Presence of N-fixing neighbors increases leaf N and δ13C in Castilleja applegatei, a root hemiparasite

Parasitic plants are known for their high transpiration rates and low water use efficiency (WUE), which the N-parasitism hypothesis posits is driven by N limitation. Thus, availability of N-fixing hosts may affect parasite’s WUE and in turn impact the surrounding plant community. Here, I investigate how the availability of an N-fixing host affects the root hemiparasite, Castilleja applegatei, and examines host-mediated effects on community structure and soil moisture. I surveyed plant diversity and percent cover and measured soil moisture in 120 1 × 1 m plots within Sagehen Experimental Forest, CA. Fifty percent of the plots included C. applegatei. In a subset of plots, I measured leaf N, C/N, δ13C, and δ15N in C. applegatei and in one N-fixer (Ceanothus prostratus) and two non-N-fixing plants (Artemisia tridentata and Wyethia mollis). In C. applegatei availability of N-fixing hosts corresponded to a significant increase in leaf %N, a distinct δ15N signature, and an increase in δ13C (which typically signifies an increased WUE). The presence of parasites was associated with a marginally significant decrease in WUE in N-fixing neighbors, but had no effect on the two non-N-fixing species. The presence of parasites did not impact diversity, percent cover, or soil moisture. These results broadly support the N-parasitism hypothesis and indicate that host type can affect parasite’s physiology and therefore have the potential to mediate parasite’s effects in the community; however, community-level impacts were not found here.

Revisiting Why Plants Become N Deficient Under Elevated CO2: Importance to Meet N Demand Regardless of the Fed-Form

An increase in plant biomass under elevated CO2 (eCO2) is usually lower than expected. N-deficiency induced by eCO2 is often considered to be a reason for this. Several hypotheses explain the induced N-deficiency: (1) eCO2 inhibits nitrate assimilation, (2) eCO2 lowers nitrate acquisition due to reduced transpiration, or (3) eCO2 reduces plant N concentration with increased biomass. We tested them using C3 (wheat, rice, and potato) and C4 plants (guinea grass, and Amaranthus) grown in chambers at 400 (ambient CO2, aCO2) or 800 (eCO2) μL L−1 CO2. In most species, we could not confirm hypothesis (1) with the measurements of plant nitrate accumulation in each organ. The exception was rice showing a slight inhibition of nitrate assimilation at eCO2, but the biomass was similar between the nitrate and urea-fed plants. Contrary to hypothesis (2), eCO2 did not decrease plant nitrate acquisition despite reduced transpiration because of enhanced nitrate acquisition per unit transpiration in all species. Comparing to aCO2, eCO2 remarkably enhanced water-use efficiency, especially in C3 plants, decreasing water demand for CO2 acquisition. As our results supported hypothesis (3) without any exception, we then examined if lowered N concentration at eCO2 indeed limits the growth using C3 wheat and C4 guinea grass under various levels of nitrate-N supply. While eCO2 significantly increased relative growth rate (RGR) in wheat but not in guinea grass, each species increased RGR with higher N supply and then reached a maximum as no longer N was limited. To achieve the maximum RGR, wheat required a 1.3-fold N supply at eCO2 than aCO2 with 2.2-fold biomass. However, the N requirement by guinea grass was less affected by the eCO2 treatment. The results reveal that accelerated RGR by eCO2 could create a demand for more N, especially in the leaf sheath rather than the leaf blade in wheat, causing N-limitation unless the additional N was supplied. We concluded that eCO2 amplifies N-limitation due to accelerated growth rate rather than inhibited nitrate assimilation or acquisition. Our results suggest that plant growth under higher CO2 will become more dependent on N but less dependent on water to acquire both CO2 and N.

Topography modulates effects of nitrogen deposition on microbial resource limitation in a nitrogen-saturated subtropical forest

Background Nitrogen (N) saturation theory proposes that an ecosystem might switch from N limitation to carbon (C), phosphorus (P), or other nutrient limitations if it receives continuous N input. Yet, after N limitation is removed, which nutrient is the most limited and whether topography modulates such change is rarely tested at a microbial level. Here, we conducted a two-year N addition experiment under two different topography positions (i.e. a slope and a valley) in a N-saturated subtropical forest. Soil enzyme activity was measured, and ecoenzymatic stoichiometry indexes were calculated as indicators of microbial resource limitation. Results In the valley, two-year N addition changed the activity of all studied enzymes to various degrees. As a result, microbial C limitation was aggravated in the valley, and consequently microbial decomposition of soil labile organic C increased, but microbial P limitation was alleviated due to the stoichiometry balance. On the slope, however, N addition did not significantly change the activity of the studied enzymes, and did not alter the status of microbial resource limitation. Conclusions These results indicate that C is a more limited element for microbial growth than P after removing N limitation, but we also highlight that topography can regulate the effect of N deposition on soil microbial resource limitation in subtropical forests. These findings provide useful supplements to the N saturation theory.

