全球气候变化对陆地植物碳氮磷生态化学计量学特征的影响

全球气候变化背景下生物地球化学循环的响应规律和陆地植物适应对策已受到广泛关注.本文在分析气候变暖和降水变化对不同生态系统植物C∶N∶P的影响、CO₂浓度升高对不同光合途径物种元素的影响,以及氮沉降对土壤 植物元素影响的短期和长期效应等基础上,从植物生理特性和土壤有效营养元素变化等方面揭示了其可能存在的内在机理,以期为研究C、N、P化学元素在土壤 植物之间传递与调节机制、陆地生态系统结构和功能,以及生物地球化学元素循环对气候变化的响应提供理论依据.最后提出了该领域研究中存在的问题及对今后研究的展望.     

Soil parent material—A major driver of plant nutrient limitations in terrestrial ecosystems

Because the capability of terrestrial ecosystems to fix carbon is constrained by nutrient availability, understanding how nutrients limit plant growth is a key contemporary question. However, what drives nutrient limitations at global scale remains to be clarified. Using global data on plant growth, plant nutritive status, and soil fertility, we investigated to which extent soil parent materials explain nutrient limitations. We found that N limitation was not linked to soil parent materials, but was best explained by climate: ecosystems under harsh (i.e., cold and or dry) climates were more N‐limited than ecosystems under more favourable climates. Contrary to N limitation, P limitation was not driven by climate, but by soil parent materials. The influence of soil parent materials was the result of the tight link between actual P pools of soils and physical–chemical properties (acidity, P richness) of soil parent materials. Some other ground‐related factors (i.e., soil weathering stage, landform) had a noticeable influence on P limitation, but their role appeared to be relatively smaller than that of geology. The relative importance of N limitation versus P limitation was explained by a combination of climate and soil parent material: at global scale, N limitation became prominent with increasing climatic constraints, but this global trend was modulated at lower scales by the effect of parent materials on P limitation, particularly under climates favourable to biological activity. As compared with soil parent materials, atmospheric deposition had only a weak influence on the global distribution of actual nutrient limitation. Our work advances our understanding of the distribution of nutrient limitation at global scale. In particular, it stresses the need to take soil parent materials into account when investigating plant growth response to environment changes.     

Aggravated phosphorus limitation on biomass production under increasing nitrogen loading: a meta‐analysis

Nitrogen (N) and phosphorus (P), either individually or in combination, have been demonstrated to limit biomass production in terrestrial ecosystems. Field studies have been extensively synthesized to assess global patterns of N impacts on terrestrial ecosystem processes. However, to our knowledge, no synthesis has been done so far to reveal global patterns of P impacts on terrestrial ecosystems, especially under different nitrogen (N) levels. Here, we conducted a meta‐analysis of impacts of P addition, either alone or with N addition, on aboveground (AGB) and belowground biomass production (BGB), plant and soil P concentrations, and N : P ratio in terrestrial ecosystems. Overall, our meta‐analysis quantitatively confirmed existing notions: (i) colimitation of N and P on biomass production and (ii) more P limitation in tropical forest than other ecosystems. More importantly, our analysis revealed new findings: (i) P limitation on biomass production was aggravated by N enrichment and (ii) plant P concentration was a better indicator of P limitation than soil P availability. Specifically, P addition increased AGB and BGB by 34% and 13%, respectively. The effect size of P addition on biomass production was larger in tropical forest than grassland, wetland, and tundra and varied with P fertilizer forms, P addition rates, or experimental durations. The P‐induced increase in biomass production and plant P concentration was larger under elevated than ambient N. Our findings suggest that the global limitation of P on biomass production will become severer under increasing N fertilizer and deposition in the future.     

