Direct evidence for phosphorus limitation on Amazon forest productivity.

The productivity of rainforests growing on highly weathered tropical soils is expected to be limited by phosphorus availability1. Yet, controlled fertilization experiments have been unable to demonstrate a dominant role for phosphorus in controlling tropical forest net primary productivity. Recent syntheses have demonstrated that responses to nitrogen addition are as large as to phosphorus2, and adaptations to low phosphorus availability appear to enable net primary productivity to be maintained across major soil phosphorus gradients3. Thus, the extent to which phosphorus availability limits tropical forest productivity is highly uncertain. The majority of the Amazonia, however, is characterized by soils that are more depleted in phosphorus than those in which most tropical fertilization experiments have taken place2. Thus, we established a phosphorus, nitrogen and base cation addition experiment in an old growth Amazon rainforest, with a low soil phosphorus content that is representative of approximately 60% of the Amazon basin. Here we show that net primary productivity increased exclusively with phosphorus addition. After 2 years, strong responses were observed in fine root (+29%) and canopy productivity (+19%), but not stem growth. The direct evidence of phosphorus limitation of net primary productivity suggests that phosphorus availability may restrict Amazon forest responses to CO2 fertilization4, with major implications for future carbon sequestration and forest resilience to climate change.

In situ short-term responses of Amazonian understory plants to elevated CO2.

The response of plants to increasing atmospheric CO2 depends on the ecological context where the plants are found. Several experiments with elevated CO2 (eCO2) have been done worldwide, but the Amazonian forest understory has been neglected. As the central Amazon is limited by light and phosphorus, understanding how understory responds to eCO2 is important for foreseeing how the forest will function in the future. In the understory of a natural forest in the Central Amazon, we installed four open-top chambers as control replicates and another four under eCO2 (+250 ppm above ambient levels). Under eCO2, we observed increases in carbon assimilation rate (67%), maximum electron transport rate (19%), quantum yield (56%), and water use efficiency (78%). We also detected an increase in leaf area (51%) and stem diameter increment (65%). Central Amazon understory responded positively to eCO2 by increasing their ability to capture and use light and the extra primary productivity was allocated to supporting more leaf and conducting tissues. The increment in leaf area while maintaining transpiration rates suggests that the understory will increase its contribution to evapotranspiration. Therefore, this forest might be less resistant in the future to extreme drought, as no reduction in transpiration rates were detected.

Tracking the impacts of El Niño drought and fire in human-modified Amazonian forests.

With humanity facing an unprecedented climate crisis, the conservation of tropical forests has never been so important - their vast terrestrial carbon stocks can be turned into emissions by climatic and human disturbances. However, the duration of these effects is poorly understood, and it is unclear whether impacts are amplified in forests with a history of previous human disturbance. Here, we focus on the Amazonian epicenter of the 2015-16 El Niño, a region that encompasses 1.2% of the Brazilian Amazon. We quantify, at high temporal resolution, the impacts of an extreme El Niño (EN) drought and extensive forest fires on plant mortality and carbon loss in undisturbed and human-modified forests. Mortality remained higher than pre-El Niño levels for 36 mo in EN-drought-affected forests and for 30 mo in EN-fire-affected forests. In EN-fire-affected forests, human disturbance significantly increased plant mortality. Our investigation of the ecological and physiological predictors of tree mortality showed that trees with lower wood density, bark thickness and leaf nitrogen content, as well as those that experienced greater fire intensity, were more vulnerable. Across the region, the 2015-16 El Niño led to the death of an estimated 2.5 ± 0.3 billion stems, resulting in emissions of 495 ± 94 Tg CO2 Three years after the El Niño, plant growth and recruitment had offset only 37% of emissions. Our results show that limiting forest disturbance will not only help maintain carbon stocks, but will also maximize the resistance of Amazonian forests if fires do occur.

Fine root dynamics across pantropical rainforest ecosystems.

