Response of grazing snails to phosphorus enrichment of modern stromatolitic microbial communities

1. The effects of added phosphorus (P) on the growth, P and RNA : DNA contents, and survivorship of snails grazing on laminated microbial mats (living ‘stromatolites’) were examined in the Rio Mesquites at Cuatro Ciénegas, Mexico (total P, c. 0.60 μmol L^?1) to test the hypothesis that strong P‐limitation of microautotroph growth produces a stoichiometric constraint on herbivores because of mineral P‐limitation. 2. In a 3‐week experiment performed in summer 2001, addition of phosphorus (+15 μmol L^?1) resulted in a strong decline in stromatolite biomass C : P ratio from very high levels (c. 2300 : 1 by atoms) to moderate levels (c. 550 : 1). The endemic hydrobiid snail Mexithauma quadripaludium responded to P‐enrichment with elevated body P content and higher RNA : DNA ratios, especially for small animals likely to be actively growing. This positive response is consistent with the existence of a stoichiometric constraint on snail growth. 3. In a longer experiment (8 weeks) involving a more moderate P enrichment (+5 μmol L^?1) in summer 2002, P enrichment reduced stromatolite C : P ratio from moderate values in control treatments (c. 750) to very low values (<100 : 1). Snails responded to stromatolite P‐enrichment with increased body P content but, in contrast to the first experiment, with lower RNA : DNA ratio, lower growth rates, and higher mortality. 4. These contrasting results suggest that both very high and very low biomass C : P ratios in stromatolites are detrimental to M. quadripaludium performance, leading us to hypothesise that these herbivores live on a ‘stoichiometric knife edge’.     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Microbial nitrogen and phosphorus co-limitation across permafrost region

The status of plant and microbial nutrient limitation have profound impacts on ecosystem carbon cycle in permafrost areas, which store large amounts of carbon and experience pronounced climatic warming. Despite the long-term standing paradigm assumes that cold ecosystems primarily have nitrogen deficiency, large-scale empirical tests of microbial nutrient limitation are lacking. Here we assessed the potential microbial nutrient limitation across the Tibetan alpine permafrost region, using the combination of enzymatic and elemental stoichiometry, genes abundance and fertilization method. In contrast with the traditional view, the four independent approaches congruently detected widespread microbial nitrogen and phosphorus co-limitation in both the surface soil and deep permafrost deposits, with stronger limitation in the topsoil. Further analysis revealed that soil resources stoichiometry and microbial community composition were the two best predictors of the magnitude of microbial nutrient limitation. High ratio of available soil carbon to nutrient and low fungal/bacterial ratio corresponded to strong microbial nutrient limitation. These findings suggest that warming-induced enhancement in soil nutrient availability could stimulate microbial activity, and probably amplify soil carbon losses from permafrost areas.     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

Carbon use efficiency of microbial communities: stoichiometry,methodology and modelling

Carbon use efficiency (CUE) is a fundamental parameter for ecological models based on the physiology of microorganisms. CUE determines energy and material flows to higher trophic levels, conversion of plant‐produced carbon into microbial products and rates of ecosystem carbon storage. Thermodynamic calculations support a maximum CUE value of ~ 0.60 (CUE _max). Kinetic and stoichiometric constraints on microbial growth suggest that CUE in multi‐resource limited natural systems should approach ~ 0.3 (CUE _max/2). However, the mean CUE values reported for aquatic and terrestrial ecosystems differ by twofold (~ 0.26 vs. ~ 0.55) because the methods used to estimate CUE in aquatic and terrestrial systems generally differ and soil estimates are less likely to capture the full maintenance costs of community metabolism given the difficulty of measurements in water‐limited environments. Moreover, many simulation models lack adequate representation of energy spilling pathways and stoichiometric constraints on metabolism, which can also lead to overestimates of CUE. We recommend that broad‐scale models use a CUE value of 0.30, unless there is evidence for lower values as a result of pervasive nutrient limitations. Ecosystem models operating at finer scales should consider resource composition, stoichiometric constraints and biomass composition, as well as environmental drivers, to predict the CUE of microbial communities.     

Global forest carbon uptake due to nitrogen and phosphorus deposition from 1850 to 2100

Spatial patterns and temporal trends of nitrogen (N) and phosphorus (P) deposition are important for quantifying their impact on forest carbon (C) uptake. In a first step, we modeled historical and future change in the global distributions of the atmospheric deposition of N and P from the dry and wet deposition of aerosols and gases containing N and P. Future projections were compared between two scenarios with contrasting aerosol emissions. Modeled fields of N and P deposition and P concentration were evaluated using globally distributed in situ measurements. N deposition peaked around 1990 in European forests and around 2010 in East Asian forests, and both increased sevenfold relative to 1850. P deposition peaked around 2010 in South Asian forests and increased 3.5‐fold relative to 1850. In a second step, we estimated the change in C storage in forests due to the fertilization by deposited N and P (?C_{ν dep}), based on the retention of deposited nutrients, their allocation within plants, and C:N and C:P stoichiometry. ?C_{ν dep} for 1997–2013 was estimated to be 0.27 ± 0.13 Pg C year^?1 from N and 0.054 ± 0.10 Pg C year^?1 from P, contributing 9% and 2% of the terrestrial C sink, respectively. Sensitivity tests show that uncertainty of ?C_{ν dep} was larger from P than from N, mainly due to uncertainty in the fraction of deposited P that is fixed by soil. ?C_{P dep} was exceeded by ?C_{N dep} over 1960–2007 in a large area of East Asian and West European forests due to a faster growth in N deposition than P. Our results suggest a significant contribution of anthropogenic P deposition to C storage, and additional sources of N are needed to support C storage by P in some Asian tropical forests where the deposition rate increased even faster for P than for N.     

