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It is well documented that microbial biomass increases during winter in cold mountain or tundra ecosystems, but the cause and mechanism of such accumulation is unclear. Results from a grassland in Iceland demonstrated that the microbial biomass carbon (MBC, measured by the fumigation-extraction method) increased in winter, while microbial DNA content remained constant. We thus hypothesized that this accumulation of microbial biomass during the cold season is driven by the decrease in temperature, that increases the carbon storage of individual cells, but not by an increase in microbial cell numbers.To test this hypothesis, we conducted a laboratory incubation experiment with soils from a grassland in Iceland sampled before the onset of winter in early October (around 9 °C). We then exposed the soils to decreasing temperatures (0.5 °C, 3 °C, 6 °C and 9 °C) over five months. We analyzed microbial biomass carbon (MBC) and quantified the DNA content. Over the course of five months, we found higher MBC values at cool temperatures compared to warm conditions. As expected, cooling did not affect the DNA content, leading to a significantly higher MBC to DNA ratio when soils were incubated at 0.5 °C compared to 9 °C. This indicates that numbers of microbial cells did not change across temperatures, but that microbes at lower temperatures stored more carbon. We also found similar patterns in soils collected at different time points in the field. Furthermore, we estimated microbial DNA production, i.e., growth rates, by measuring the incorporation of 18O from labelled water into DNA. We observed lower microbial growth rates under field conditions in winter, indicating that increasing biomass carbon was not due to increased growth and that growth and turnover was balanced at all temperatures. Instead, we suggest that carbon uptake (which was decreased at lower temperatures) was less affected by cold temperatures than growth, so that microbial carbon could accumulate. We also verified this pattern in growth and carbon uptake rates with decreasing temperatures in the laboratory incubation experiment.Decreasing growth (cell division) and turnover rates with decreasing temperatures, at a lower but sustained carbon uptake rate, suggest that the cell size of soil microorganisms may increase when exposed to cooling. We will show and discuss first results from measurements with a suspended microchannel resonator (SMR), that, together with microscopic imaging allows to assess the mass (size) of individual cells of microorganisms at different temperatures.

High-latitude soils are particularly vulnerable to temperature-driven C losses and may contribute substantially to the increasing atmospheric CO2 concentrations. The magnitude of their contribution is, however, uncertain, and largely dependent on the interactions between C and nitrogen (N) biogeochemical cycles, soil microbial activities and the feedbacks between plants and soil microbes. Warming may cause a particularly pronounced acceleration of soil N transformation in N-poor cold regions. The consequent alleviation of plant N limitations in cold ecosystems may increase plant productivity and C inputs to the soil, compensating the expected soil C loss, at least partially. Alternatively, warming may desynchronize or unbalance the intimate coupling between microbial N mineralization and vegetation N uptake, leading to potential soil N loss, but also higher soil C losses. We aimed to elucidate potential mechanisms of ecosystem N losses in subarctic grasslands by determining the effects of soil warming on the seasonal patterns of plant N acquisition and microbial net N immobilization. For this, we performed a seasonal isotope tracing experiment using a mix of 15N-labelled amino-acids along soil temperature gradients in geothermal systems in Iceland.Soil microbial biomass acted as a temporal reservoir of N by increasing N immobilization particularly during unfavorable winter periods for vegetation, likely due to the alleviated microbial C limitation. However, soil warming exacerbated microbial C limitation and decreased the N storage capacity of soil microbes during snowmelt periods. As a result, a higher proportion of N remained in the extractable soil fraction susceptible to leaching losses. , however, this increased plant N uptake did not compensate for the lower microbial biomass N storage, leading to ecosystem N losses. Our results highlight the relevant role of soil microbes to safely store and immobilize N when plants do not need it and to release N when plants require it. Warming can weaken this particularly important soil microbial function in cold regions, leading to substantial ecosystem N and fertility losses, which may also promote irreversible soil C losses in these ecosystems.

Soil nutrient availability is a key constraint on tropical forest growth. On highly weathered soils, intact forests' carbon-nutrient cycles have developed over pedogenic time scales, leading to tight nutrient cycling of some depleted elements, such as phosphorus. Ecosystem nutrient recycling and plant nutrient limitation are predominant in lowland Amazonia, controlling the forests' response to disturbance and climate change. Deforestation, biomass removal, and fire lead to the loss of carbon and nutrients previously stored in vegetation, potentially enforcing nutrient limitation and reducing carbon storage in regrowing forests. Here, we employ the process-based terrestrial biosphere model QUINCY (Thum et al., 2019), coupled with the microbial-explicit soil model JSM (Yu et al., 2020) to simulate carbon and nutrient cycling rates at intact Amazonian forest sites, which span a natural gradient of 30 to 727 mg phosphorus g dry soil-1, and from 2 to 81% clay. The model QUINCY-JSM accounts for dynamic plant carbon investment in growth and nutrient acquisition, and microbial-explicit growth, turnover, and nutrient cycling. We confront the ecosystems with historical changes in atmospheric CO2 and climate, and simulate an experimental harvest of the entire forest stand to assess the consequences of that nutrient loss on the regrowing forest stand. Simulations of the Amazon forest sites are in good agreement with forest census data on vegetation carbon dynamics. Biologically-driven nutrient mineralization represents the main source of nutrients for plants, with negligible contribution of inorganic nutrient cycling in highly weathered sites. After experimental deforestation, we find that the inorganic nutrient supply is insufficient and restricts forest growth, leading to lower vegetation biomass equilibrium after deforestation. Our simulations suggest that forest degradation may occur through biomass removal in tropical forests.Thum, T., Caldararu, S., Engel, J., Kern, M., Pallandt, M., Schnur, R., et al. (2019). A new model of the coupled carbon, nitrogen, and phosphorus cycles in the terrestrial biosphere (QUINCY v1.0; revision 1996). Geoscientific Model Development, 12(11), 4781–4802. https://doi.org/10.5194/gmd-12-4781-2019Yu, L., Ahrens, B., Wutzler, T., Schrumpf, M., & Zaehle, S. (2020). Jena Soil Model (JSM v1.0; Revision 1934): A microbial soil organic carbon model integrated with nitrogen and phosphorus processes. Geoscientific Model Development, 13(2), 783–803. https://doi.org/10.5194/gmd-13-783-202

Arbuscular mycorrhizal fungi (AMF) form mutualistic associations with roughly 70% of vascular plant species, supporting the nutrient acquisition of their host plants and deriving carbon in return. AMF and plant communities are linked to each other by host-specificity and the ecological selection of favorable nutrient and carbon trading strategies. Changing soil nutrient availabilities can affect both plant and AMF communities directly and also indirectly via the response of their partners. We aimed to elucidate the combined response of AMF (belowground) and plant (aboveground) community compositions to changing soil nutrient availabilities.We sampled soil and roots from a long-term nutrient deficiency experimental grassland in Admont (Styria, Austria). The grassland plots have been fertilized with different combinations of nitrogen (N), phosphorus (P), and potassium (K) over 70 years. Aboveground biomass cuts were removed three times each year, leading to long-term deficiencies of nutrients not replaced by fertilizers. Soil and root AMF community compositions were measured by DNA and RNA amplicon sequencing of the 18S rRNA gene. In addition, we assessed the plant community composition of the sampled roots by amplicon sequencing of the chloroplast rbcL (RuBisCo large subunit) gene region, and visually recorded the plant community composition on each investigated plot.Our results demonstrate that N and P deficiencies influenced soil AMF community composition, whereas K deficiency had a major impact on root AMF community composition. Interestingly, the plant community composition was affected by N and P, similar to the soil AMF community composition. Both, soil and root AMF community compositions were significantly correlated to plant community composition across all treatments, the correlation was however stronger for soil AMF communities (R2 = 0.55, p< 0.001). By using bipartite network analysis, we identified several fungus-plant pairs that responded consistently to treatments.Our results indicate that the response of grasslands to nutrient deficiencies is potentially driven by strong feedbacks between plant and belowground AMF community compositions. We here demonstrate that the known interactions between grassland plants and AMF - which are often investigated from a single plant or monoculture perspective - are major drivers of how diverse plant community compositions will respond to environmental change, such as fertilization. In conclusion, considering the ecology of the subsurface AMF communities may strongly benefit our understanding of plant communities in a future environment.