Constraints to nitrogen acquisition of terrestrial plants under elevated CO2

A key part of the uncertainty in terrestrial feedbacks on climate change is related to how and to what extent nitrogen (N) availability constrains the stimulation of terrestrial productivity by elevated CO₂ (eCO₂), and whether or not this constraint will become stronger over time. We explored the ecosystem‐scale relationship between responses of plant productivity and N acquisition to eCO₂ in free‐air CO₂ enrichment (FACE) experiments in grassland, cropland and forest ecosystems and found that: (i) in all three ecosystem types, this relationship was positive, linear and strong (r² = 0.68), but exhibited a negative intercept such that plant N acquisition was decreased by 10% when eCO₂ caused neutral or modest changes in productivity. As the ecosystems were markedly N limited, plants with minimal productivity responses to eCO₂ likely acquired less N than ambient CO₂‐grown counterparts because access was decreased, and not because demand was lower. (ii) Plant N concentration was lower under eCO₂, and this decrease was independent of the presence or magnitude of eCO₂‐induced productivity enhancement, refuting the long‐held hypothesis that this effect results from growth dilution. (iii) Effects of eCO₂ on productivity and N acquisition did not diminish over time, while the typical eCO₂‐induced decrease in plant N concentration did. Our results suggest that, at the decennial timescale covered by FACE studies, N limitation of eCO₂‐induced terrestrial productivity enhancement is associated with negative effects of eCO₂ on plant N acquisition rather than with growth dilution of plant N or processes leading to progressive N limitation.     

氮添加对森林植物磷含量的影响及其机制

氮沉降对森林生态系统磷循环产生了不可忽视的影响, 尤其是加剧了植物生长的磷限制, 从而使得氮沉降背景下植物磷含量变化备受关注。该文综述了氮添加对森林植物磷含量的影响, 认为氮添加通过促进土壤磷酸酶活性进而提高土壤有效磷含量, 有利于植物的磷吸收并增加植物磷含量。同时, 森林植物磷含量对氮添加的响应还受物种、生活型以及施氮时间长短等因素的影响。基于森林植物磷含量对氮添加响应的差异性, 该文进一步探讨氮富集背景下森林植物磷含量变化的可能机制: 1)外源氮输入通过改变土壤中有效磷含量从而对植物磷的来源产生影响; 2)通过影响植物的根系分泌物、菌根共生和根系形态结构等进而影响植物的磷吸收能力; 3)通过影响植物的磷养分再分配、磷养分重吸收对植物磷利用效率产生影响。综上所述, 外源氮输入使植物磷含量发生改变, 首要原因是土壤有效磷含量的改变, 其次是植物磷吸收能力和磷利用效率的改变起调控作用。     

Effects of climate on soil phosphorus cycle and availability in natural terrestrial ecosystems

Climate is predicted to change over the 21st century. However, little is known about how climate change can affect soil phosphorus (P) cycle and availability in global terrestrial ecosystems, where P is a key limiting nutrient. With a global database of Hedley P fractions and key‐associated physiochemical properties of 760 (seminatural) natural soils compiled from 96 published studies, this study evaluated how climate pattern affected soil P cycle and availability in global terrestrial ecosystems. Overall, soil available P, indexed by Hedley labile inorganic P fraction, significantly decreased with increasing mean annual temperature (MAT) and precipitation (MAP). Hypothesis‐oriented path model analysis suggests that MAT negatively affected soil available P mainly by decreasing soil organic P and primary mineral P and increasing soil sand content. MAP negatively affected soil available P both directly and indirectly through decreasing soil primary mineral P; however, these negative effects were offset by the positive effects of MAP on soil organic P and fine soil particles, resulting in a relatively minor total MAP effect on soil available P. As aridity degree was mainly determined by MAP, aridity also had a relatively minor total effect on soil available P. These global patterns generally hold true irrespective of soil depth (≤10 cm or >10 cm) or site aridity index (≤1.0 or >1.0), and were also true for the low‐sand (≤50%) soils. In contrast, available P of the high‐sand (>50%) soils was positively affected by MAT and aridity and negatively affected by MAP. Our results suggest that temperature and precipitation have contrasting effects on soil P availability and can interact with soil particle size to control soil P availability.     

