Divergence in plant and microbial allocation strategies explains continental patterns in microbial allocation and biogeochemical fluxes

Allocation trade‐offs shape ecological and biogeochemical phenomena at local to global scale. Plant allocation strategies drive major changes in ecosystem carbon cycling. Microbial allocation to enzymes that decompose carbon vs. organic nutrients may similarly affect ecosystem carbon cycling. Current solutions to this allocation problem prioritise stoichiometric tradeoffs implemented in plant ecology. These solutions may not maximise microbial growth and fitness under all conditions, because organic nutrients are also a significant carbon resource for microbes. I created multiple allocation frameworks and simulated microbial growth using a microbial explicit biogeochemical model. I demonstrate that prioritising stoichiometric trade‐offs does not optimise microbial allocation, while exploiting organic nutrients as carbon resources does. Analysis of continental‐scale enzyme data supports the allocation patterns predicted by this framework, and modelling suggests large deviations in soil C loss based on which strategy is implemented. Therefore, understanding microbial allocation strategies will likely improve our understanding of carbon cycling and climate.     

Global climate change and soil carbon stocks; predictions from two contrasting models for the turnover of organic carbon in soil

Enhanced release of CO₂ to the atmosphere from soil organic carbon as a result of increased temperatures may lead to a positive feedback between climate change and the carbon cycle, resulting in much higher CO₂ levels and accelerated global warming. However, the magnitude of this effect is uncertain and critically dependent on how the decomposition of soil organic C (heterotrophic respiration) responds to changes in climate. Previous studies with the Hadley Centre's coupled climate–carbon cycle general circulation model (GCM) (HadCM3LC) used a simple, single‐pool soil carbon model to simulate the response. Here we present results from numerical simulations that use the more sophisticated ‘RothC’ multipool soil carbon model, driven with the same climate data. The results show strong similarities in the behaviour of the two models, although RothC tends to simulate slightly smaller changes in global soil carbon stocks for the same forcing. RothC simulates global soil carbon stocks decreasing by 54 Gt C by 2100 in a climate change simulation compared with an 80 Gt C decrease in HadCM3LC. The multipool carbon dynamics of RothC cause it to exhibit a slower magnitude of transient response to both increased organic carbon inputs and changes in climate. We conclude that the projection of a positive feedback between climate and carbon cycle is robust, but the magnitude of the feedback is dependent on the structure of the soil carbon model.     

Large losses of inorganic nitrogen from tropical rainforests suggest a lack of nitrogen limitation

Inorganic nitrogen losses from many unpolluted mature tropical forests are over an order of magnitude higher than losses from analogous temperate forests. This pattern could either reflect a lack of N limitation or accelerated plant-soil N cycling under tropical temperatures and moisture. We used a simple analytical framework of the N cycle and compared our predictions with data of N in stream waters of temperate and tropical rainforests. While the pattern could be explained by differences in N limitation, it could not be explained based on up-regulation of the internal N cycle without invoking the unlikely assumption that tropical plants are two to four times less efficient at taking up N than temperate plants. Our results contrast with the idea that a tropical climate promotes and sustains an up-regulated and leaky - but N-limited - internal N cycle. Instead, they are consistent with the notion that many tropical rainforests exist in a state of N saturation.     

On multiple-nutrient-limited growth of microorganisms,with special reference to dual limitation by carbon and nitrogen substrates

Simultaneous limitation of microbial growth by two or more nutrients is discussed for dual carbon/nitrogen-limited growth in continuous culture. The boundaries of the zone where double-limited growth occurs can be clearly defined from both cultivation data and cellular composition and they can be also predicted from growth yield data measured under single-substrate-limited conditions. It is demonstrated that for the two nutrients carbon and nitrogen the zone of double nutrient limitation is dependent on both the C:N ratio of the growth medium and the growth (dilution) rate. The concept on double-(carbon/nitrogen)-limited growth presented here can be extended to other binary and multiple combinations of nutrients.     

