Temperature and the metabolic balance of streams

1. It is becoming increasingly clear that fresh waters play a major role in the global C cycle. Stream ecosystem respiration (ER) and gross primary productivity (GPP) exert a significant control on organic carbon fluxes in fluvial networks. However, little is known about how climate change will influence these fluxes. 2. Here, we used a ‘natural experiment’ to demonstrate the role of temperature and nutrient cycling in whole‐system metabolism (ER, GPP and net ecosystem production – NEP), in naturally heated geothermal (5–25 °C) Icelandic streams. 3. We calculated ER and GPP with a new, more accurate method, which enabled us to take into account the additional uncertainties owing to stream spatial heterogeneity in oxygen concentrations within a reach. ER ranged 1–25 g C m^?2 day^?1 and GPP 1–10 g C m^?2 day^?1. The median uncertainties (based on 1 SD) in ER and GPP were 50% and 20%, respectively. 4. Despite extremely low water nutrient concentrations, high metabolic rates in the warm streams were supported by fast cycling rates of nutrients, as revealed from inorganic nutrient (N, P) addition experiments. 5. ER exceeded GPP in all streams (with average GPP/ER = 0.6) and was more strongly related to temperature than GPP, resulting in elevated negative NEP with warming. We show that, as a first approximation based on summer investigations, global stream carbon emission to the atmosphere would nearly double from 0.12 Pg C year^?1 at 13 °C to 0.21 (0.15–0.33) Pg C year^?1 with a 5 °C warming. 6. Compared to previous studies from natural systems (including terrestrial ecosystems), the temperature dependence of stream metabolism was not confounded by latitude or altitude, seasonality, light and nutrient availability, water chemistry, space availability (water transient storage), and water availability. 7. Consequently, stream nutrient processing is likely to increase with warming, protecting downstream ecosystems (rivers, estuaries, coastal marine systems) during the summer low flows from nutrient enrichment, but at the cost of increased CO₂ flux back to the atmosphere.     

A synthesis: The role of nutrients as constraints on carbon balances in boreal and arctic regions

As in many ecosystems, carbon (C) cycling in arctic and boreal regions is tightly linked to the cycling of nutrients: nutrients (particularly nitrogen) are mineralized through the process of organic matter decomposition (C mineralization), and nutrient availability strongly constrains ecosystem C gain through primary production. This link between C and nutrient cycles has implications for how northern systems will respond to future climate warming and whether feedbacks to rising concentrations of atmospheric CO₂ from these regions will be positive or negative. Warming is expected to cause a substantial release of C to the atmosphere because of increased decomposition of the large amounts of organic C present in high-latitude soils (a positive feedback to climate warming). However, increased nutrient mineralization associated with this decomposition is expected to stimulate primary production and ecosystem C gain, offsetting or even exceeding C lost through decomposition (a negative feedback to climate warming). Increased primary production with warming is consistent with results of numerous experiments showing increased plant growth with nutrient enrichment. Here we examine key assumptions behind this scenario: (1) temperature is a primary control of decomposition in northern regions, (2) increased decomposition and associated nutrient release are tightly coupled to plant nutrient uptake, and (3) short-term manipulations of temperature and nutrient availability accurately predict long-term responses to climate change.     

The stoichiometry of nitrogen and phosphorus spiralling in heterotrophic and autotrophic streams

1. Nutrient spiralling provides a conceptual framework and a whole‐system approach to investigate ecosystem responses to environmental changes. We use spiralling metrics to examine how the coupling of nitrogen and phosphorus uptake varies between streams dominated by either heterotrophic (i.e. bacteria‐dominated) or autotrophic (algal‐dominated) microbial communities. 2. Algae generally exhibit greater capacity to store nutrients than bacteria because of differences in cellular structures. These differences led us to hypothesise that the uptake of N and P in heterotrophic ecosystems should have reduced stoichiometric variation in response to changes in supply N : P compared to autotrophic ecosystems when assimilation dominates nutrient uptake. 3. To test this hypothesis, we used an array of serial nutrient additions in several streams in the South Fork Eel River watershed in Northern California. In one set of experiments, N and P were added alone and simultaneously in separate experiments to two small, heterotrophic streams to assess uptake rates and interactions between nutrient cycles. In a second set of experiments, N and P were added simultaneously at a range of N : P in one heterotrophic and one autotrophic stream to assess differences in uptake responses to changes in supply N : P. 4. Results of these experiments suggest two important conclusions. First, increased N supply significantly shortened P uptake lengths, while P addition had little impact on N uptake in both streams, indicating that uptake of non‐limiting nutrients is tightly coupled to the availability of the limiting element. Second, changes in P uptake and uptake ratios (U_N : U_P) with increased supply N : P supported our hypothesis that heterotrophic streams are more homeostatic in their responses to changes in nutrient supply than autotrophic streams, suggesting that physiological controls on nutrient use scale up to influence ecosystem‐scale patterns in nutrient cycling.     