The terrestrial soil organic matter (SOM) pool size depends on the balance between SOM formation and stabilization of decomposing plant litter relative to mineralization as CO2. Decomposition and mineralization processes are to large extents mediated by microbial decomposer communities. In addition, labile fractions released by decomposing litter can be stabilized in mineral associations (MAOM). High latitude ecosystems are particularly affected by global warming. Increasing temperatures can stimulate litter decomposition and increase nutrient mineralization, thereby increasing nutrient (e.g. nitrogen) availability for plants allowing higher productivity and subsequent plant organic matter inputs. On the other hand, if warming accelerates SOM decomposition stronger than formation processes it can cause large carbon and nutrient losses.Tracing 13C/15N labelled above-ground litter we aimed to disentangle if soil warming changes the balance between litter derived SOM formation through microbial communities, over particulate organic matter (POM) and MAOM stabilization across a decadal geothermal soil warming gradient (from ambient up to +5 °C) in a grassland in Iceland. In addition, we added three levels of inorganic N to disentangle potential direct warming from indirect (over plant feedbacks) effects of increased warming on SOM dynamics. We found that over the course of two years warming accelerated litter decomposition rates and litter-derived carbon turnover by the microbial community, and we could recover more litter-derived carbon recovered in particulate organic matter (POM) nor MAOM. Nitrogen additions triggered a faster decomposition of structural above-ground litter compounds, but did not influence carbon turnover in different soil fractions. On the other, hand we found that overall the absolute amount of soil carbon decreased in response to warming. We therefore conclude that direct warming not only increased SOM formation, but also decomposition and mineralization processes, and – at least in the studied grassland – plant inputs may not have counterbalanced warming induced losses.

In early stages of forest succession plants have a high nutrient demand, but it is still a matter of debate if regrowth success of pioneer species is related to plant functional traits favoring fast soil colonization and nutrient acquisition. In general, we would expect trade-offs between plant growth performance and fine root morphological properties in association with different plant life-history strategies. Hence, we hypothesized that fast growing plants should have a more efficient root system that allows them to outcompete slow-growing neighbors in a resource-limited environment.To test our hypothesis we monitored plant successional growth dynamics in a tropical lowland rainforest reforestation experiment conducted in southwest Costa Rica. We collected absorptive roots (

There is increasing evidence to suggest that soil nutrient availability can limit the carbon sink capacity of forests, a particularly relevant issue considering today's changing climate. This question is especially important in the tropics, where most part of the Earth's plant biomass is stored. To assess whether tropical forest growth is limited by soil nutrients and to explore N and P limitations, we analyzed stem growth and foliar elemental composition of the five stem widest trees per plot at two sites in French Guiana after 3 years of nitrogen (N), phosphorus (P), and N + P addition. We also compared the results between potential N-fixer and non-N-fixer species. We found a positive effect of N fertilization on stem growth and foliar N, as well as a positive effect of P fertilization on stem growth, foliar N, and foliar P. Potential N-fixing species had greater stem growth, greater foliar N, and greater foliar P concentrations than non-N-fixers. In terms of growth, there was a negative interaction between N-fixer status, N + P, and P fertilization, but no interaction with N fertilization. Because N-fixing plants do not show to be completely N saturated, we do not anticipate N providing from N-fixing plants would supply non-N-fixers. Although the soil-age hypothesis only anticipates P limitation in highly weathered systems, our results for stem growth and foliar elemental composition indicate the existence of considerable N and P co-limitation, which is alleviated in N-fixing plants. The evidence suggests that certain mechanisms invest in N to obtain the scarce P through soil phosphatases, which potentially contributes to the N limitation detected by this study.

In temperate soil systems, microbial biomass often increases during winter and decreases again in spring. This build-up and release of microbial carbon could potentially lead to a stabilization of soil carbon during winter times. Whether this increase is caused by changes in microbial physiology, in community composition or by changed substrate allocation within microbes or communities is unclear. In a laboratory incubation study, we looked into microbial respiration and growth, as well as microbial glucose uptake and carbon resource partitioning in response to cooling. Soils taken from a temperate forest and an agricultural system in October 2020, were cooled down from field temperature of 11 °C to 1 °C. We determined microbial growth using 18O-incorporation into DNA immediately after cooling and after an acclimation phase of 7 days; in addition, we traced 13C-labelled glucose into microbial biomass, CO2 respired from the soil, and into microbial phospholipid fatty acids (PLFAs). Our results show that the studied soil microbial communities responded immediately to soil cooling. Independent of soil type and acclimation period, total respiration, as well as 18O based growth, and thus cell division were strongly reduced when soils were cooled from 11 °C to 1 °C, while glucose uptake and glucose-derived respiration were unchanged. We found that microbes increased the investment of glucose-derived carbon in unsaturated phospholipid fatty acids at colder temperatures. Since unsaturated fatty acids retain fluidity at lower temperatures compared to saturated fatty acids, this could be interpreted as a precaution to reduced temperatures. Together with the maintained glucose uptake and reduced cell division, our findings show an immediate response of soil microorganisms to soil cooling, potentially to prepare for freeze-thaw events. The discrepancy between C uptake and cell division, further hints at a mechanism that could explain previously observed high microbial biomass carbon in temperate soils in winter.

Nutrient cycles are tightly linked to the carbon cycle in tropical forests, controlling its responses to environmental change such as elevated CO2 concentrations (eCO2). In tropical wet forests of the Amazon, plants tend to grow slower in low fertility soils, while their relative investments in nutrient acquisition are likely higher due to costly mechanisms of nutrient mobilization. In low fertility soils where weatherable minerals have been depleted, decomposition and mineralization of soil organic matter and plant litter by free-living microorganisms may represent the dominant nutrient source for plants. The availability of soil nutrients for plants is a key constraint on tropical forest growth under future climate change. However, soil microbial communities not only drive the mineralization, but require nutrients for growth and biomass production themselves, and therefore from a plant’s perspective, soil microbial communities may act as a source or sink of nutrients under eCO2.Here, we employ the process-based terrestrial biosphere model QUINCY (Thum et al., 2019), coupled to the microbial-explicit soil model JSM (Yu et al., 2020) to model shifts in coupled nutrient and carbon cycling rates at field sites with differing soil fertility in the Amazon forest sites, and to confront the modeled ecosystems with elevated CO2. QUINCY-JSM reflects our current understanding of governing carbon and nutrient cycling feedbacks, allowing for dynamic plant carbon investment in growth and nutrient acquisition, and microbial-explicit growth, turnover, and nutrient cycling. We compiled a unique dataset of forest growth, soil and litter chemistry, as well as microbial growth and stoichiometry from a set of Amazon forest plots that cover a large gradient in soil fertility. Microbial stoichiometry and soil texture data are used to calibrate QUINCY-JSM, and data on forest aboveground growth, microbial growth, litter chemistry, and soil carbon and nutrients are used for model evaluation. We test the hypothesis that the contribution of microbial-driven nutrient mineralization to the nutrient supply of plants increases with lowering soil fertility and explore the soil microbial-induced nutrient feedback to eCO2-induced carbon sequestration in wet lowland Amazon forest sites. We examine the consequences of model assumptions on the conditions in space and time under which soil microorganisms alleviate or enforce the plants’ nutrient limitation under eCO2. Directly testable hypotheses for old-growth wet forests’ response to elevated CO2 in ecosystem-scale experiments like AmazonFACE are formulated.Thum, T., Caldararu, S., Engel, J., Kern, M., Pallandt, M., Schnur, R., et al. (2019). A new model of the coupled carbon, nitrogen, and phosphorus cycles in the terrestrial biosphere (QUINCY v1.0; revision 1996). Geoscientific Model Development, 12(11), 4781–4802. https://doi.org/10.5194/gmd-12-4781-2019Yu, L., Ahrens, B., Wutzler, T., Schrumpf, M., & Zaehle, S. (2020). Jena Soil Model (JSM v1.0; Revision 1934): A microbial soil organic carbon model integrated with nitrogen and phosphorus processes. Geoscientific Model Development, 13(2), 783–803. https://doi.org/10.5194/gmd-13-783-2020