Nutrient limitation and soil development: Experimental test of a biogeochemical theory

Walker & Syers (1976) proposed a conceptual model that describesthe pattern and regulation of soil nutrient pools and availability during long-term soil and ecosystem development. Their model implies that plantproduction generally should be limited by N on young soils and by P on oldsoils; N and P supply should more or less equilibrate onintermediate-aged soils. We tested the application of this model to nutrientlimitation, using a well characterized substrate age sequence in Hawaiianmontane rain forest. Earlier experiments had evaluated nutrient limitationin forests on a young (300 y) and an old (4,100,000 y) substrate on the samedevelopmental sequence; N alone limited tree growth on the youngsubstrate, while P alone did so on the old one. An additional fertilizerexperiment based on replicated treatments with N, P, and all othernutrients combined, applied in individually and in all factorialcombinations, was established in an intermediate-aged site in theLaupahoehoe Forest Reserve, Hawaii. Here, diameter increments of thedominant tree Metrosideros polymorpha increased slightly with Nadditions, and nearly doubled when N and P were added together.Additions of elements other than N and P had no significant effecton growth. These results show that N and P had equilibrated (relativeto plant requirements) in the intermediate aged site. Together withthe earlier experiments, these results suggest that the Walker and Syersmodel provides a useful starting point for explaining the nature anddistribution of nutrient limitation in forest ecosystems.     

Climate-driven spatial and temporal patterns in peatland pool biogeochemistry

Peatland pools are freshwater bodies that are highly dynamic aquatic ecosystems because of their small size and their development in organic-rich sediments. However, our ability to understand and predict their contribution to both local and global biogeochemical cycles under rapidly occurring environmental change is limited because the spatiotemporal drivers of their biogeochemical patterns and processes are poorly understood. We used (1) pool biogeochemical data from 20 peatlands in eastern Canada, the United Kingdom, and southern Patagonia and (2) multi-year data from an undisturbed peatland of eastern Canada, to determine how climate and terrain features drive the production, delivering and processing of carbon (C), nitrogen (N), and phosphorus (P) in peatland pools. Across sites, climate (24%) and terrain (13%) explained distinct portions of the variation in pool biogeochemistry, with climate driving spatial differences in pool dissolved organic C (DOC) concentration and aromaticity. Within the multi-year dataset, DOC, carbon dioxide (CO₂), total N concentrations, and DOC aromaticity were highest in the shallowest pools and at the end of the growing seasons, and increased gradually from 2016 to 2021 in relation to a combination of increases in summer precipitation, mean air temperature for the previous fall, and number of extreme summer heat days. Given the contrasting effects of terrain and climate, broad-scale terrain characteristics may offer a baseline for the prediction of small-scale pool biogeochemistry, while broad-scale climate gradients and relatively small year-to-year variations in local climate induce a noticeable response in pool biogeochemistry. These findings emphasize the reactivity of peatland pools to both local and global environmental change and highlight their potential to act as widely distributed climate sentinels within historically relatively stable peatland ecosystems.     

Biological nitrogen fixation: rates,patterns and ecological controls in terrestrial ecosystems

New techniques have identified a wide range of organisms with the capacity to carry out biological nitrogen fixation (BNF)—greatly expanding our appreciation of the diversity and ubiquity of N fixers—but our understanding of the rates and controls of BNF at ecosystem and global scales has not advanced at the same pace. Nevertheless, determining rates and controls of BNF is crucial to placing anthropogenic changes to the N cycle in context, and to understanding, predicting and managing many aspects of global environmental change. Here, we estimate terrestrial BNF for a pre-industrial world by combining information on N fluxes with ¹⁵N relative abundance data for terrestrial ecosystems. Our estimate is that pre-industrial N fixation was 58 (range of 40–100) Tg N fixed yr⁻¹; adding conservative assumptions for geological N reduces our best estimate to 44 Tg N yr⁻¹. This approach yields substantially lower estimates than most recent calculations; it suggests that the magnitude of human alternation of the N cycle is substantially larger than has been assumed.     