Warming alters coupled carbon and nutrient cycles in experimental streams

Although much effort has been devoted to quantifying how warming alters carbon cycling across diverse ecosystems, less is known about how these changes are linked to the cycling of bioavailable nitrogen and phosphorus. In freshwater ecosystems, benthic biofilms (i.e. thin films of algae, bacteria, fungi, and detrital matter) act as biogeochemical hotspots by controlling important fluxes of energy and material. Understanding how biofilms respond to warming is thus critical for predicting responses of coupled elemental cycles in freshwater systems. We developed biofilm communities in experimental streamside channels along a gradient of mean water temperatures (7.5–23.6 °C), while closely maintaining natural diel and seasonal temperature variation with a common water and propagule source. Both structural (i.e. biomass, stoichiometry, assemblage structure) and functional (i.e. metabolism, N₂‐fixation, nutrient uptake) attributes of biofilms were measured on multiple dates to link changes in carbon flow explicitly to the dynamics of nitrogen and phosphorus. Temperature had strong positive effects on biofilm biomass (2.8‐ to 24‐fold variation) and net ecosystem productivity (44‐ to 317‐fold variation), despite extremely low concentrations of limiting dissolved nitrogen. Temperature had surprisingly minimal effects on biofilm stoichiometry: carbon:nitrogen (C:N) ratios were temperature‐invariant, while carbon:phosphorus (C:P) ratios declined slightly with increasing temperature. Biofilm communities were dominated by cyanobacteria at all temperatures (>91% of total biovolume) and N₂‐fixation rates increased up to 120‐fold between the coldest and warmest treatments. Although ammonium‐N uptake increased with temperature (2.8‐ to 6.8‐fold variation), the much higher N₂‐fixation rates supplied the majority of N to the ecosystem at higher temperatures. Our results demonstrate that temperature can alter how carbon is cycled and coupled to nitrogen and phosphorus. The uncoupling of C fixation from dissolved inorganic nitrogen supply produced large unexpected changes in biofilm development, elemental cycling, and likely downstream exports of nutrients and organic matter.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

Temporal Variation in the Importance of a Dominant Consumer to Stream Nutrient Cycling

Animal excretion can be a significant nutrient flux within ecosystems, where it supports primary production and facilitates microbial decomposition of organic matter. The effects of excretory products on nutrient cycling have been documented for various species and ecosystems, but temporal variation in these processes is poorly understood. We examined variation in excretion rates of a dominant grazing snail, Elimia clavaeformis, and its contribution to nutrient cycling, over the course of 14 months in a well-studied, low-nutrient stream (Walker Branch, east Tennessee, USA). Biomass-specific excretion rates of ammonium varied over twofold during the study, coinciding with seasonal changes in food availability (measured as gross primary production) and water temperature (multiple linear regression, R ² = 0.57, P = 0.053). The contribution of ammonium excretion to nutrient cycling varied with seasonal changes in both biological (that is, nutrient uptake rate) and physical (that is, stream flow) variables. On average, ammonium excretion accounted for 58% of stream water ammonium concentrations, 26% of whole-stream nitrogen demand, and 66% of autotrophic nitrogen uptake. Phosphorus excretion by Elimia was contrastingly low throughout the year, supplying only 1% of total dissolved phosphorus concentrations. The high average N:P ratio (89:1) of snail excretion likely exacerbated phosphorus limitation in Walker Branch. To fully characterize animal excretion rates and effects on ecosystem processes, multiple measurements through time are necessary, especially in ecosystems that experience strong seasonality.     

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.     