Does long-term soil warming affect microbial element limitation? A test by short-term assays of microbial growth responses to labile C,N and P additions

Increasing global temperatures have been reported to accelerate soil carbon (C) cycling, but also to promote nitrogen (N) and phosphorus (P) dynamics in terrestrial ecosystems. However, warming can differentially affect ecosystem C, N and P dynamics, potentially intensifying elemental imbalances between soil resources, plants and soil microorganisms. Here, we investigated the effect of long-term soil warming on microbial resource limitation, based on measurements of microbial growth (¹⁸O incorporation into DNA) and respiration after C, N and P amendments. Soil samples were taken from two soil depths (0–10, 10–20 cm) in control and warmed (>14 years warming, +4°C) plots in the Achenkirch soil warming experiment. Soils were amended with combinations of glucose-C, inorganic/organic N and inorganic/organic P in a full factorial design, followed by incubation at their respective mean field temperatures for 24 h. Soil microbes were generally C-limited, exhibiting 1.8-fold to 8.8-fold increases in microbial growth upon C addition. Warming consistently caused soil microorganisms to shift from being predominately C limited to become C-P co-limited. This P limitation possibly was due to increased abiotic P immobilization in warmed soils. Microbes further showed stronger growth stimulation under combined glucose and inorganic nutrient amendments compared to organic nutrient additions. This may be related to a prolonged lag phase in organic N (glucosamine) mineralization and utilization compared to glucose. Soil respiration strongly positively responded to all kinds of glucose-C amendments, while responses of microbial growth were less pronounced in many of these treatments. This highlights that respiration–though easy and cheap to measure—is not a good substitute of growth when assessing microbial element limitation. Overall, we demonstrate a significant shift in microbial element limitation in warmed soils, from C to C-P co-limitation, with strong repercussions on the linkage between soil C, N and P cycles under long-term warming.     

模拟增温对中亚热带杉木人工林土壤磷有效性的影响

气候变暖改变与土壤磷循环相关的生物地球化学过程,对陆地生态系统磷循环产生直接或间接影响。为研究亚热带地区杉木人工林土壤磷有效性对增温的响应,开展了模拟增温实验。实验设置对照组及增温组(5℃),经过1.5a的短期增温,对杉木人工林的土壤全磷、有机磷、微生物量磷、有效磷、酸性磷酸酶活性及相关土壤理化性质进行测定,结果表明:增温处理下,土壤酸性磷酸酶活性提高约1.5倍,土壤全磷、微生物量磷以及有机磷含量分别减少了6%、34%和12%,土壤有效磷含量增加25%。可见,短期增温通过提高土壤磷酸酶活性进而促进土壤有机磷矿化和降低土壤微生物固磷量,从而增加土壤磷有效性,但是增温导致潜在可利用的土壤微生物量磷大幅度的降低,将有可能加剧亚热带杉木人工林土壤磷限制。     

Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

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.     

Long‐term antagonistic effect of increased precipitation and nitrogen addition on soil respiration in a semiarid steppe

Changes in water and nitrogen (N) availability due to climate change and atmospheric N deposition could have significant effects on soil respiration, a major pathway of carbon (C) loss from terrestrial ecosystems. A manipulative experiment simulating increased precipitation and atmospheric N deposition has been conducted for 9 years (2005–2013) in a semiarid grassland in Mongolian Plateau, China. Increased precipitation and N addition interactively affect soil respiration through the 9 years. The interactions demonstrated that N addition weakened the precipitation‐induced stimulation of soil respiration, whereas increased precipitation exacerbated the negative impacts of N addition. The main effects of increased precipitation and N addition treatment on soil respiration were 15.8% stimulated and 14.2% suppressed, respectively. Moreover, a declining pattern and 2‐year oscillation were observed for soil respiration response to N addition under increased precipitation. The dependence of soil respiration upon gross primary productivity and soil moisture, but not soil temperature, suggests that resources C substrate supply and water availability are more important than temperature in regulating interannual variations of soil C release in semiarid grassland ecosystems. The findings indicate that atmospheric N deposition may have the potential to mitigate soil C loss induced by increased precipitation, and highlight that long‐term and multi‐factor global change studies are critical for predicting the general patterns of terrestrial C cycling in response to global change in the future.     