Phosphorus (P) is one of the main limiting nutrients for forest productivity in Amazonia. To meet P needs, plants invest resources in different strategies which may increase their P-use efficiency, e.g., by resorbing P from senescing organs, or increase their P-acquisition efficiency, e.g., by acclimating fine root traits (architectural, morphological, physiological, and symbiotic). P-acquisition strategies can be categorized into foraging strategies related to the uptake of plant-available P or mining strategies related to the mobilization and uptake of less available forms of P. However, little is known about the effects of soil P on plant P-use and -acquisition strategies in Amazonia. Therefore, we have conducted a literature review and synthesized the current knowledge on the variation of different P-use and -acquisition strategies across soil P fertility gradients and their response to P fertilization in Amazonia and other tropical forests (Reichert et al., in press). We provide a conceptual framework on the distribution of these strategies in Amazonia and propose that, at the plant community level, foraging strategies (via fine roots and arbuscular mycorrhizas) are more prevalent and may contribute most for plant P uptake in soils with intermediate to high P availability, and leaf P resorption and mining strategies (via root exudation of acid phosphatases and organic acids) in soils with intermediate to low P availability (Reichert et al., in press). Here, we suggest that the investment in different P-acquisition strategies may be partially explained by the energy cost per unit P acquired. Based on the assumption that this cost varies with strategy and the form of P and its concentration in the soil (Raven et al., 2018), we have developed a stand-alone theoretical model to predict plant investments in P acquisition and test our conceptual framework. We constrain the model with field observations on forest growth and soil nutrients from sites in Amazonia and explore possible shifts in P-acquisition strategies along soil P fertility gradients.Raven JA, Lambers H, Smith SE, Westoby M. 2018. Costs of acquiring phosphorus by vascular land plants: patterns and implications for plant coexistence. New Phytologist 217(4): 1420-1427.Reichert T, Rammig A, Fuchslueger L, Lugli LF, Quesada CA, Fleischer K. In press. Plant phosphorus-use and -acquisition strategies in Amazonia. New Phytologist.

Background It is relatively unknown if and how seasonal fluctuations of tropical microbial activity affect soil nutrient availability. In tropical forests, nutrient economics are often considered to be centered around phosphorus, which might be a limiting factor to sustain crucial ecosystem processes, such as primary production and decomposition of organic material, thus in turn affecting microbial processes and associated nutrient dynamics of the forest ecosystem. Aims We investigate seasonal fluctuations in extracellular hydrolytic soil enzyme activities and soil nutrients and its relationship with precipitation and litterfall input, in a lowland tropical forest in the Central Amazon region. Methods We analyzed data obtained from monitoring microbial enzyme activity and nutrient dynamics in litter and soil and use stoichiometric enzyme theory and proportional vectors for assessing relative nutrient limitation throughout a year. Results Our results show that precipitation seasonality was driving leaf litterfall, which was subsequently synchronized with extracellular enzyme activities in soil, such that both litterfall and enzyme activities peaked during the dry season. Conclusions Our study indicates that soil extractable nutrient concentrations were positively related to microbial enzyme activities, which thus highlights the importance of soil microbial processes for nutrient cycling in this phosphorus limited ecosystem. Our results suggest that projected shifts in climate seasonality that result in longer and more pronounced dry seasons, might desynchronize seasonal patterns of aboveground nutrient input and belowground microbial activity, and thus leading to a decoupling of nutrient cycling in tropical forest ecosystems.

Long-term soil warming and nitrogen (N) availability have been shown to affect microbial biomass and community composition. Altered assimilation patterns of recent plant-derived C and changes in soil C stocks following warming as well as increased N availability are critical in mediating the direction and magnitude of these community shifts. A 13C pulse labelling experiment was done on a warming gradient in an Icelandic grassland (Sigurdsson et al. 2016), to investigate the role of recent plant-derived C and warming on the microbial community structure and size. We observed an overall increase of microbial 13C (e.g., root-exudate) uptake, while warming led to significant microbial biomass loss in all microbial groups. The increase of microbial 13C uptake with warming differed between microbial groups: an increase was only observed in the general and Gram-positive bacterial phospholipid fatty acid (PLFA) markers and in the PLFA and neutral lipid fatty acid (NLFA) markers of arbuscular mycorrhizal fungi (AMF). Nitrogen addition of 50 kg ha-1 y-1 for two years had no effect on the microbial uptake, microbial biomass or community composition, indicating that microbes were not N limited, and no plant-mediated N addition effects occurred. Additionally, we show that both warming and soil C depletion were responsible for the microbial biomass loss. Soil warming caused stronger loss in microbial groups with higher 13C uptake. In our experiment, warming caused a general reduction of microbial biomass, despite a relative increase in microbial 13C uptake, and altered microbial community composition. The warming effects on microbial biomass and community composition were partly mediated through soil C depletion with warming and changes in recent plant-derived C uptake patterns of the microbial community.

Global warming may lead to carbon transfers from soils to the atmosphere, yet this positive feedback to the climate system remains highly uncertain, especially in subsoils (Ilyina and Friedlingstein, 2016; Shi et al., 2018). Using natural geothermal soil warming gradients of up to +6.4 ∘C in subarctic grasslands (Sigurdsson et al., 2016), we show that soil organic carbon (SOC) stocks decline strongly and linearly with warming (−2.8 t ha−1 ∘C−1). Comparison of SOC stock changes following medium-term (5 and 10 years) and long-term (>50 years) warming revealed that all SOC stock reduction occurred within the first 5 years of warming, after which continued warming no longer reduced SOC stocks. This rapid equilibration of SOC observed in Andosol suggests a critical role for ecosystem adaptations to warming and could imply short-lived soil carbon–climate feedbacks. Our data further revealed that the soil C loss occurred in all aggregate size fractions and that SOC stock reduction was only visible in topsoil (0–10 cm). SOC stocks in subsoil (10–30 cm), where plant roots were absent, showed apparent conservation after >50 years of warming. The observed depth-dependent warming responses indicate that explicit vertical resolution is a prerequisite for global models to accurately project future SOC stocks for this soil type and should be investigated for soils with other mineralogies.

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.

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.

Climate change is altering the frequency and severity of drought events. Recent evidence indicates that drought may produce legacy effects on soil microbial communities. However, it is unclear whether precedent drought events lead to ecological memory formation, i.e., the capacity of past events to influence current ecosystem response trajectories. Here, we utilize a long-term field experiment in a mountain grassland in central Austria with an experimental layout comparing 10 years of recurrent drought events to a single drought event and ambient conditions. We show that recurrent droughts increase the dissimilarity of microbial communities compared to control and single drought events, and enhance soil multifunctionality during drought (calculated via measurements of potential enzymatic activities, soil nutrients, microbial biomass stoichiometry and belowground net primary productivity). Our results indicate that soil microbial community composition changes in concert with its functioning, with consequences for soil processes. The formation of ecological memory in soil under recurrent drought may enhance the resilience of ecosystem functioning against future drought events., Legacies of past ecological disturbances are expected but challenging to demonstrate. Here the authors report a 10-year field experiment in a mountain grassland that shows ecological memory of soil microbial community and functioning in response to recurrent drought.

