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Global terrestrial models currently predict that the Amazon rainforest will continue to act as a carbon sink in the future, primarily owing to the rising atmospheric carbon dioxide (CO2) concentration. Soil phosphorus impoverishment in parts of the Amazon basin largely controls its functioning, but the role of phosphorus availability has not been considered in global model ensembles—for example, during the Fifth Climate Model Intercomparison Project. Here we simulate the planned free-air CO2 enrichment experiment AmazonFACE with an ensemble of 14 terrestrial ecosystem models. We show that phosphorus availability reduces the projected CO2-induced biomass carbon growth by about 50% to 79 ± 63 g C m−2 yr−1 over 15 years compared to estimates from carbon and carbon–nitrogen models. Our results suggest that the resilience of the region to climate change may be much less than previously assumed. Variation in the biomass carbon response among the phosphorus-enabled models is considerable, ranging from 5 to 140 g C m−2 yr−1, owing to the contrasting plant phosphorus use and acquisition strategies considered among the models. The Amazon forest response thus depends on the interactions and relative contributions of the phosphorus acquisition and use strategies across individuals, and to what extent these processes can be upregulated under elevated CO2.

Extracellular enzymes (EE) play a vital role in soil nutrient cycling and thus affect terrestrial ecosystem functioning. Yet the drivers that regulate microbial activity, and therefore EE activity, remain under debate. In this study we investigate the temporal variation of soil EE in a tropical terra-firme forest. We found that EE activity peaked during the drier season in association with increased leaf litterfall, which was also reflected in negative relationships between EE activities and precipitation. Soil nutrients were weakly related to EE activities, although extractable N was related to EE activities in the top 5 cm of the soil. These results suggest that soil EE activity is synchronized with precipitation-driven substrate inputs and depends on the availability of N. Our results further indicate high investments in P acquisition, with a higher microbial N demand in the month before the onset of the drier season, shifting to higher P demand towards the end of the drier season. These seasonal fluctuations in the potential acquisition of essential resources imply dynamic shifts in microbial activity in coordination with climate seasonality and resource limitation of central-eastern Amazon forests.

Purpose The tropical phosphorus cycle and its relation to soil phosphorus (P) availability are a major uncertainty in projections of forest productivity. In highly weathered soils with low P concentrations, plant and microbial communities depend on abiotic and biotic processes to acquire P. We explored the seasonality and relative importance of drivers controlling the fluctuation of common P pools via processes such as litter production and decomposition, and soil phosphatase activity. Methods We analyzed intra-annual variation of tropical soil phosphorus pools using a modified Hedley sequential fractionation scheme. In addition, we measured litterfall, the mobilization of P from litter and soil extracellular phosphatase enzyme activity and tested their relation to fluctuations in P- fractions. Results Our results showed clear patterns of seasonal variability of soil P fractions during the year. We found that modeled P released during litter decomposition was positively related to change in organic P fractions, while net change in organic P fractions was negatively related to phosphatase activities in the top 5 cm. Conclusion We conclude that input of P by litter decomposition and potential soil extracellular phosphatase activity are the two main factors related to seasonal soil P fluctuations, and therefore the P economy in P impoverished soils. Organic soil P followed a clear seasonal pattern, indicating tight cycling of the nutrient, while reinforcing the importance of studying soil P as an integrated dynamic system in a tropical forest context.

Background and aims Ancient Amazon soils are characterised by low concentrations of soil phosphorus (P). Therefore, it is hypothesised that plants may invest a substantial proportion of their resources belowground to adjust their P-uptake strategies, including root morphological, physiological (phosphatase enzyme activities) and biotic (arbuscular mycorrhizal (AM) associations) adaptations. Since these strategies are energy demanding, we hypothesise that trade-offs between morphological traits and root phosphatase exudation and symbiotic associations would occur. Specifically, we expected that plants which invest in finer roots, and therefore have greater ability to explore large soil volumes, would have a high investment in physiological adaptations such as enhanced phosphatase production. In contrast, we expected that plants with predominantly thicker roots would invest more in symbiotic associations, in which carbon is traded for P acquired from AM fungal communities. Methods We collected absorptive roots (<2 mm diameter) from a lowland Central Amazon forest near Manaus, Brazil. We measured fine root diameter, specific root length (SRL), specific root area (SRA), root tissue density (RTD), root phosphatase activity (APase) and arbuscular mycorrhizal (AM) fungi colonisation. Results Root morphological traits were related to APase activity, with higher APase activity in roots with higher SRL and SRA but lower RTD. However, the degree of AM colonisation was not related to any measured root morphological trait. Conclusions Fine absorptive roots likely benefit from having low RTD, high SRL, SRA and APase exudation to acquire P efficiently. However, because AM colonisation was not related to root morphology, we suggest that investment in multiple P-uptake strategies is required for maintaining productivity in Central Amazon forests.