Ecosystem recovery: heathland response to a reduction in nitrogen deposition

Atmospheric deposition of nitrogen is responsible for widespread changes in the structure and function of sensitive seminatural ecosystems. The proposed reduction in emissions of nitrogenous pollutants in Europe under the Gothenburg Protocol raises the question of whether affected ecosystems have the potential to recover to their previous condition and, if so, over what timescale. Since 1998, we have monitored the response of a lowland heathland in southern England following the cessation of a long‐term nitrogen addition experiment, and subsequent management, assessing changes in vegetation growth and chemistry, soil chemistry and the soil microbial community. Persistent effects of earlier nutrient loading on Calluna growth and phenology, and on the abundance of lichens, were apparent up to 8 years after nitrogen additions ceased, indicating the potential for long‐term effects of modest nutrient loading (up to 15.4 kg N ha^?1 yr^?1, over 7 years) on heathland ecosystems. The size and activity of the soil microbial community was elevated in former N‐treated plots, 6–8 years after additions ceased, suggesting a prolonged effect on the rate of nutrient cycling. Although habitat management in 1998 reduced nitrogen stores in plant biomass, effects on belowground nitrogen stores were small. Although some parameters (e.g. soil pH) recover pretreatment levels relatively rapidly, others (e.g. vegetation cover and microbial activity) respond much more slowly, indicating that the ecological effects of even small increases in nitrogen deposition will persist for many years after deposition inputs are reduced. Indeed, calculations suggest that the additional soil nitrogen storage associated with 7 years of experimental nitrogen inputs could sustain the observed effects on plant growth and phenology for several decades. Carry over effects on plant phenology and sensitivity to drought suggest that the persistence of vegetation responses to nitrogen deposition should be integrated into long‐term assessments of the impact of global climate change on sensitive ecosystems.     

Global patterns and substrate‐based mechanisms of the terrestrial nitrogen cycle

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses.     

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。     

A global analysis of fine root production as affected by soil nitrogen and phosphorus

Fine root production is the largest component of belowground production and plays substantial roles in the biogeochemical cycles of terrestrial ecosystems. The increasing availability of nitrogen (N) and phosphorus (P) due to human activities is expected to increase aboveground net primary production (ANNP), but the response of fine root production to N and P remains unclear. If roots respond to nutrients as ANNP, fine root production is anticipated to increase with increasing soil N and P. Here, by synthesizing data along the nutrient gradient from 410 natural habitats and from 469 N and/or P addition experiments, we showed that fine root production increased in terrestrial ecosystems with an average increase along the natural N gradient of up to 0.5 per cent with increasing soil N. Fine root production also increased with soil P in natural conditions, particularly at P < 300 mg kg(-1). With N, P and combined N + P addition, fine root production increased by a global average of 27, 21 and 40 per cent, respectively. However, its responses differed among ecosystems and soil types. The global average increases in fine root production are lower than those of ANNP, indicating that above- and belowground counterparts are coupled, but production allocation shifts more to aboveground with higher soil nutrients. Our results suggest that the increasing fertilizer use and combined N deposition at present and in the future will stimulate fine root production, together with ANPP, probably providing a significant influence on atmospheric CO(2) emissions.     

Shifts in symbiotic associations in plants capable of forming multiple root symbioses across a long‐term soil chronosequence

Changes in soil nutrient availability during long‐term ecosystem development influence the relative abundances of plant species with different nutrient‐acquisition strategies. These changes in strategies are observed at the community level, but whether they also occur within individual species remains unknown. Plant species forming multiple root symbioses with arbuscular mycorrhizal (AM) fungi, ectomycorrhizal (ECM) fungi, and nitrogen‐(N) fixing microorganisms provide valuable model systems to examine edaphic controls on symbioses related to nutrient acquisition, while simultaneously controlling for plant host identity. We grew two co‐occurring species, Acacia rostellifera (N₂‐fixing and dual AM and ECM symbioses) and Melaleuca systena (AM and ECM dual symbioses), in three soils of contrasting ages (c. 0.1, 1, and 120 ka) collected along a long‐term dune chronosequence in southwestern Australia. The soils differ in the type and strength of nutrient limitation, with primary productivity being limited by N (0.1 ka), co‐limited by N and phosphorus (P) (1 ka), and by P (120 ka). We hypothesized that (i) within‐species root colonization shifts from AM to ECM with increasing soil age, and that (ii) nodulation declines with increasing soil age, reflecting the shift from N to P limitation along the chronosequence. In both species, we observed a shift from AM to ECM root colonization with increasing soil age. In addition, nodulation in A. rostellifera declined with increasing soil age, consistent with a shift from N to P limitation. Shifts from AM to ECM root colonization reflect strengthening P limitation and an increasing proportion of total soil P in organic forms in older soils. This might occur because ECM fungi can access organic P via extracellular phosphatases, while AM fungi do not use organic P. Our results show that plants can shift their resource allocation to different root symbionts depending on nutrient availability during ecosystem development.     