Terrestrial biosphere models typically use the biochemical model of Farquhar, von Caemmerer and Berry (1980) to simulate photosynthesis, which requires accurate values of photosynthetic capacity of different biomes. However, data on tropical forests are sparse and highly variable due to the high species diversity, and it is still highly uncertain how these tropical forests respond to nutrient limitation in terms of C uptake. Tropical forests often grow on phosphorus (P)-poor soils and are, in general, assumed to be P- rather than nitrogen (N)-limited. However, the relevance of P as a control of photosynthetic capacity is still debated. Here, we provide a comprehensive dataset of vertical profiles of photosynthetic capacity and important leaf traits, including leaf N and P concentrations, from two three-year, large-scale nutrient addition experiments conducted in two tropical rainforests in French Guiana. These data present a unique source of information to further improve model representations of the roles of N, P, and other leaf nutrients, in photosynthesis in tropical forests. To further facilitate the use of our data in syntheses and model studies, we provide an elaborate list of ancillary data, including important soil properties and nutrients, along with the leaf data. As environmental drivers are key to improve our understanding of carbon (C)-nutrient cycle interactions, this comprehensive dataset will aid to further enhance our understanding of how nutrient availability interacts with C uptake in tropical forests. The data are available at DOI 10.5281/zenodo.4719242 (Verryckt, 2021).

In tropical forests, free-living Biological nitrogen (N) fixation (BNF) in soil and litter tends to decrease when substrate N concentrations increase, whereas increasing phosphorus (P) and molybdenum (Mo) soil and litter concentrations have been shown to stimulate free-living BNF rates. Yet, very few studies explored the effects of adding N, P, and Mo together in a single large-scale fertilization experiment, which would teach us which of these elements constrain or limit BNF activities. At two distinct forest sites in French Guiana, we performed a 3-year in situ nutrient addition study to explore the effects of N, P, and Mo additions on leaf litter and soil BNF. Additionally, we conducted a short-term laboratory study with the same nutrient addition treatments (+N, +N+P, +P, +Mo, and +P+Mo). We found that N additions alone suppressed litter free-living BNF in the field, but not in the short-term laboratory study, while litter free-living BNF remained unchanged in response to N+P additions. Additionally, we found that P and P+Mo additions stimulated BNF in leaf litter, both in the field and in the lab, while Mo alone yielded no changes. Soil BNF increased with P and P+Mo additions in only one of the field sites, while in the other site soil BNF increased with Mo and P+Mo additions. We concluded that increased substrate N concentrations suppress BNF. Moreover, both P and Mo have the potential to limit free-living BNF in these tropical forests, but the balance between P versus Mo limitation is determined by site-specific characteristics of nutrient supply and demand.

Purpose. The tropical phosphorus-cycle and its impacts on 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 biological and physical 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 variation of standard tropical soil phosphorus pools over one year. In addition, we measured litterfall, its decomposition rates and soil extracellular phosphatase enzyme activity and tested their relation to the fluctuations in P-fractions. Results. Our results show clear patterns of seasonal variability of soil P fractions during the year. We found that modeled P released during litter decomposition is 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 organic P forms by litter decomposition and organically produced phosphatases are the two main factors controlling seasonal soil P fluctuation, and therefore the P economy in P impoverished soils. Organic soil P follows 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.

Tropical forest ecosystems are important components of global carbon (C) and nutrient cycles. Many tropical rainforests grow on old and highly weathered soils depleted in phosphorus (P) and other rock-derived nutrients. While plants in such forests are usually P limited, it remains unclear if heterotrophic microbial communities are also limited by P or rather by C or energy. Elemental limitations of microorganisms in soil are often approached by measurements of changes in respiration rates or microbial biomass in response to additions of nutrients or carbon. However, it has been argued lately, that microbial growth rather than respiration or biomass should be used to assess microbial limitations. In this study we asked the question whether the growth of heterotrophic microbial communities in tropical soil is limited by available P or by C. We sampled soils along a topographic gradient (plateau, slope, bottom) differing in soil texture and total and available P concentrations from a highly weathered site in French Guiana. We incubated these soils in the laboratory with cellulose as a C source, phosphate (pH adjusted) and with a combination of both. We determined microbial growth by measuring the incorporation of 18O from labelled water into microbial DNA. In general, plateau soils were higher in microbial C, while bottom soils were higher in microbial P, leading to increased microbial C:P ratios in plateau soils compared to bottom soils. Microbial C, N and P did not respond to the addition of cellulose. Microbial P on the other hand was significantly increased by P additions, with no interactive effect between cellulose and P. Although microbial C was significantly higher in plateau soils, respiration rates were similar to those of bottom soils. This led to similar mass specific respiration rates in plateau and slope soils, with bottom soils being significantly higher. Moreover, we found that C and P addition increased mass specific respiration rates and both nutrient additions showed a positive interactive effect. Gross microbial growth rates were stimulated by P additions but were unresponsive to C additions alone. However, the addition of carbon further stimulated the effect of P on growth. The observed interactive effect of C and P additions on gross microbial growth rates suggests a co-limitation of microorganisms by C and P in highly weathered soils. We argue that co-limitation bears significant ecological advantages for microorganisms as it minimizes the investments in acquiring nutrients for growth.We further conclude that microorganisms in tropical soils are highly efficient in taking up and storing P from the environment. In our experiment, microbial P almost doubled in the six days after P addition, while microbial C was not enhanced. This also means that the microbes were not homeostatic with regard to their C:P ratios. Finally, our study demonstrates the importance of investigating gross microbial growth rates, rather than respiration or biomass, for inferring nutrient limitations.

Fertilization experiments provide insights into elemental imbalances in soil microbial communities and their consequences for soil nutrient cycling. By addition of selected nutrients, other nutrients become deficient and limiting for soil microorganisms as well as for plants. In this study we focused on microbial nitrogen (N) cycling in a long-term nutrient manipulation experiment. In many soils, the rate-limiting step in N cycling is depolymerization of high-molecular-weight nitrogen compounds (e.g., proteins) to oligomers (e.g., peptides) and monomers (e.g., amino acids) rather than the subsequent steps of mineralization (ammonification) and nitrification. The aim of our study was to determine whether nutrient deficiency directly or indirectly – via changes in plant carbon (C) inputs - affects soil microbial N processing.We collected soil samples from a fertilization experiment, established in 1946 on a hay meadow close to Admont (Styria, Austria). The field experiment consisted of a full factorial combination of inorganic N, P, and K fertilization and a control with no fertilizers. Furthermore, liming (Ca-addition) and organic fertilizer application treatments (solid manure and liquid slurry) were established. In the experiment, plant biomass is harvested three times per year, inducing strong nutrient limitation in plots that have not received nutrient additions (fully deficient or deficient in a single element). We determined gross rates of microbial protein depolymerization, N-mineralization and nitrification via isotope pool dilution assays with 15N-labeled amino acids, NH4+, and NO3-. We hypothesized that N deficiency (lack of N fertilization) would stimulate microbial N mining (depolymerization), and reduce subsequent N mineralization and nitrification. In contrast, we expected that organic fertilization would alleviate microbial C and N limitations, reducing N depolymerization rates and increasing mineralization and nitrification.Our results show that organically fertilized and limed soils have significantly lower gross protein depolymerization rates than plots receiving inorganic N. No significant differences were found comparing gross N-mineralization and gross nitrification rates across the different treatments. Given the higher rates of protein depolymerization in inorganically fertilized soils as compared to organically fertilized and limed soils, microbial N processes seem to be controlled by plant C input and/or soil pH rather than by direct soil nutrient availability. However, depolymerization of macromolecular N does not only supply N to the soil microbial community but also organic C. Thus, the reduced plant C input compared to fully fertilized soils may have caused microorganisms to increase their mining for a C-containing energy source, thereby increasing protein depolymerization rates. In summary, this study suggests that long term nutrient deficiency or nutrient imbalances may affect soil nutrient cycling indirectly by changing plant C inputs (via reduced primary production) and/or changing soil pH, rather than directly, by nutrient availability. This further indicates that soil microbial communities are rather C than nutrient limited.