Long-term ecosystem nitrogen limitation from foliar δ15N data and a land surface model

The effect of nutrient availability on plant growth and the terrestrial carbon sink under climate change and elevated CO2 remains one of the main uncertainties of the terrestrial carbon cycle. This is partially due to the difficulty of assessing nutrient limitation at large scales over long periods of time. Consistent declines in leaf nitrogen (N) content and leaf δ15N have been used to suggest that nitrogen limitation has increased in recent decades, most likely due to the concurrent increase in atmospheric CO2. However, such datasets are often not straightforward to interpret due to the complex factors that contribute to the spatial and temporal variation in leaf N and isotope concentration. We use the land surface model QUINCY, which has the unique capacity to represent N isotopic processes, in conjunction with two large datasets of foliar N and N isotope content. We run the model with different scenarios to test whether foliar δ15N isotopic data can be used to infer large scale nitrogen limitation and if the observed trends are caused by increasing atmospheric CO2, changes in climate or changes in sources of anthropogenic N deposition. We show that while the model can capture the observed change in leaf N content and predicts widespread increases in N limitation, it does not capture the pronounced, but very spatially heterogeneous, decrease in foliar δ15N observed in the data across the globe. The addition of an observed temporal trend in isotopic composition of N deposition leads to a more pronounced decrease in simulated leaf δ15N. Our results show that leaf δ15N observations should not, on their own, be used to assess global scale N limitation and that using such a dataset in conjunction with a land surface model can reveal the drivers behind the observed patterns.

A novel representation of biological nitrogen fixation and competitive dynamics between nitrogen-fixing and non-fixing plants in a land model (GFDL LM4.1-BNF)

Representing biological nitrogen fixation (BNF) is an important challenge for coupled carbon (C) and nitrogen (N) land models. Initial representations of BNF in land models applied simplified phenomenological relationships. More recent representations of BNF are mechanistic and include the dynamic response of symbiotic BNF to N limitation of plant growth. However, they generally do not include the competitive dynamics between N-fixing and non-fixing plants, which is a key ecological mechanism that determines ecosystem-scale symbiotic BNF. Furthermore, asymbiotic BNF is generally not included in land models. Here, we present LM4.1-BNF, a novel representation of BNF (asymbiotic and symbiotic) and an updated representation of N cycling in the Geophysical Fluid Dynamics Laboratory Land Model 4.1 (LM4.1). LM4.1-BNF incorporates a mechanistic representation of asymbiotic BNF by soil microbes, a representation of the competitive dynamics between N-fixing and non-fixing plants, and distinct asymbiotic and symbiotic BNF temperature responses derived from corresponding observations. LM4.1-BNF makes reasonable estimations of major carbon (C) and N pools and fluxes and their temporal dynamics, in comparison to the previous version of LM4.1 with N cycling (LM3-SNAP) and to previous representations of BNF in land models generally (phenomenological representations and those without competitive dynamics between N-fixing and non-fixing plants and/or asymbiotic BNF) at a temperate forest site. LM4.1-BNF effectively reproduces asymbiotic BNF rate (13 kgNha-1yr-1) in comparison to observations (11 kgNha-1yr-1). LM4.1-BNF effectively reproduces the temporal dynamics of symbiotic BNF rate: LM4.1-BNF simulates a symbiotic BNF pulse in early succession that reaches 73 kgNha-1yr-1 at 15 years and then declines to ∼0 kgNha-1yr-1 at 300 years, similarly to observed symbiotic BNF, which reaches 75 kgNha-1yr-1 at 17 years and then declines to ∼0 kgNha-1yr-1 in late successional forests. As such, LM4.1-BNF can be applied to project the dynamic response of vegetation to N limitation of plant growth and the degree to which this will constrain the terrestrial C sink under elevated atmospheric CO2 concentration and other global change factors.