• Soil nutrient availability can strongly affect root traits. In tropical forests, phosphorus (P) is often considered the main limiting nutrient for plants. However, support for the P paradigm is limited, and N and cations might also control tropical forests functioning. • We used a large-scale experiment to determine how the factorial addition of nitrogen (N), P and cations affected root productivity and traits related to nutrient acquisition strategies (morphological traits, phosphatase activity, arbuscular mycorrhizal colonisation and nutrient contents) in a primary rainforest growing on low-fertility soils in Central Amazonia after 1 yr of fertilisation. • Multiple root traits and productivity were affected. Phosphorus additions increased annual root productivity and root diameter, but decreased root phosphatase activity. Cation additions increased root productivity at certain times of year, also increasing root diameter and mycorrhizal colonisation. P and cation additions increased their element concentrations in root tissues. No responses were detected with N addition. • Here we showed that rock-derived nutrients determined root functioning in low-fertility Amazonian soils, demonstrating not only the hypothesised importance of P, but also highlighting the role of cations. The changes in fine root traits and productivity indicated that even slow-growing tropical rainforests can respond rapidly to changes in resource availability.

Purpose Large parts of the Amazon rainforest grow on weathered soils depleted in phosphorus and rock-derived cations. We tested the hypothesis that in this ecosystem, fine roots stimulate decomposition and nutrient release from leaf litter biochemically by releasing enzymes, and by exuding labile carbon stimulating microbial decomposers. Methods We monitored leaf litter decomposition in a Central Amazon tropical rainforest, where fine roots were either present or excluded, over 188 days and added labile carbon substrates (glucose and citric acid) in a fully factorial design. We tracked litter mass loss, remaining carbon, nitrogen, phosphorus and cation concentrations, extracellular enzyme activity and microbial carbon and nutrient concentrations. Results Fine root presence did not affect litter mass loss but significantly increased the loss of phosphorus and cations from leaf litter. In the presence of fine roots, acid phosphatase activity was 43.2% higher, while neither microbial stoichiometry, nor extracellular enzyme activities targeting carbon- and nitrogen-containing compounds changed. Glucose additions increased phosphorus loss from litter when fine roots were present, and enhanced phosphatase activity in root exclusions. Citric acid additions reduced litter mass loss, microbial biomass nitrogen and phosphorus, regardless of fine root presence or exclusion. Conclusions We conclude that plant roots release significant amounts of acid phosphatases into the litter layer and mobilize phosphorus without affecting litter mass loss. Our results further indicate that added labile carbon inputs ( i.e . glucose) can stimulate acid phosphatase production by microbial decomposers, highlighting the potential importance of plant-microbial feedbacks in tropical forest ecosystems.

The response of lowland tropical forest on highly weathered P-limited soils to changes in atmospheric composition is likely to be regulated by P-availability from litter and soil. Our aim was to gain insight into possible mechanisms that may affect P-availability and C sequestration. We collected litter and soil samples along a transect in Central Amazonia. We examined litter and soil properties, determined C, N, and P contents and performed density fractionation to obtain organic-C, -N and -P fractions. To assess microbial demand and mineralization, we analysed C, N and P associated enzyme activities. The litter layer has an estimated turnover time of about one year or less and C, N and P contents were much smaller than in the mineral soil. Total soil C, N and P decreased with depth, while P was depleted relatively more. Most organic-C, -N and -P is stabilized by adsorption onto mineral surfaces. Non-adsorbed organic matter fractions were depleted of P. Nearly all organic-P appeared to be present in the mineral-associated fractions from where it, despite high phosphatase activities, appears not to be available for mineralization. Enzyme activities revealed that microbial C-demand in litter is higher than in soil, while P-demand was relatively high in soil and increased with depth. Hypothesized higher phosphatase activity under increasing atmospheric CO2 concentrations may probably not alleviate P-limitation in these terra firme forests due to the limited availability of mineralizable organic-P.

The tropical phosphorus cycle and its relation to soil phosphorus (P) availability are a major uncertainty in projections of forest productivity. In highly weathered soils with low P concentrations, plant and microbial communities depend on abiotic and biotic processes to acquire P. We explored the seasonality and relative importance of drivers controlling the fluctuation of common P pools via processes such as litter production and decomposition, and soil phosphatase activity. We analyzed intra-annual variation of tropical soil phosphorus pools using a modified Hedley sequential fractionation scheme. In addition, we measured litterfall, the mobilization of P from litter and soil extracellular phosphatase enzyme activity and tested their relation to fluctuations in P- fractions. Our results showed clear patterns of seasonal variability of soil P fractions during the year. We found that modeled P released during litter decomposition was positively related to change in organic P fractions, while net change in organic P fractions was negatively related to phosphatase activities in the top 5 cm. We conclude that input of P by litter decomposition and potential soil extracellular phosphatase activity are the two main factors related to seasonal soil P fluctuations, and therefore the P economy in P impoverished soils. Organic soil P followed a clear seasonal pattern, indicating tight cycling of the nutrient, while reinforcing the importance of studying soil P as an integrated dynamic system in a tropical forest context.