The raise of atmospheric CO2 concentrations, with consequent increase in global warming and the likelihood of severe droughts, is altering the terrestrial biogeochemical carbon (C) cycle, with potential feedback to climate change. Microbial physiology, i.e. growth, turnover and carbon use efficiency, control soil carbon fluxes to the atmosphere. Thus, improving our ability to accurately quantify microbial physiology, and how it is affected by climate change, is essential. Recent advances in the field have allowed the quantification of community-level microbial growth and carbon use efficiency in dry conditions via an 18O water vapor equilibration technique, allowing for the first time to evaluate microbial growth rates under drought conditions.We modified the water vapor equilibration method using 2H-labelled water to estimate microbial community growth via deuterium incorporation into fatty acids. First, we verified that a rapid equilibration of 2H with soil water is possible. Then, we applied this approach to soil samples collected from a long-term climate change experiment (https://www.climgrass.at/) where warming, elevated atmospheric CO2 (eCO2) and drought are manipulated in a full factorial combination. Samples were taken in the field during peak drought and one week after rewetting. We used a high-throughput method to extract phospho- and neutral- lipid fatty acids (PLFA and NLFA) and we measured 2H enrichment in these compounds via GC-IRMS.Our results show that within 48 h, 2H in water vapor was in equilibrium with soil water and was detectable in microbial PLFA and NLFAs. We were able to quantify growth rates for different groups of microorganisms (Gram-positive, Gram-negative, Fungi and Actinobacteria) and calculate community level carbon use efficiency. We showed that a reduction of carbon use efficiency in the combined warming + eCO2 treatment was caused by a reduced growth of fungi and overall higher respiration rates. During drought, all groups showed a reduction in growth rates, albeit the reduction was stronger in bacteria than in fungi. Moreover, fungi accumulated high amounts of 2H into NLFAs, representing up to one third of the amount in PLFAs and indicating enhanced investment into storage compounds. This investment was still higher than in control plots two days after rewetting and returned to control levels within a week.Our study demonstrates that climate change can have strong effects on microbial physiology, with group-specific responses to different climate change factors. Our approach has the benefit of using fatty acid biomarkers to improve resolution into community level growth responses to climate change. This allowed a quantification of group-specific growth rates and concomitantly a measurement of investment into reserve compounds.

In temperate soil systems microbial biomass often increases during winter and decreases again in spring. This build up and release of microbial carbon could potentially lead to a build-up of stabilized soil carbon during winter times. The mechanism behind the increase in microbial carbon is not well understood. In this laboratory incubation study, we looked into microbial physiology as well as microbial glucose uptake and partitioning during cooling. Soils from a temperate forest and agricultural system were cooled down from field temperature of 11°C to 1°C. We added 13C-labelled glucose immediately and after an acclimation phase of 7 days and traced the 13C into microbial biomass, CO2 respired from the soil and phospholipid fatty acids. In addition we determined microbial growth using 18O-incorporation into DNA.First results show that while total respiration was strongly reduced when soils were cooled, glucose-derived respiration was as high in soils at 1°C as at 11°C. The same general pattern was found in soils during fast cooling and after an acclimation phase in agricultural and forest soils. We also saw an increased investment of glucose-derived carbon in unsaturated PLFAs. Since unsaturated fatty acids retain fluidity at lower temperatures compared to saturated fatty acids, this could be interpreted as precaution to reduced temperatures and potential freezing.Our results show a distinct response of the soil microbial community to cooling. The maintained glucose-derived respiration and incorporation into PLFAs at low temperatures compared to field temperature might indicate a preferential use of labile C forms during cooling. Moreover, the 13C incorporation into PLFAs may signal the buildup of cooling resistant cell membranes. These findings will be discussed with results from the 13C label tracing into microbial biomass, extractable organic carbon and total soil carbon as well as data on microbial growth and carbon use efficiency.

Introduction Reoccurring drought events can severely restrict forage production. However, experimentally drought stressed temperate forage grasslands have recently been reported to recover quickly after drought stress and re-wetting (DRW) and to be even more productive after drought than non-drought stressed control plots (Hofer et al., 2017; Hahn et al., in press). Although several studies show increased nutrient availability and microbial activity after DRW in grasslands (Bünemann et al., 2013; Sundert et al., 2020), an in-depth understanding if or how these mechanisms determine forage yield recovery is still missing.Methods This study examined the effect of a 2-month experimental summer drought under different nitrogen (N) fertilizer applications on the recovery of high-input Lolium perenne swards after re-wetting. Yield performance, physiology, nutrient availability and soil microbial activity were assessed over 2 years during and after the drought treatment. In addition, a post-drought transplantation experiment of control and DRW soil and plants withdrawn from the field was conducted to disentangle plant physiological and soil nutrient cycling effects on yield recovery after drought.Results Under all N applications, DRW outperformed the control yield in the field and showed higher N mineralization rates and higher soil N and K availabilities. Transplanted DRW plants showed longer but thinner leaves and decreased yields compared to control plants, irrespective of the soil’s DRW treatment. In contrast, DRW soils induced strongly increased L. perenne yields (on average +25%) compared to control soils. In summary, our data show that despite impaired plant growth after DRW, formerly drought stressed swards surpass control yields by profiting of higher mineralization rates and higher nutrient availability.ReferencesBünemann EK, Keller B, Hoop D, Jud K, Boivin P, Frossard E (2013) Increased availability of phosphorus after drying and rewetting of a grassland soil: processes and plant use. Plant and Soil 370: 511–526Hahn C, Lüscher A, Ernst-HaslerS, Suter M, Kahmen A (in press) Timing of drought in the growing season and strong legacy effects determine the annual productivity of temperate grasses in a changing climate. Biogeosciences.Hofer D, Suter M, Buchmann N, Lüscher A (2017) Nitrogen status of functionally different forage species explains resistance to severe drought and post-drought overcompensation. Agriculture, Ecosystems & Environment 236: 312–322Sundert KV, Brune V, Bahn M, Deutschmann M, Hasibeder R, Nijs I, Vicca S (2020) Post-drought rewetting triggers substantial K release and shifts in leaf stoichiometry in managed and abandoned mountain grasslands. Plant Soil 448: 353–368

The Amazon rainforest is an important sink for atmospheric CO2 counteracting increased emissions from anthropogenic fossil fuel combustion and land use change storing large amounts of carbon in plant biomass and soils. However, large parts of the Amazon Basin are characterized by highly weathered soils (ultisols and oxisols) with low availability of rock-derived phosphorus (and cations), which are mostly occluded in soil or bound in organic matter. Such low phosphorus availability is thought to be (co-)limiting plant productivity. However, much less is known whether low phosphorus availability influences the activity of heterotrophic microbial communities controlling litter and soil organic matter decomposition and thereby long-term carbon sequestration in tropical soils.In tropical soils high temperature and humid conditions allow overall high microbial activity. Over a larger soil phosphorus fertility gradient across several Amazonian rainforest sites, at low P sites almost 40 % of total P was stored in microbial biomass, highlighting the competitive strength of microorganisms and their importance as P reservoir. Across all sites soil microbial biomass was a significant predictor for soil microbial respiration, but mass-specific respiration rates (normalized by microbial biomass C) rather decreased at higher soil P. Using the incorporation of 18O from labelled water into DNA (i.e., a substrate-independent method) to determine microbial growth, we found significantly lower microbial growth rates per unit of microbial biomass at higher soil P. This resulted in a lower microbial carbon use efficiency, at a narrower C:P stoichiometry in soils with higher P levels, and pointed towards a microbial co-limitation of phosphorus and carbon at low soil P levels. Furthermore, data from a multi-year nutrient manipulation experiment in French Guiana and from short-term lab incubations suggest that microbial communities thriving at low P levels are highly efficient in taking up and storing added P, but do not necessarily respond with increased growth.Soil microbial communities play a crucial role in soil carbon and phosphorus cycling in tropical soils as potent competitors for available P. They also play an important role in storing and buffering P losses from highly weathered tropical soils. The potential non-homoeostatic stoichiometric behavior of microbial communities in P cycling is important to consider in soil and ecosystem models based on stoichiometric relationships.