Elevated CO2 promotes long‐term nitrogen accumulation only in combination with nitrogen addition

Biogeochemical models that incorporate nitrogen (N) limitation indicate that N availability will control the magnitude of ecosystem carbon uptake in response to rising CO₂. Some models, however, suggest that elevated CO₂ may promote ecosystem N accumulation, a feedback that in the long term could circumvent N limitation of the CO₂ response while mitigating N pollution. We tested this prediction using a nine‐year CO₂xN experiment in a tidal marsh. Although the effects of CO₂ are similar between uplands and wetlands in many respects, this experiment offers a greater likelihood of detecting CO₂ effects on N retention on a decadal timescale because tidal marshes have a relatively open N cycle and can accrue soil organic matter rapidly. To determine how elevated CO₂ affects N dynamics, we assessed the three primary fates of N in a tidal marsh: (1) retention in plants and soil, (2) denitrification to the atmosphere, and (3) tidal export. We assessed changes in N pools and tracked the fate of a ¹⁵N tracer added to each plot in 2006 to quantify the fraction of added N retained in vegetation and soil, and to estimate lateral N movement. Elevated CO₂ alone did not increase plant N mass, soil N mass, or ¹⁵N label retention. Unexpectedly, CO₂ and N interacted such that the combined N+CO₂ treatment increased ecosystem N accumulation despite the stimulation in N losses indicated by reduced ¹⁵N label retention. These findings suggest that in N‐limited ecosystems, elevated CO₂ is unlikely to increase long‐term N accumulation and circumvent progressive N limitation without additional N inputs, which may relieve plant–microbe competition and allow for increased plant N uptake.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Nutrient stoichiometry of linked catchment‐lake systems along a gradient of land use

1. Catchments export nutrients to aquatic ecosystems at rates and ratios that are strongly influenced by land use practices, and within aquatic ecosystems nutrients can be processed, retained, lost to the atmosphere, or exported downstream. The stoichiometry of carbon and nutrients can influence ecosystem services such as water quality, nutrient limitation, biodiversity, eutrophication and the sequestration of nutrients and carbon in sediments. However, we know little about how nutrient stoichiometry varies along the pathway from terrestrial landscapes through aquatic systems. 2. We studied the stoichiometry of nitrogen and phosphorus exported by three catchments of contrasting land use (forest versus agriculture) and in the water column and sediments of downstream reservoirs. We also related stoichiometry to phytoplankton nutrient limitation and the abundance of heterocystous cyanobacteria. 3. The total N : P of stream exports varied greatly among catchments and was 18, 54 and 140 (molar) in the forested, mixed‐use and agricultural catchment, respectively. Total N : P in the mixed layers of the lakes was less variable but ordered similarly: 35, 52 132 in the forested, mixed‐use and agricultural lake, respectively. In contrast, there was little variation among systems in the C : N and C : P ratios of catchment exports or in reservoir seston. 4. Phytoplankton in the forested lake were consistently N limited, those in the agricultural lake were consistently P limited, and those in the mixed‐use lake shifted seasonally from P‐ to N limitation, reflecting N : P supply ratios. Total phytoplankton and cyanobacteria biomass were highest in the agricultural lake, but heterocystous (potentially N fixing) cyanobacteria were most abundant in the forested lake, corresponding to low N : P ratios. 5. Despite large differences in catchment export and water column N : P ratios, the N : P of sediment burial (integrated over several decades) was very low and remarkably similar (4.3–7.3) across reservoirs. N and P budgets constructed for the agricultural reservoir suggested that denitrification could be a major loss of N, and may help explain the relatively low N : P of buried sediment. 6. Our results show congruence between the catchment export N : P, reservoir N : P, phytoplankton N versus P limitation and the dominance of heterocystous cyanobacteria. However, the N : P stoichiometry of sediments retained in the lakes was relatively insensitive to catchment stoichiometry, suggesting that a common set of biogeochemical processes constrains sediment N : P across lakes of contrasting catchment land use.