Fine roots constitute a significant component of the net primary productivity (NPP) of forest ecosystems but are much less studied than aboveground NPP. Comparisons across sites and regions are also hampered by inconsistent methodologies, especially in tropical areas. Here, we present a novel dataset of fine root biomass, productivity, residence time, and allocation in tropical old-growth rainforest sites worldwide, measured using consistent methods, and examine how these variables are related to consistently determined soil and climatic characteristics. Our pantropical dataset spans intensive monitoring plots in lowland (wet, semi-deciduous, and deciduous) and montane tropical forests in South America, Africa, and Southeast Asia (n = 47). Large spatial variation in fine root dynamics was observed across montane and lowland forest types. In lowland forests, we found a strong positive linear relationship between fine root productivity and sand content, this relationship was even stronger when we considered the fractional allocation of total NPP to fine roots, demonstrating that understanding allocation adds explanatory power to understanding fine root productivity and total NPP. Fine root residence time was a function of multiple factors: soil sand content, soil pH, and maximum water deficit, with longest residence times in acidic, sandy, and water-stressed soils. In tropical montane forests, on the other hand, a different set of relationships prevailed, highlighting the very different nature of montane and lowland forest biomes. Root productivity was a strong positive linear function of mean annual temperature, root residence time was a strong positive function of soil nitrogen content in montane forests, and lastly decreasing soil P content increased allocation of productivity to fine roots. In contrast to the lowlands, environmental conditions were a better predictor for fine root productivity than for fractional allocation of total NPP to fine roots, suggesting that root productivity is a particularly strong driver of NPP allocation in tropical mountain regions.

Wood Nutrient-Water-Density Linkages Are Influenced by Both Species and Environment.

Tropical trees store a large amount of nutrients in their woody tissues, thus triggering the question of what the functional association of these elements with other wood traits is. Given the osmotic activity of mineral elements such as potassium, sodium, and calcium, these elements should be strong candidates in mediating the water storing capacity in tropical trees. We investigated the role of wood nutrients in facilitating wood water storage in trees by using branch samples from 48 tropical tree species in South America and examined their associations with wood density (ρ). Wood density varied from 316 kg/m3 in Peru plots, where the soil nutrient status is relatively higher, to 908 kg/m3 in Brazil plots, where the nutrient availability is lower. Phosphorus content in wood varied significantly between plots with lowest values found in French Guiana (1.2 mol/m3) and plots with highest values found in Peru (43.6 mol/m3). Conversely, potassium in woody tissues showed a significant cross-species variation with Minquartia guianensis in Brazil showing the lowest values (8.8 mol/m3) and with Neea divaricata in Peru having the highest values (114 mol/m3). We found that lower wood density trees store more water in their woody tissues with cations, especially potassium, having a positive association with water storage. Specific relationships between wood cation concentrations and stem water storage potential nevertheless depend on both species' identity and growing location. Tropical trees with increased water storage capacity show lower wood density and have an increased reliance on cations to regulate this reservoir. Our study highlights that cations play a more important role in tropical tree water relations than has previously been thought, with potassium being particularly important.

Tropical Tree Branch-Leaf Nutrient Scaling Relationships Vary With Sampling Location.

Bivariate relationships between plant tissue nutrient concentration have largely been studied across broad environmental scales regardless of their covariation with soil and climate. Comparing leaf and branch wood concentrations of C, Ca, K, Mg, N, Na, and P for trees growing in tropical forests in Amazonia and Australia we found that the concentrations of most elements varied with sampling location, but with foliar and branch woody tissues varying from site to site in different ways. Using a Mixed Effect Model (MEM) approach it was further found that relationships between branch and leaf concentrations within individual plots differed in terms of both slope and/or significance to the ordinary least squares (OLS) estimates for most elements. Specifically, using MEM we found that within plots only K and Mg were correlated across organs, but with the K cross-organ intercept estimates varying significantly between sites. MEM analyses further showed that within-plot wood density variations were also negatively related to wood K and Na, suggesting a potentially important role for these cations in water transport and/or storage in woody tissues. The OLS method could not detect significant correlations in any of the above cases. By contrast, although Ca, N, and P leaf and wood tissue concentrations showed similar patterns when individual elements were compared across sites, MEM analyses suggested no consistent association within sites. Thus, for all these three elements, strong within-tree scaling relationships were inferred when data were analyzed across sites using OLS, even though there was no relationship within individual sites. Thus (as for Ca, N, and P) not only can a pooling of data across sites result in trait (co)variations attributable to the environment potentially being incorrectly attributed solely to the species and/or individual (the so-called "ecological fallacy"), but in some cases (as was found here for K and Na) the opposite can also sometimes occur with significant within-site covariations being obscured by large site-site variations. We refer to the latter phenomenon as "environmental obfuscation."