Arbuscular mycorrhiza (AM) fungi are associated with almost all land plants and provide soil nutrients and other benefits to their plant hosts in exchange for photosynthetic products. While fertilization regimes in managed grasslands or agricultural systems are tailored for increasing plant biomass, their potential effects on AM fungi are rarely taken into account. Nutrient-driven changes in abundance and community composition of AM fungi, however, may feedback on ecosystem performance in the long term. Therefore, it is necessary to get a better understanding on how AM fungal communities respond to changes and imbalances in soil nutrient availabilities.Here, we evaluated how long-term nutrient deficiency of phosphorus (P), nitrogen (N) and potassium (K) affects the abundance and community composition of AM fungi in a mountainous grassland. In addition, we investigated how the responses of AM fungi to those deficiencies were modulated by liming and the type of fertilizer addition (inorganic versus organic).Our study was carried out on a long-term nutrient deficiency experimental grassland site in Admont (Styria, Austria), established in 1946. Different fertilization treatments were applied for more than 70 years in a randomized block design, including numerous combinations of inorganic (P, N, K with/without lime) and organic (solid manure and liquid slurry) fertilizers. The hay meadow at the site is cut three times per year and biomass is not returned to the system. Therefore, biomass and nutrients have been continuously removed for decades, leading to different types of soil nutrient deficiency. In this study, we collected both root and soil samples in July 2019 and quantified AM fungi and other microbial groups by measuring neutral fatty acid (NLFA) and phospholipid fatty acid (PLFA) biomarkers, respectively. Additionally, we applied DNA and RNA-based amplicon sequencing of the 18S rRNA gene to identify AM fungal community composition.Our data shows that deficiencies of one or more elements had a major impact on both AM fungal biomass and community composition. AM fungal biomass was higher in plots that received no fertilizers compared to inorganically fertilized plots, but lower in plots which were deficient only in certain single or multiple elements, specifically in plots fertilized with inorganic N only (i.e., deficient in P and K). Conversely, liming and organic fertilizer amendments increased AM fungal biomass compared to plots containing inorganic fertilizers without lime. Across all treatments, AM fungal biomass was positively correlated with pH and soil water content, and negatively with dissolved N compounds, indicating indirect effects via responses of other soil parameters to nutrient deficiency. Long-term nutrient deficiency also altered plant community composition, which may also have indirectly affected AM fungal communities.We conclude that long-term nutrient deficiency, and in particular the stoichiometry of available nutrients, strongly affects the abundance and community composition of AM fungi in grassland soil. This response may be linked to changes in plant community composition or soil chemistry both as a result and as a cause, emphasizing the complexity of feedbacks determining the response of grassland ecosystems to changing nutrient conditions.

Biological nitrogen fixation (BNF) isa fundamental part of nitrogen cycling in tropical forests, yet little is known about the contribution made by free-living nitrogen fixers inhabiting the often-extensive forest canopy. We used the acetylene reduction assay, calibrated with N-15(2), to measure free-living BNF on forest canopy leaves, vascular epiphytes, bryophytes and canopy soil, as well as on the forest floor in leaf litter and soil. We used a combination of calculated and published component densities to upscale free-living BNF rates to the forest level. We found that bryophytes and leaves situated in the canopy in particular displayed high mass-based rates of freeliving BNF. Additionally, we calculated that nearly 2 kg of nitrogen enters the forest ecosystem through free-living BNF every year, 40% of which was fixed by the various canopy components. Our results reveal that in the studied tropical lowland forest a large part of the nitrogen input through free-living BNF stems from the canopy, but also that the total nitrogen inputs by free-living BNF are lower than previously thought and comparable to the inputs of reactive nitrogen by atmospheric deposition. (C) 2020 Elsevier B.V. All rights reserved.

The abstract is available here: https://uscholar.univie.ac.at/o:1603267

Priming is the change of microbial soil organic matter (SOM) decomposition induced by a labile carbon (C) source. It is recognised as an important mechanism influencing soil C dynamics and C storage in terrestrial ecosystems. Microbial nitrogen (N) mining in SOM and preferential substrate utilisation, i.e., a shift in microbial carbon use from SOM to more labile energy sources, are possible, counteracting, mechanisms driving the priming effect. Climate warming and increased N availability might affect these mechanisms, and thus determine the direction and magnitude of the priming effect. Hence, these abiotic factors can indirectly affect soil C stocks, which makes their understanding crucial for predicting the soil C feedback in a warming world. We conducted a short-term incubation experiment (6 days) with soils from a subarctic grassland that had been subjected to long-term geothermal warming (55 years) by 2-4°C above unwarmed soil. Soil samples were amended with 13C-labelled glucose and 15N-labelled NH4NO3. We found a significantly negative relationship between in situ warming and cumulative primed C, with negative priming in the warmed soils. The negative priming suggests that preferential substrate utilisation was a key mechanism in our experiment. Our results indicate that changes in SOM characteristics associated with the in situ warming gradient can play a major role in determining the rate and direction of the priming effect. Additionally, we found that neither microbial N limitation nor N addition affected the priming effect, providing evidence that in our experiment, N mining did not lead to positive priming.

• 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 one year 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 show that rock‐derived nutrients determine 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 indicate that even slow‐growing tropical rainforests can respond rapidly to changes in resource availability.

The rapid rise in atmospheric CO2 concentration over the past century is unprecedented. It has unambiguously influenced Earth’s climate system and terrestrial ecosystems. Elevated atmospheric CO2 concentrations (eCO2) have induced an increase in biomass and thus, a carbon sink in forests worldwide. It is assumed that eCO2 stimulates photosynthesis and plant productivity and enhances water-use efficiency – the so-called CO2-fertilization effect, which may provide an important buffering effect for plants during adverse climate conditions. For these reasons, current global climate simulations consistently predict that tropical forests will continue to sequester more carbon in aboveground biomass, partially compensating human emissions and decelerating climate change by acting as a carbon sink. In contrast to model simulations, several lines of evidence point towards a decreasing carbon sink strength of the Amazon rainforest. Reliable predictions of eCO2 effects in the Amazon rainforest are hindered by a lack of process-based information gained from ecosystem scale eCO2 experiments. Here we report on baseline measurements from the Amazon Free Air CO2 Enrichment (AmazonFACE) experiment and preliminary results from open-top chamber (OTC) experiments with eCO2. After three months of eCO2, we find that understory saplings increased carbon assimilation by 17% (under light saturated conditions) and water use efficiency by 39% in the OTC experiment. We present our main hypotheses for the FACE experiment, and discuss our expectations on the potential driving processes for limiting or stimulating the Amazon rainforest carbon sink under eCO2. We focus on possible effects of eCO2 on carbon uptake and allocation, nutrient cycling, water-use and plant-herbivore interactions, which need to be implemented in dynamic vegetation models to estimate future changes of the Amazon carbon sink.

The Amazon rainforest faces immense pressures from human-induced deforestation and climate change and its future existence is largely indeterminate. Accurately projecting the forest’s response to future conditions, and thus preparing for the best possible outcome, requires a sound process-based understanding of its ecological and biogeochemical functioning. The intact forest acts as a sink of atmospheric carbon dioxide (CO2), however, this invaluable function is slowing down for unclear reasons, according to long-term plot measurements of tree growth. Earth system models, on the other hand, assume a continuous sink of carbon into the 21st century, predominantly driven by CO2 fertilization, concurrently buffering against adverse effects by climate change. Advancing empirical and experimental evidence points to strong nutrient constraints on the Amazon carbon sink, foremostly by phosphorus and other cations, so that the projected strength of the future carbon sink is certainly unrealistic. It is highly uncertain, however, to which degree nutrients are and will diminish elevated CO2-induced productivity, and to which extent plant-based mechanisms may upregulate phosphorus supply or optimize phosphorus use to facilitate the increasing demand by elevated CO2. Site-scale ecosystem model ensemble analysis underscores the diverging hypotheses on phosphorus feedbacks we are currently facing. In addition, heterogeneous soil phosphorus availability across the Amazon basin, in combination with a hyperdiverse plant community, challenges current efforts to project phosphorus constraints on the future of the Amazon carbon sink. We here give an outlook of current progress and future research needs of model-experiment integration to tackle this pressing question.

