within the GRIPAP1 gene; P = 6.2 × 10–9; OR, 1.33; 95% CI, 1.21–1.46). Indeed, rs7059064 tags a unique linkage disequilibrium block including 7 genes: TIMM17B, PQBP1, PIM2, SLC35A2, OTUD5, KCND1, and GRIPAP1, as well as a superenhancer (GH0XJ048933 within OTUD5) targeting all these genes. GH0XJ048933 is also predicted to target FOXP3, the main T-regulatory cell lineage specification factor. Consistently, OTUD5 and FOXP3 RNA levels were up-regulated in PBC case patients (1.75- and 1.64-fold, respectively). Conclusions: This work represents the first comprehensive study, to our knowledge, of the chrX contribution to the genetics of an autoimmune liver disease and shows a novel PBC-related genome-wide significant locus.

Published
2021

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