Importance of Terrestrial Arthropods as Subsidies in Lowland Neotropical Rain Forest Stream Ecosystems

The importance of terrestrial arthropods has been documented in temperate stream ecosystems, but little is known about the magnitude of these inputs in tropical streams. Terrestrial arthropods falling from the canopy of tropical forests may be an important subsidy to tropical stream food webs and could also represent an important flux of nitrogen (N) and phosphorus (P) in nutrient‐poor headwater streams. We quantified input rates of terrestrial insects in eight streams draining lowland tropical wet forest in Costa Rica. In two focal headwater streams, we also measured capture efficiency by the fish assemblage and quantified terrestrially derived N‐ and P‐excretion relative to stream nutrient uptake rates. Average input rates of terrestrial insects ranged from 5 to 41 mg dry mass/m²/d, exceeding previous measurements of aquatic invertebrate secondary production in these study streams, and were relatively consistent year‐round, in contrast to values reported in temperate streams. Terrestrial insects accounted for half of the diet of the dominant fish species, Priapicthys annectens. Although terrestrially derived fish excretion was found to be a small flux relative to measured nutrient uptake rates in the focal streams, the efficient capture and processing of terrestrial arthropods by fish made these nutrients available to the local stream ecosystem. This aquatic‐terrestrial linkage is likely being decoupled by deforestation in many tropical regions, with largely unknown but potentially important ecological consequences.     

Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

Nutrient uptake in streams draining agricultural catchments of the midwestern United States

1. Agriculture is a major contributor of non‐point source pollution to surface waters in the midwestern United States, resulting in eutrophication of freshwater aquatic ecosystems and development of hypoxia in the Gulf of Mexico. Agriculturally influenced streams are diverse in morphology and have variable nutrient concentrations. Understanding how nutrients are transformed and retained within agricultural streams may aid in mitigating increased nutrient export to downstream ecosystems. 2. We studied six agriculturally influenced streams in Indiana and Michigan to develop a more comprehensive understanding of the factors controlling nutrient retention and export in agricultural streams using nutrient addition and isotopic tracer studies. 3. Metrics of nutrient uptake indicated that nitrate uptake was saturated in these streams whereas ammonium and phosphorus uptake increased with higher concentrations. Phosphorus uptake was likely approaching saturation as evidenced by decreasing uptake velocities with concentration; ammonium uptake velocity also declined with concentration, though not significantly. 4. Higher whole‐stream uptake rates of phosphorus and ammonium were associated with the observed presence of stream autotrophs (e.g. algae and macrophytes). However, there was no significant relationship between measures of nutrient uptake and stream metabolism. Water‐column nutrient concentrations were positively correlated with gross primary production but not community respiration. 5. Overall, nutrient uptake and metabolism were affected by nutrient concentrations in these agriculturally influenced streams. Biological uptake of ammonium and phosphorus was not saturated, although nitrate uptake did appear to be saturated in these ecosystems. Biological activity in agriculturally influenced streams is higher relative to more pristine streams and this increased biological activity likely influences nutrient retention and transport to downstream ecosystems.     

去叶时间对半干旱草原植物养分回收和干草生产的影响

去叶时间对半干旱草原植物养分回收和干草生产的影响 养分回收是植物养分保存的重要策略,其对环境和管理变化的响应关系到生态系统的养分循环和生产。去叶(刈割)是影响草地植物养分回收和生产的重要途径,而去叶时间的影响尚不清楚。本研究以内蒙古典型草原生态系统为对象,设置早期去叶(生物量高峰期之前)、峰期去叶(生物量高峰期)、晚期去叶(养分回收开始后)和不去叶(对照)四个处理,探讨了去叶时间对植物养分回收和生产的影响。通过测定植物物种和群落水平氮(N)和磷(P)回收特征,量化了植物N、P回收通量以及凋落物归还通量和干草输出通量,并评估了不同去叶时间处理下割草草地系统干草产量和质量。研究结果显示,峰期和晚期去叶降低植物群落N、P回收度,而早期去叶则对二者无影响;不同去叶时间处理下植物N、P回收效率相对稳定,仅晚期去叶降低N回收效率。峰期和晚期去叶降低植物群落N、P回收通量和凋落物N、P归还通量,而早期去叶并不影响这些参数。去叶时间降低植物群落养分回收通量,但未改变植物根系养分储存,说明根系养分吸收增加可补偿养分回收通量的降低。草地干草产量和质量在峰期去叶处理下最高,晚期去叶处理下最低。本研究结果为割草草地生态系统养分循环提供了新见解,通过调整刈割时间可以平衡草原的保护与生产,在植物生物量高峰期之前割草可实现保护和可持续生产的双重目标。     

Roles of benthic algae in the structure,function, and assessment of stream ecosystems affected by acid mine drainage