Seasonal variations in soil water in two woodland savannas of central Brazil with different fire history.

Changes in soil water content were determined in two cerrado (sensu stricto) areas with contrasting fire history and woody vegetation density. The study was undertaken near Brasília, Brazil, from 1999 to 2001. Soil water content was measured with a neutron probe in three access tubes per site to a depth of 4.7 m. One site has been protected from fire for more than 30 years and, as a consequence, has a high density of woody plants. The other site had been frequently burned, and has a high herbaceous vegetation density and less woody vegetation. Soil water uptake patterns were strongly seasonal, and despite similarities in hydrological processes, the protected area systematically used more water than the burned area. Three temporarily contiguous patterns of water absorption were differentiated, characterized by variation in the soil depth from which water was extracted. In the early dry season, vegetation used water from throughout the soil profile but with a slight preference for water in the upper soil layers. Toward the peak of the dry season, vegetation had used most or all available water from the surface to a depth of 1.7 m, but continued to extract water from greater depths. Following the first rains, all water used was from the recently wetted upper soil layers only. Evaporation rates were a linear function of soil water availability, indicating a strong coupling of atmospheric water demand and the physiological response of the vegetation.

Foliar trait contrasts between African forest and savanna trees: genetic versus environmental effects.

Variations in leaf mass per unit area (Ma) and foliar concentrations of N, P, C, K, Mg and Ca were determined for 365 trees growing in 23 plots along a West African precipitation gradient ranging from 0.29 to 1.62m a-1. Contrary to previous studies, no marked increase in Ma with declining precipitation was observed, but savanna tree foliar [N] tended to be higher at the drier sites (mass basis). Generally, Ma was slightly higher and [N] slightly lower for forest vs savanna trees with most of this difference attributable to differences in soil chemistry. No systematic variations in [P], [Mg] and [Ca] with precipitation or between trees of forest vs savanna stands were observed. We did, however, find a marked increase in foliar [K] of savanna trees as precipitation declined, with savanna trees also having a significantly lower [K] than those of nearby forest. These differences were not related to differences in soil nutrient status and were accompanied by systematic changes in [C] of opposite sign. We suggest an important but as yet unidentified role for K in the adaption of savanna species to periods of limited water availability; with foliar [K] being also an important factor differentiating tree species adapted to forest vs savanna soils within the 'zone of transition' of Western Africa.

Variations in Amazon forest productivity correlated with foliar nutrients and modelled rates of photosynthetic carbon supply.

The rate of above-ground woody biomass production, W(P), in some western Amazon forests exceeds those in the east by a factor of 2 or more. Underlying causes may include climate, soil nutrient limitations and species composition. In this modelling paper, we explore the implications of allowing key nutrients such as N and P to constrain the photosynthesis of Amazon forests, and also we examine the relationship between modelled rates of photosynthesis and the observed gradients in W(P). We use a model with current understanding of the underpinning biochemical processes as affected by nutrient availability to assess: (i) the degree to which observed spatial variations in foliar [N] and [P] across Amazonia affect stand-level photosynthesis; and (ii) how these variations in forest photosynthetic carbon acquisition relate to the observed geographical patterns of stem growth across the Amazon Basin. We find nutrient availability to exert a strong effect on photosynthetic carbon gain across the Basin and to be a likely important contributor to the observed gradient in W(P). Phosphorus emerges as more important than nitrogen in accounting for the observed variations in productivity. Implications of these findings are discussed in the context of future tropical forests under a changing climate.