Highly weathered soils depleted in minerals and phosphorus (P) support large tracts of the tropical rainforests in the Central Amazon, which significantly contribute to the global carbon (C) sink. In these soils (oxisols and ferrasols), P is either occluded in Al/Fe-oxides, bound to the soil mineral matrix or in soil organic matter, and therefore not directly available for uptake as inorganic phosphate (Pi). To liberate Pi for plant or microbial uptake two processes are key: (i) changes of sorption-desorption equilibria of Pi with the soil matrix and (ii) the release of Pi from organic compounds (Po) catalyzed by enzymes, such as phosphatases. Plant roots and soil microbes have developed strategies to stimulate the release of P by accelerating P dissolution and desorption and by releasing extracellular phosphatases into the soil environment, which requires however C and energy investment. Because of P limitation in this ecosystem, the relative contributions of abiotic and biotic controls over P mineralization is of pivotal importance. Yet conclusive results are still scarce.We therefore aimed to disentangle abiotic and biotic controls over P mineralization in tropical soils. To achieve this, we collected forest soils from the Amazon Basin covering a range of soil texture and P concentrations, determined soil mineralogy and measured gross P desorption and mineralization rates using a 33P isotope pool dilution assay. Moreover, we determined acid phosphatase activity rates and microbial biomass C and P. We found significant differences between the studied sites in gross P influx and efflux rates into the Pi pool. Gross influx rates (i.e. the sum of Pi desorption and organic P mineralization) exceeded efflux (i.e. sorption or biotic Pi uptake rates) only in sandy and silty soils, while in clayey soils efflux rates dominated P fluxes indicating a very high Pi sorption capacity. However, gross influx and efflux rates were not related to total or dissolved P. Microbial biomass and acid phosphatase activity normalized to microbial biomass C were highest in sites with overall low total P microbial biomass P accounting for up to 40 % of total P in low P soils. We therefore conclude that in low P soils organic P turnover plays a major role in soil P cycling, and despite of the high P sorption capacity of clay rich soils, microbes can be strong competitors for plant available P.

Soils store more carbon than the atmosphere and total land plant biomass combined. Soil organic matter (SOM) can be classified into different physical pools characterized by their degree of protection and turnover rates. Usually, these pools are isolated by dividing soils in different water-stable aggregate size classes and, inside these classes, SOM fractions with differing densities and properties: Stable mineral-associated organic matter (MOM) and labile particulate organic matter (POM). Increasing temperatures are known to initially enhance microbial decomposition rates, releasing C from soils which could further accelerate climate change. The magnitude of this feedback depends on which C pool is affected the most by increased decomposition. Since MOM, thought to be the best protected carbon pool, holds most of the soil C, losses from this pool would potentially have the biggest impact on global climate. Experimental results are inconclusive so far, as most studies are based on short-term field warming (years rather than decades), leaving the ecosystem response to decades to century of warming uncertain.We made use of a geothermal warming platform in Iceland (ForHot; https://forhot.is/) to compare the effect of short-term (STW, 5-8 years) and long-term (LTW, more than 50 years) warming on soil organic carbon and nitrogen (SOC, SON) and its carbon and nitrogen isotope composition (δ13C and δ15N) in soil aggregates of different sizes in a subarctic grassland. OM fractions were isolated via density fractionation and ultrasonication both in macro- and microaggregates: Inter-aggregate free POM (fPOM), POM occluded within aggregates (iPOM) and MOM.MOM, containing most of the SOC and SON, showed a similar response to warming for both macro- and microaggregates. Compared to LTW plots, STW plots overall had higher C and N stocks. But warming reduced the carbon content more strongly in STW plot than in LTW plots. δ13C of MOM soil increased with temperature on the STW sites, indicating higher overall SOM turnover rates at higher temperatures, in line with the higher SOC losses. For LTW, δ13C decreased with warming except for the most extreme treatment (+16°C). Warming duration had no impact on iPOM-C. fPOM-C decreased in STW sites with increasing temperature, while it increased on the LTW sites.Overall our results demonstrate warming-induced C losses from the MOM-C-pool, thought to be most stable soil carbon pool. Thus, warming stimulated microbes to decompose both labile fPOM and more stable MOM. After decades of warming, C losses are less pronounced compared to the short-term warmed plots, pointing to a replenishment of the carbon pools at higher temperatures in the long-term. This might be explained by adaptations of the primary productivity and/or substrate-limitation of microbial growth.

The Amazon rainforest is the biggest tropical rainforest in the world and provides significant global climate regulation services. However, the future of the Amazon rainforest carbon sink under elevated CO2 is uncertain. The potential fertilization effect of elevated CO2 on the Amazon rainforest carbon sink may be constrained by phosphorus availability. Phosphorus is an essential element involved in all major plant processes and is considered to be the primary limiting nutrient in the Amazon rainforest. To cope with phosphorus limitation, plants have developed different strategies to increase the use efficiency, uptake, and availability of phosphorus. Vegetation models have identified phosphorus use and acquisition strategies as crucial to the projections of the future of the Amazon rainforest. Although some of the strategies are explicitly or implicitly represented in vegetation models, the mechanistic representations diverge due to the lack of empirical knowledge. Here, we synthesized the current understanding of the main strategies and how they may play out in the Amazon rainforest, namely, root phosphorus foraging, arbuscular mycorrhizal symbiosis, phosphatase and organic acids exudation, and leaf phosphorus resorption. We focus on mechanisms, drivers, and plasticity along soil phosphorus gradients of the named strategies, aiming to inform models and highlight important knowledge gaps, offering an opportunity to bring modeling and experimental research together.

Climate warming has been suggested to impact high latitude grasslands severely, causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts soil C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. We hypothesized that warming would increase belowground C allocation, while enhanced N availability would decrease it, and that their interactive effects would be additive.We studied a subarctic grassland located at a natural geothermal soil warming gradient close to Hveragerði, Iceland, which was established by an earthquake in 2008. We chose 14 plots along the gradient with soil warming temperatures ranging from 0 to 10°C above ambient, and fertilized a subset of plots with 50kg ha-1 y-1 of NH4NO3 twice a year prior to the study. We performed 13CO2 canopy pulse labeling for an hour and tracked the 13C pulse through the plant-microbe-soil system and into soil respiration for ten days after labeling.Our preliminary results show that at higher temperatures microbial activity increased, causing higher C turnover and a higher respiration of recently assimilated C from the soil. Warming significantly decreased microbial biomass, however, the recent C allocated from roots to microbes increased. This indicates a higher microbial C-limitation and a tighter root-microbe coupling under warming. Nitrogen addition increased the allocation of recent C to roots, microbial biomass, and soil respiration. The effects of N addition and warming were additive with no interaction. Our results indicate that the microbes in warmed soil may not be N limited, but could be C limited and depend more on the supply of recent C from plants. We conclude that in a future climate with warmer soils, more C may be allocated belowground, however, its residence time may decrease.

Temperature governs most biotic processes, yet we know little about how warming affects whole ecosystems. Here we examined the responses of 128 components of a subarctic grassland to either 5-8 or >50 years of soil warming. Warming of >50 years drove the ecosystem to a new steady state possessing a distinct biotic composition and reduced species richness, biomass and soil organic matter. However, the warmed state was preceded by an overreaction to warming, which was related to organism physiology and was evident after 5-8 years. Ignoring this overreaction yielded errors of >100% for 83 variables when predicting their responses to a realistic warming scenario of 1 degrees C over 50 years, although some, including soil carbon content, remained stable after 5-8 years. This study challenges long-term ecosystem predictions made from short-term observations, and provides a framework for characterization of ecosystem responses to sustained climate change. Comparing grassland ecosystem responses of 128 biotic and abiotic variables to geothermal warming, the authors find that short-term (5-8 years) responses are poor predictors of change over the long term (>50 years).