Tens of thousands of stream kilometers worldwide are degraded by a legacy of acid loads, high metal concentrations, and altered habitat caused by acid mine drainage (AMD) from abandoned underground and surface mines. As the primary production base in streams, the condition of algal‐dominated periphyton communities is particularly important to nutrient cycling, energy flow, and higher trophic levels. Here, we synthesize current knowledge regarding how AMD‐associated stressors affect (i) algal communities and their use as ecological indicators, (ii) their functional roles in stream ecosystems, and (iii) how these findings inform management decisions and evaluation of restoration effectiveness. A growing body of research has found ecosystem simplification caused by AMD stressors. Species diversity declines, productivity decreases, and less efficient nutrient uptake and retention occur as AMD severity increases. New monitoring approaches, indices of biological condition, and attributes of algal community structure and function effectively assess AMD severity and effectiveness of management practices. Measures of ecosystem processes, such as nutrient uptake rates, extracellular enzyme activities, and metabolism, are increasingly being used as assessment tools, but remain in their infancy relative to traditional community structure‐based approaches. The continued development, testing, and implementation of functional measures and their use alongside community structure metrics will further advance assessments, inform management decisions, and foster progress toward restoration goals. Algal assessments will have important roles in making progress toward improving and sustaining the water quality, ecological condition, and ecosystem services of streams in regions affected by the legacy of unregulated coal mining.     

不同基因型甘蔗种质资源的表型遗传多样性

植物的碳、氮、磷化学计量特征能反映植物对土壤营养元素的利用效率,岩溶区植物经过长期的进化形成了自身独特的生理生态和生态化学计量特征,通过岩溶区植物叶片碳、氮、磷化学计量可以揭示岩溶生态系统各组分之间的养分循环规律.该研究在桂林毛村岩溶区次生林中选择3个20 m×20 m 的样方,采用多元统计方法分析了岩溶区森林12种典型植物叶片共186个样品的碳、氮、磷的生态化学计量特征,研究它们之间的相互关系,探讨碳、氮、磷化学计量学在岩溶生态系统中的生态指示作用.结果表明：(1)虽然岩溶区石灰土氮和磷的含量较高,但由于其有效性低,植物对养分的吸收和利用缓慢,岩溶区石灰土植物的生长仍然受到 N 和 P 的共同限制;(2)由于岩溶区植物叶片中 N 和 P 的含量显著偏低导致较高的 C∶N 和 C∶P 值(C∶N 的平均值为80．86;C∶P 的平均值为639．65);(3)利用 N∶P＜14表明氮受限制,N∶P＞16表明磷受限制,14＜N∶P＜16表明 N 和 P 共同限制的标准判断植物叶片受 N 或 P 的限制在岩溶区不完全适合;(4)元素间相关性分析表明,叶片的 C 和 N 呈极显著负相关关系(P ＜0．01),C 和 P 呈显著负相关关系(P ＜0．05),N 和 P之间呈现极显著正相关关系(P ＜0．01).这体现了植物体内两营养元素含量需求变化的相对一致性.研究结果有助于了解岩溶区森林植物的适生机制及其生态地球化学过程,可为岩溶区生态治理提供理论依据.     

Photosynthesis and growth reduction with warming are driven by nonstomatal limitations in a Mediterranean semi‐arid shrub

Whereas warming enhances plant nutrient status and photosynthesis in most terrestrial ecosystems, dryland vegetation is vulnerable to the likely increases in evapotranspiration and reductions in soil moisture caused by elevated temperatures. Any warming‐induced declines in plant primary production and cover in drylands would increase erosion, land degradation, and desertification. We conducted a four‐year manipulative experiment in a semi‐arid Mediterranean ecosystem to evaluate the impacts of a ~2°C warming on the photosynthesis, transpiration, leaf nutrient status, chlorophyll content, isotopic composition, biomass growth, and postsummer survival of the native shrub Helianthemum squamatum. We predicted that warmed plants would show reduced photosynthetic activity and growth, primarily due to the greater stomatal limitation imposed by faster and more severe soil drying under warming. On average, warming reduced net photosynthetic rates by 36% across the study period. Despite this strong response, warming did not affect stomatal conductance and transpiration. The reduction of peak photosynthetic rates with warming was more pronounced in a drought year than in years with near‐average rainfall (75% and 25–40% reductions relative to controls, respectively), with no indications of photosynthetic acclimation to warming through time. Warmed plants had lower leaf N and P contents, δ¹³C, and sparser and smaller leaves than control plants. Warming reduced shoot dry mass production by 31%. However, warmed plants were able to cope with large reductions in net photosynthesis, leaf area, and shoot biomass production without changes in postsummer survival rates. Our findings highlight the key role of nonstomatal factors (biochemical and/or nutritional) in reducing net carbon assimilation rates and growth under warming, which has important implications for projections of plant carbon balance under the warmer and drier climatic scenario predicted for drylands worldwide. Projected climate warming over the coming decades could reduce net primary production by about one‐third in semi‐arid gypsum shrublands dominated by H. squamatum.     