氮添加诱导的磷限制改变了亚热带黄山松林土壤微生物群落结构

土壤微生物在陆地生态系统的生物地球化学循环中起着重要作用。然而目前尚不清楚氮(N)添加量及其持续时间如何影响土壤微生物群落结构,以及微生物群落结构变化与微生物相对养分限制状况是否存在关联。本研究在亚热带黄山松林开展了N添加试验以模拟N沉降,并设置3个处理:对照(CK, 0 kg N·hm^-2·a^-1)、低N(LN, 40 kg N·hm^-2·a^-1)和高N(HN, 80 kg N·hm^-2·a^-1)。在N添加满1年和3年时测定土壤基本理化性质、磷脂脂肪酸含量和碳(C)、N、磷(P)获取酶活性,并通过生态酶化学计量分析土壤微生物的相对养分限制状况。结果表明: 1年N添加对土壤微生物群落结构无显著影响,3年LN处理显著提高了革兰氏阳性菌(G⁺)、革兰氏阴性菌(G^-)、放线菌(ACT)和总磷脂脂肪酸(TPLFA)含量,而3年HN处理对微生物的影响不显著,表明细菌和ACT对N添加可能更为敏感。N添加加剧了微生物C和P限制,而P限制是土壤微生物群落结构变化的最佳解释因子。这表明,N添加诱导的P限制可能更有利于部分贫营养菌(如G⁺)和参与P循环的微生物(如ACT)的生长,从而改变亚热带黄山松林土壤微生物群落结构。     

Methane emissions from wetlands: biogeochemical,microbial, and modeling perspectives from local to global scales

Understanding the dynamics of methane (CH₄) emissions is of paramount importance because CH₄ has 25 times the global warming potential of carbon dioxide (CO₂) and is currently the second most important anthropogenic greenhouse gas. Wetlands are the single largest natural CH₄ source with median emissions from published studies of 164 Tg yr^?1, which is about a third of total global emissions. We provide a perspective on important new frontiers in obtaining a better understanding of CH₄ dynamics in natural systems, with a focus on wetlands. One of the most exciting recent developments in this field is the attempt to integrate the different methodologies and spatial scales of biogeochemistry, molecular microbiology, and modeling, and thus this is a major focus of this review. Our specific objectives are to provide an up‐to‐date synthesis of estimates of global CH₄ emissions from wetlands and other freshwater aquatic ecosystems, briefly summarize major biogeophysical controls over CH₄ emissions from wetlands, suggest new frontiers in CH₄ biogeochemistry, examine relationships between methanogen community structure and CH₄ dynamics in situ, and to review the current generation of CH₄ models. We highlight throughout some of the most pressing issues concerning global change and feedbacks on CH₄ emissions from natural ecosystems. Major uncertainties in estimating current and future CH₄ emissions from natural ecosystems include the following: (i) A number of important controls over CH₄ production, consumption, and transport have not been, or are inadequately, incorporated into existing CH₄ biogeochemistry models. (ii) Significant errors in regional and global emission estimates are derived from large spatial‐scale extrapolations from highly heterogeneous and often poorly mapped wetland complexes. (iii) The limited number of observations of CH₄ fluxes and their associated environmental variables loosely constrains the parameterization of process‐based biogeochemistry models.     

Global vegetation biomass production efficiency constrained by models and observations

Plants use only a fraction of their photosynthetically derived carbon for biomass production (BP). The biomass production efficiency (BPE), defined as the ratio of BP to photosynthesis, and its variation across and within vegetation types is poorly understood, which hinders our capacity to accurately estimate carbon turnover times and carbon sinks. Here, we present a new global estimation of BPE obtained by combining field measurements from 113 sites with 14 carbon cycle models. Our best estimate of global BPE is 0.41 ± 0.05, excluding cropland. The largest BPE is found in boreal forests (0.48 ± 0.06) and the lowest in tropical forests (0.40 ± 0.04). Carbon cycle models overestimate BPE, although models with carbon–nitrogen interactions tend to be more realistic. Using observation‐based estimates of global photosynthesis, we quantify the global BP of non‐cropland ecosystems of 41 ± 6 Pg C/year. This flux is less than net primary production as it does not contain carbon allocated to symbionts, used for exudates or volatile carbon compound emissions to the atmosphere. Our study reveals a positive bias of 24 ± 11% in the model‐estimated BP (10 of 14 models). When correcting models for this bias while leaving modeled carbon turnover times unchanged, we found that the global ecosystem carbon storage change during the last century is decreased by 67% (or 58 Pg C).     