Tropical rainforests harbor exceptionally high biodiversity and store large amounts of carbon in vegetation biomass. However, regional variation in plant species richness and vegetation carbon stock can be substantial, and may be related to the heterogeneity of topoedaphic properties. Therefore, aboveground vegetation carbon storage typically differs between geographic forest regions in association with the locally dominant plant functional group. A better understanding of the underlying factors controlling tropical forest diversity and vegetation carbon storage could be critical for predicting tropical carbon sink strength in response to projected climate change. Based on regionally replicated 1-ha forest inventory plots established in a region of high geomorphological heterogeneity we investigated how climatic and edaphic factors affect tropical forest diversity and vegetation carbon storage. Plant species richness (of all living stems >10 cm in diameter) ranged from 69 to 127 ha−1 and vegetation carbon storage ranged from 114 to 200 t ha−1. While plant species richness was controlled by climate and soil water availability, vegetation carbon storage was strongly related to wood density and soil phosphorus availability. Results suggest that local heterogeneity in resource availability and plant functional composition should be considered to improve projections of tropical forest ecosystem functioning under future scenarios. Las selvas tropicales albergan una biodiversidad excepcionalmente alta y almacenan grandes cantidades de carbono en la biomasa de la vegetación. Sin embargo, la variación regional en la riqueza de especies de plantas y las existencias de carbono de la vegetación puede ser sustancial y puede estar relacionada con la heterogeneidad de las propiedades topoedáficas. Por lo tanto, el almacenamiento de carbono de la vegetación aérea normalmente difiere entre las regiones forestales geográficas en asociación con el grupo funcional de plantas localmente dominante. Una mejor comprensión de los factores subyacentes que controlan la diversidad de los bosques tropicales y el almacenamiento de carbono de la vegetación podría ser fundamental para predecir la fuerza de los sumideros de carbono tropical en respuesta al cambio climático proyectado. Con base en parcelas de inventario forestal de 1 ha replicadas regionalmente establecidas en una región de alta heterogeneidad geomorfológica, investigamos cómo los factores climáticos y edáficos afectan la diversidad de los bosques tropicales y el almacenamiento de carbono de la vegetación. La riqueza de especies de plantas (de todos los tallos vivos> 10 cm de diámetro) varió de 69 a 127 ha - 1 y el almacenamiento de carbono de la vegetación varió de 114 a 200 t ha - 1. Si bien la riqueza de especies de plantas estuvo controlada por el clima y la disponibilidad de agua del suelo, la vegetación El almacenamiento de carbono estaba fuertemente relacionado con la densidad de la madera y la disponibilidad de fósforo en el suelo. Los resultados sugieren que se debe considerar la heterogeneidad local en la disponibilidad de recursos y la composición funcional de las plantas para mejorar las proyecciones del funcionamiento del ecosistema forestal tropical en escenarios futuros. International Institute for Applied Systems Analysis, Austria Universidad de Costa Rica University of Antwerp, Belgium Universidad Nacional, Costa Rica University of Vienna, Austria University of Natural Resources and Life Sciences, Austria Universidad de Salamanca, España Nanyang Technological University, Singapore Cardiff University, United Kingdom Coordenação de Dinâmica Ambiental, Brasil Technical University of Munich, Germany University Park, United Kingdom Escuela de Ciencias Ambientales

Numerous studies have demonstrated that fertilization with nutrients such as nitrogen, phosphorus, and potassium increases plant productivity in both natural and managed ecosystems, demonstrating that primary productivity is nutrient limited in most terrestrial ecosystems. In contrast, it has been demonstrated that heterotrophic microbial communities in soil are primarily limited by organic carbon or energy. While this concept of contrasting limitations, that is, microbial carbon and plant nutrient limitation, is based on strong evidence that we review in this paper, it is often ignored in discussions of ecosystem response to global environment changes. The plant-centric perspective has equated plant nutrient limitations with those of whole ecosystems, thereby ignoring the important role of the heterotrophs responsible for soil decomposition in driving ecosystem carbon storage. To truly integrate carbon and nutrient cycles in ecosystem science, we must account for the fact that while plant productivity may be nutrient limited, the secondary productivity by heterotrophic communities is inherently carbon limited. Ecosystem carbon cycling integrates the independent physiological responses of its individual components, as well as tightly coupled exchanges between autotrophs and heterotrophs. To the extent that the interacting autotrophic and heterotrophic processes are controlled by organisms that are limited by nutrient versus carbon accessibility, respectively, we propose that ecosystems by definition cannot be 'limited' by nutrients or carbon alone. Here, we outline how models aimed at predicting non-steady state ecosystem responses over time can benefit from dissecting ecosystems into the organismal components and their inherent limitations to better represent plant-microbe interactions in coupled carbon and nutrient models.

Quantifying soil carbon dynamics is of utmost relevance in the context of global change because soils play an important role in land-atmosphere gas exchange. Our current understanding of both present and future carbon dynamics is limited because we fail to accurately represent soil processes across temporal and spatial scales, partly because of the paucity of data on the relative importance and hierarchical relationships between microbial, geochemical and climatic controls. Here, using observations from a 3,000-kyr-old soil chronosequence preserved in alluvial terrace deposits of the Merced River, California, we show how soil carbon dynamics are driven by the relationship between short-term biotic responses and long-term mineral weathering. We link temperature sensitivity of heterotrophic respiration to biogeochemical soil properties through their relationship with microbial activity and community composition. We found that soil mineralogy, and in particular changes in mineral reactivity and resulting nutrient availability, impacts the response of heterotrophic soil respiration to warming by altering carbon inputs, carbon stabilization, microbial community composition and extracellular enzyme activity. We demonstrate that biogeochemical alteration of the soil matrix (and not short-term warming) controls the composition of microbial communities and strategies to metabolize nutrients. More specifically, weathering first increases and then reduces nutrient availability and retention, as well as the potential of soils to stabilize carbon.

Grassland management can modify soil microbial carbon (C) and nitrogen (N) cycling, affecting the resistance to extreme weather events, which are predicted to increase in frequency and magnitude in the near future. However, effects of grassland management on microbial C and N cycling and their responses to extreme weather events, such as droughts and heatwaves, have rarely been tested in a combined approach. We therefore investigated whether grassland management affects microbial C and N cycling responses to drought and temperature manipulation. We collected soils from in situ drought experiments conducted in an extensively managed and an abandoned mountain grassland and incubated them at two temperature levels. We measured microbial respiration and substrate incorporation, as well as gross rates of organic and inorganic N cycling to estimate microbial C and N use efficiencies (CUE and NUE). The managed grassland was characterized by lower microbial biomass, lower fungi to bacteria ratio, and higher microbial CUE, but only slightly different microbial NUE. At both sites drought induced a shift in microbial community composition driven by an increase in Gram-positive bacterial abundance. Drought significantly reduced C substrate respiration and incorporation by microbes at both sites, while microbial CUE remained constant. In contrast, drought increased gross rates of N mineralization at both sites, whereas gross amino acid uptake rates only marginally changed. We observed a significant direct, as well as interactive effect between land management and drought on microbial NUE. Increased temperatures significantly stimulated microbial respiration and reduced microbial CUE independent of drought or land management. Although microbial N processing rates showed no clear response, microbial NUE significantly decreased at higher temperatures. In summary in our study, microbial CUE, in particular respiration, is more responsive to temperature changes. Although N processing rates were stronger responding to drought than to temperature microbial NUE was affected by both drought and temperature increase. We conclude that direct effects of drought and heatwaves can induce different responses in soil microbial C and N cycling similarly in the studied land management systems.