Biomass loss and change in species dominance shift stream community excretion stoichiometry during severe drought

1. Animals contribute significantly to nutrient cycling through excretion, but most studies consider their effects under relatively benign abiotic conditions. Disturbances such as drought may alter animals’ nutrient contributions through shifts in species composition and biomass. Headwater streams are particularly vulnerable to extreme climate events and thus might show rapid changes in stream biota and their ecosystem effects.
2. We tested how biomass and subsequent ecosystem effects (nutrient cycling) of an intermittent prairie stream community changed during a drought. We quantified the biomass and contributions to nutrient cycling for assemblages comprising fishes, crayfish, and tadpoles in 12 isolated pools over 3 months encompassing the harshest drought on record for Kings Creek, KS, U.S.A. We predicted that macroconsumer biomass would decline with pool surface area and that differences in macroconsumer biomass and taxonomic composition would lead to different contributions of pool assemblages to nutrient cycling.
3. The biomass of pool assemblages declined with decreasing pool size, a pattern apparently driven by mortality, emigration, or metamorphosis. We also observed a change in assemblage structure of drying pools during drought relative to pool size, shifting dominance toward species with more drought-resistant traits. Accordingly, assemblage nitrogen (N) excretion rates declined as pool biomass was reduced, leading to a 58% reduction in N available to epilithic biofilms. Phosphorus (P) excretion rates declined from June to July, but increased in August, as species with high P excretion rates maintained similar proportional biomass and biomass of a non-native fish increased. Molar N:P of pool assemblage excretion declined significantly throughout the drought and coincided with loss of southern redbelly dace (Chrosomus erythrogaster: Cyprinidae).
4. Animal-mediated nutrient cycling was altered by the loss of biomass and stoichiometric traits of taxa that differed in their occurrences and ability to tolerate abiotic conditions during drought. Elevated availability of dissolved N in isolated pools may increase N uptake rates by biofilms during drought conditions, indicating the importance of N excreted by aggregated macroconsumers, especially those with unique stoichiometric traits. While the significance of shifts in the composition of freshwater communities to ecosystems is not entirely known, additional losses in ecosystem function and changes in community structure may follow episodes of severe drought.

     

Nitrogen and phosphorus translocation of forest floor mosses as affected by a pulse of these nutrients

藓类是亚高山森林、北方针叶林和极地苔原等原始生态系统的重要组成部分。养分沉降是这些生态系统的主要养分来源。然而,对 藓类如何利用大气沉降这一时间上异质性较高的养分来源的认识缺乏,限制了我们对藓类在生态系统养分循环中作用的认识,也制约了模 拟和预测生态系统服务功能动态变化的有效性。本研究中,我们重点关注如下问题：(1)藓类新生部分的氮、磷含量随氮或磷沉降量的变化; (2)氮或磷养分沉降如何影响藓类的氮磷转运;(3)氮磷转运与藓类新生部分养分含量的关联性？为此,我们在位于青藏高原东部的亚高山 冷杉林中开展了设置有不同氮、磷施加量水平的突发养分沉降实验。以林下优势度最高的塔藓和锦丝藓为对象,我们在施加实验处理8天和1年后,分别测定了不同年龄藓类茎段中的氮、磷含量。研究结果发现,养分处理8天后,锦丝藓和塔藓的所有茎段的氮或磷含量均随处理中氮或磷施加量的增加而增加,表明它们能够高效率地吸收施加的氮或磷。养分处理1年后,形成于养分处理后的新生茎段中的氮、磷含量以及氮磷比与处理后8天的藓类茎段中氮、磷含量以及氮磷比呈正相关关系,表明藓类将吸收的养分转运至新生部分,转运过程受源—库关系的调控。磷转运效率解释了新生茎段21%–23%的氮磷比变化,而氮转运与新生茎段氮磷比没有显著关系,结果暗示磷转运对藓类生长的重要性。上述结果表明,养分转运是藓类植物利用间断的沉降养分的关键机制,也使藓类成为生态系统中重要的养分库。     