Ensemble projections elucidate effects of uncertainty in terrestrial nitrogen limitation on future carbon uptake

The magnitude of the nitrogen (N) limitation of terrestrial carbon (C) storage over the 21st century is highly uncertain because of the complex interactions between the terrestrial C and N cycles. We use an ensemble approach to quantify and attribute process‐level uncertainty in C‐cycle projections by analysing a 30‐member ensemble representing published alternative representations of key N cycle processes (stoichiometry, biological nitrogen fixation (BNF) and ecosystem N losses) within the framework of one terrestrial biosphere model. Despite large differences in the simulated present‐day N cycle, primarily affecting simulated productivity north of 40°N, ensemble members generally conform with global C‐cycle benchmarks for present‐day conditions. Ensemble projections for two representative concentration pathways (RCP 2.6 and RCP 8.5) show that the increase in land C storage due to CO₂ fertilization is reduced by 24 ± 15% due to N constraints, whereas terrestrial C losses associated with climate change are attenuated by 19 ± 20%. As a result, N cycling reduces projected land C uptake for the years 2006–2099 by 19% (37% decrease to 3% increase) for RCP 2.6, and by 21% (40% decrease to 9% increase) for RCP 8.5. Most of the ensemble spread results from uncertainty in temperate and boreal forests, and is dominated by uncertainty in BNF (10% decrease to 50% increase for RCP 2.6, 5% decrease to 100% increase for RCP 8.5). However, choices about the flexibility of ecosystem C:N ratios and processes controlling ecosystem N losses regionally also play important roles. The findings of this study demonstrate clearly the need for an ensemble approach to quantify likely future terrestrial C–N cycle trajectories. Present‐day C‐cycle observations only weakly constrain the future ensemble spread, highlighting the need for better observational constraints on large‐scale N cycling, and N cycle process responses to global change.     

Global patterns of foliar nitrogen isotopes and their relationships with climate, mycorrhizal fungi, foliar nutrient concentrations, and nitrogen availability

Ratios of nitrogen (N) isotopes in leaves could elucidate underlying patterns of N cycling across ecological gradients. To better understand global-scale patterns of N cycling, we compiled data on foliar N isotope ratios (δ¹⁵N), foliar N concentrations, mycorrhizal type and climate for over 11 000 plants worldwide. Arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal plants were depleted in foliar δ¹⁵N by 2‰, 3.2‰, 5.9‰, respectively, relative to nonmycorrhizal plants. Foliar δ¹⁵N increased with decreasing mean annual precipitation and with increasing mean annual temperature (MAT) across sites with MAT ≥ −0.5°C, but was invariant with MAT across sites with MAT < −0.5°C. In independent landscape-level to regional-level studies, foliar δ¹⁵N increased with increasing N availability; at the global scale, foliar δ¹⁵N increased with increasing foliar N concentrations and decreasing foliar phosphorus (P) concentrations. Together, these results suggest that warm, dry ecosystems have the highest N availability, while plants with high N concentrations, on average, occupy sites with higher N availability than plants with low N concentrations. Global-scale comparisons of other components of the N cycle are still required for better mechanistic understanding of the determinants of variation in foliar δ¹⁵N and ultimately global patterns in N cycling.     

Belowground carbon flux links biogeochemical cycles and resource‐use efficiency at the global scale

Nutrient limitation is pervasive in the terrestrial biosphere, although the relationship between global carbon (C) nitrogen (N) and phosphorus (P) cycles remains uncertain. Using meta‐analysis we show that gross primary production (GPP) partitioning belowground is inversely related to soil‐available N : P, increasing with latitude from tropical to boreal forests. N‐use efficiency is highest in boreal forests, and P‐use efficiency in tropical forests. High C partitioning belowground in boreal forests reflects a 13‐fold greater C cost of N acquisition compared to the tropics. By contrast, the C cost of P acquisition varies only 2‐fold among biomes. This analysis suggests a new hypothesis that the primary limitation on productivity in forested ecosystems transitions from belowground resources at high latitudes to aboveground resources at low latitudes as C‐intensive root‐ and mycorrhizal‐mediated nutrient capture is progressively replaced by rapidly cycling, enzyme‐derived nutrient fluxes when temperatures approach the thermal optimum for biogeochemical transformations.     

Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems

The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation.     

Integrating plant–soil interactions into global carbon cycle models

1.  Plant–soil interactions play a central role in the biogeochemical carbon (C), nitrogen (N) and hydrological cycles. In the context of global environmental change, they are important both in modulating the impact of climate change and in regulating the feedback of greenhouse gas emissions (CO₂, CH₄ and N₂O) to the climate system.
2.  Dynamic global vegetation models (DGVMs) represent the most advanced tools available to predict the impacts of global change on terrestrial ecosystem functions and to examine their feedbacks to climate change. The accurate representation of plant–soil interactions in these models is crucial to improving predictions of the effects of climate change on a global scale.
3.  In this paper, we describe the general structure of DGVMs that use plant functional types (PFTs) classifications as a means to integrate plant–soil interactions and illustrate how models have been developed to improve the simulation of: (a) soil carbon dynamics, (b) nitrogen cycling, (c) drought impacts and (d) vegetation dynamics. For each of these, we discuss some recent advances and identify knowledge gaps.
4.  We identify three ongoing challenges, requiring collaboration between the global modelling community and process ecologists. First, the need for a critical evaluation of the representation of plant–soil processes in global models; second, the need to supply and integrate knowledge into global models; third, the testing of global model simulations against large-scale multifactor experiments and data from observatory gradients.
5.   Synthesis . This paper reviews how plant–soil interactions are represented in DGVMs that use PFTs and illustrates some model developments. We also identify areas of ecological understanding and experimentation needed to reduce uncertainty in future carbon coupled climate change predictions.     

Persistent nitrogen limitation of stream biofilm communities along climate gradients in the Arctic

Climate change is rapidly reshaping Arctic landscapes through shifts in vegetation cover and productivity, soil resource mobilization, and hydrological regimes. The implications of these changes for stream ecosystems and food webs is unclear and will depend largely on microbial biofilm responses to concurrent shifts in temperature, light, and resource supply from land. To study those responses, we used nutrient diffusing substrates to manipulate resource supply to biofilm communities along regional gradients in stream temperature, riparian shading, and dissolved organic carbon (DOC) loading in Arctic Sweden. We found strong nitrogen (N) limitation across this gradient for gross primary production, community respiration and chlorophyll‐a accumulation. For unamended biofilms, activity and biomass accrual were not closely related to any single physical or chemical driver across this region. However, the magnitude of biofilm response to N addition was: in tundra streams, biofilm response was constrained by thermal regimes, whereas variation in light availability regulated this response in birch and coniferous forest streams. Furthermore, heterotrophic responses to experimental N addition increased across the region with greater stream water concentrations of DOC relative to inorganic N. Thus, future shifts in resource supply to these ecosystems are likely to interact with other concurrent environmental changes to regulate stream productivity. Indeed, our results suggest that in the absence of increased nutrient inputs, Arctic streams will be less sensitive to future changes in other habitat variables such as temperature and DOC loading.     

Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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Global patterns of root turnover for terrestrial ecosystems

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands ( r ² = 0.48) and forests ( r ² = 0.17) and for total root biomass in shrublands ( r ² = 0.55). On the basis of the best-fit exponential model, the Q ₁₀ for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature ( r ² = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.     

土壤有机碳和氮分解对温度变化的响应趋势与研究方法

总结了土壤中碳和氮贮量与温度的关系、土壤碳和氮分解对温度时空差异和直接加热升温的响应,以及土壤碳和氮分解对低温冻结及冻融循环的响应趋势,讨论了其研究方法的误差和不确定性,并对今后的研究提出了一些建议．气候变暖在短期内将使土壤碳和氮分解加速并引起CO2释放量增加,而长期过程中却并不一定会引起土壤碳和氮分解加速．合理解释不同研究结果的差异,除了需要系统分析土壤碳和氮分解对温度变化响应的机制外,还需要充分认识土壤碳和氮分解对温度变化响应的长期过程和短期过程的差异,以及研究方法、植被、土壤和气候等因素的影响．