Enhanced carbon acquisition and use efficiency alleviate microbial carbon relative to nitrogen limitation under soil acidification

Background Soil microbial communities cope with an imbalanced supply of resources by adjusting their element acquisition and utilization strategies. Although soil pH has long been considered an essential driver of microbial growth and community composition, little is known about how soil acidification affects microbial acquisition and utilization of carbon (C) and nitrogen (N). To close the knowledge gap, we simulated soil acidification and created a pH gradient by adding eight levels of elemental sulfur (S) to the soil in a meadow steppe. Results We found that S-induced soil acidification strongly enhanced the ratio of fungi to bacteria (F:B) and microbial biomass C to N (MBC:MBN) and subsequently decreased the C:N imbalance between microbial biomass and their resources. The linear decrease in the C:N imbalance with decreasing soil pH implied a conversion from N limitation to C limitation. To cope with enhanced C versus N limitation, soil microbial communities regulated the relative production of enzymes by increasing the ratio of β-glucosidase (BG, C-acquiring enzyme) to leucine aminopeptidase (LAP, N-acquiring enzyme), even though both enzymatic activities decreased with S addition. Structural equation modeling (SEM) suggested that higher C limitation and C:N-acquiring enzyme stimulated microbial carbon-use efficiency (CUE), which counteracted the negative effect of metal stress (i.e., aluminum and manganese) under soil acidification. Conclusions Overall, these results highlight the importance of stoichiometric controls in microbial adaption to soil acidification, which may help predict soil microbial responses to future acid deposition.

JULES-CN: a coupled terrestrial carbon–nitrogen scheme (JULES vn5.1)

Understanding future changes in the terrestrial carbon cycle is important for reliable projections of climate change and impacts on ecosystems. It is well known that nitrogen (N) could limit plants' response to increased atmospheric carbon dioxide and it is therefore important to include a representation of the N cycle in Earth system models. Here we present the implementation of the terrestrial nitrogen cycle in the Joint UK Land Environment Simulator (JULES) – the land surface scheme of the UK Earth System Model (UKESM). Two configurations are discussed – the first one (JULES-CN) has a bulk soil biogeochemical model and the second one is a development configuration that resolves the soil biogeochemistry with depth (JULES-CNlayer). In JULES the nitrogen (N) cycle is based on the existing carbon (C) cycle and represents all the key terrestrial N processes in a parsimonious way. Biological N fixation is dependent on net primary productivity, and N deposition is specified as an external input. Nitrogen leaves the vegetation and soil system via leaching and a bulk gas loss term. Nutrient limitation reduces carbon-use efficiency (CUE – ratio of net to gross primary productivity) and can slow soil decomposition. We show that ecosystem level N limitation of net primary productivity (quantified in the model by the ratio of the potential amount of C that can be allocated to growth and spreading of the vegetation compared with the actual amount achieved in its natural state) falls at the lower end of the observational estimates in forests (approximately 1.0 in the model compared with 1.01 to 1.38 in the observations). The model shows more N limitation in the tropical savanna and tundra biomes, consistent with the available observations. Simulated C and N pools and fluxes are comparable to the limited available observations and model-derived estimates. The introduction of an N cycle improves the representation of interannual variability of global net ecosystem exchange, which was more pronounced in the C-cycle-only versions of JULES (JULES-C) than shown in estimates from the Global Carbon Project. It also reduces the present-day CUE from a global mean value of 0.45 for JULES-C to 0.41 for JULES-CN and 0.40 for JULES-CNlayer, all of which fall within the observational range. The N cycle also alters the response of the C fluxes over the 20th century and limits the CO2 fertilisation effect, such that the simulated current-day land C sink is reduced by about 0.5 Pg C yr−1 compared to the version with no N limitation. JULES-CNlayer additionally improves the representation of soil biogeochemistry, including turnover times in the northern high latitudes. The inclusion of a prognostic land N scheme marks a step forward in functionality and realism for the JULES and UKESM models.

Coexistence of dominant marine phytoplankton species sustained by nutrient specialization

Prochlorococcus and Synechococcus are the two dominant picocyanobacteria in the low-nutrient surface waters of the subtropical ocean, but the basis for their coexistence in these biomes is still unclear. Here we combine in situ microcosm experiments and an ecological model to show that this coexistence can arise from specialization in the uptake of distinct nitrogen (N) substrates. In field incubations, the response of both Prochlorococcus and Synechococcus to nanomolar N amendments demonstrates N limitation of growth in both populations, but Prochlorococcus showed a higher affinity to ammonium whereas Synechococcus was more adapted to nitrate uptake. A simple ecological model demonstrates that the differential nutrient affinity of these species can explain their coexistence. Phylogenetic analysis of the presence of nitrate reductase and nitrite reductase further support the higher nitrate affinity of Synechococcus compared to Prochlorococcus. Our study suggests that the evolution of differential nutrient affinities is an important mechanism for sustaining coexistence of species under resource competition.