Tight Coupling Between Shoot Level Foliar N and P,Leaf Area,and Shoot Growth in Arctic Dwarf Shrubs Under Simulated Climate Change

Nutrient availability limits productivity of arctic ecosystems, and this constraint means that the amount of nitrogen (N) in plant canopies is an exceptionally strong predictor of vegetation productivity. However, climate change is predicted to increase nutrient availability leading to increases in carbon sequestration and shifts in community structure to more productive species. Despite tight coupling of productivity with canopy nutrients at the vegetation scale, it remains unknown how species/shoot level foliar nutrients couple to growth, or how climate change may influence foliar nutrients–productivity relationships to drive changes in ecosystem carbon gain and community structure. We investigated the influence of climate change on arctic plant growth relationships to shoot level foliar N and phosphorus (P) in three dominant subarctic dwarf shrubs using an 18-year warming and nutrient addition experiment. We found a tight coupling between total leaf N and P per shoot, leaf area and shoot extension. Furthermore, a steeper shoot length-leaf N relationship in deciduous species (Vaccinium myrtillus and Vaccinium uliginosum) under warming manipulations suggests a greater capacity for nitrogen to stimulate growth under warmer conditions in these species. This mechanism may help drive the considerable increases in deciduous shrub cover observed already in some arctic regions. Overall, our work provides the first evidence at the shoot level of tight coupling between foliar N and P, leaf area and growth i.e. consistent across species, and provides mechanistic insight into how interspecific differences in alleviation of nutrient limitation will alter community structure and primary productivity in a warmer Arctic.     

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.     

Biotic and abiotic controls on the ecosystem significance of consumer excretion in two contrasting tropical streams

1. Excretion of nitrogen (N) and phosphorus (P) is a direct and potentially important role for aquatic consumers in nutrient cycling that has recently garnered increased attention. The ecosystem‐level significance of excreted nutrients depends on a suite of abiotic and biotic factors, however, and few studies have coupled measurements of excretion with consideration of its likely importance for whole‐system nutrient fluxes. 2. We measured rates and ratios of N and P excretion by shrimps (Xiphocaris elongata and Atya spp.) in two tropical streams that differed strongly in shrimp biomass because a waterfall excluded predatory fish from one site. We also made measurements of shrimp and basal resource carbon (C), N and P content and estimated shrimp densities and ecosystem‐level N and P excretion and uptake. Finally, we used a 3‐year record of discharge and NH₄‐N concentration in the high‐biomass stream to estimate temporal variation in the distance required for excretion to turn over the ambient NH₄‐N pool. 3. Per cent C, N, and P body content of Xiphocaris was significantly higher than that of Atya. Only per cent P body content showed significant negative relationships with body mass. C:N of Atya increased significantly with body mass and was higher than that of Xiphocaris. N : P of Xiphocaris was significantly higher than that of Atya. 4. Excretion rates ranged from 0.16–3.80 μmol NH₄‐N shrimp^?1 h^?1, 0.23–5.76 μmol total dissolved nitrogen (TDN) shrimp^?1 h^?1 and 0.002–0.186 μmol total dissolved phosphorus (TDP) shrimp^?1 h^?1. Body size was generally a strong predictor of excretion rates in both taxa, differing between Xiphocaris and Atya for TDP but not NH₄‐N and TDN. Excretion rates showed statistically significant but weak relationships with body content stoichiometry. 5. Large between‐stream differences in shrimp biomass drove differences in total excretion by the two shrimp communities (22.3 versus 0.20 μmol NH₄‐N m^?2 h^?1, 37.5 versus 0.26 μmol TDN m^?2 h^?1 and 1.1 versus 0.015 μmol TDP m^?2 h^?1), equivalent to 21% and 0.5% of NH₄‐N uptake and 5% and <0.1% of P uptake measured in the high‐ and low‐biomass stream, respectively. Distances required for excretion to turn over the ambient NH₄‐N pool varied more than a hundredfold over the 3‐year record in the high‐shrimp stream, driven by variability in discharge and NH₄‐N concentration. 6. Our results underscore the importance of both biotic and abiotic factors in controlling consumer excretion and its significance for nutrient cycling in aquatic ecosystems. Differences in community‐level excretion rates were related to spatial patterns in shrimp biomass dictated by geomorphology and the presence of predators. Abiotic factors also had important effects through temporal patterns in discharge and nutrient concentrations. Future excretion studies that focus on nutrient cycling should consider both biotic and abiotic factors in assessing the significance of consumer excretion in aquatic ecosystems.