Nitrogen and phosphorus content of some temperate and tropical freshwater fishes

The nitrogen and phosphorus content of two temperate fishes, Rutilus rutilus and Perca fluviatilis , and six tropical fishes, Oreochromis niloticus , Cichla monoculus , Serrassalmus rhombeus , Plagioscion squamosissimus , Prochilodus brevis and Hoplias malabaricus , were investigated to test the hypothesis that variation in body P content and N:P ratio is related to body size. Regressions of %P and N:P ratios against fish size (length and mass) confirmed the hypothesis for P. fluviatilis and P. squamosissimus , suggesting that body size is an important factor driving body P content and N:P ratios in some fishes. Moreover, significant increases in %N and N:P ratio with body size was found for H. malabaricus , a common piscivorous fish of the Neotropics. Interspecific variation in %P and N:P ranged two-fold and significant differences ( P < 0·05) were found among the tested species. The mean ± s . d . elemental content across all fishes ( n = 170) was 10·35 ± 1·29% for N and 3·05 ± 0·82% for P, while the N:P ratio was 8·00 ± 2·14. Data on fish body nutrient content and ratio will improve parameterization of bioenergetics and mass balance models and help clarify the role of fishes in nutrient cycles in both temperate and tropical freshwaters.     

土壤酶计量揭示了武夷山黄山松林土壤微生物沿海拔梯度的碳磷限制变化

了解土壤胞外酶活性和酶计量的变化对评估山地生态系统土壤养分有效性和微生物的营养限制状况具有重要意义.然而,亚热带山地森林土壤微生物的营养限制状况对海拔梯度变化的响应及其驱动因素尚不清楚.本研究以武夷山不同海拔(1200～2000 m)黄山松林为对象,测定了土壤基本性质、微生物生物量以及与碳(C)、氮(N)、磷(P)循环...     

Atmospheric nitrogen deposition is associated with elevated phosphorus limitation of lake zooplankton

Here, we present data that for the first time suggests that the effects of atmospheric nitrogen (N) deposition on nutrient limitation extend into the food web. We used a novel and sensitive assay for an enzyme that is over‐expressed in animals growing under dietary phosphorus (P) deficiency (alkaline phosphatase activity, APA) to assess the nutritional status of major crustacean zooplankton taxa in lakes across a gradient of atmospheric N deposition in Norway. Lakes receiving high N deposition had suspended organic matter (seston) with significantly elevated carbon:P and N:P ratios, indicative of amplified phytoplankton P limitation. This P limitation appeared to be transferred up the food chain, as the cosmopolitan seston‐feeding zooplankton taxa Daphnia and Holopedium had significantly increased APA. These results indicate that N deposition can impair the efficiency of trophic interactions by accentuating stoichiometric food quality constraints in lake food webs.     

Influence of phosphorus concentration on the growth kinetics and stoichiometry of the microalga Scenedesmus obliquus

The growth of the freshwater microalga Scenedesmus obliquus was studied at 30°C in a mineral culture medium with phosphorus concentrations of between 0 and 372 μ . The values for the specific growth rates, between and , fitted a semistructured substrate-limitation model with μ_m1 = 0·0466 h⁻¹, μ_m2 = 0·0256 h⁻¹ and . The specific uptake rate of phosphorus reached a maximum value of q_Sm1 = 658·01 × 10⁻⁴ μmol P mg⁻¹ biomass h⁻¹.     

Effects of resource supplies on the structure and function of microbial communities

The supplies of nutrients, and their elemental stoichiometry, can have significant impacts upon the structure and function of microbial communities. This review focuses on the effects of nutrient supplies on the biodegradation of organic matter, and on the dynamics of host-pathogen interactions. Analyses of data from the literature suggest significant effects of nitrogen:phosphorus supply ratios on the microbial decomposition of organic matter, and it is argued that these stoichiometric effects may have important implications for the fate and fluxes of carbon in natural ecosystems. In addition, it is shown that the supplies of nitrogen and phosphorus to the host can strongly influence the outcome of infections in both terrestrial and aquatic plants, suggesting that resource availability and resource supply ratios potentially may have significant effects on the health of many plant communities. This revised version was published online in August 2006 with corrections to the Cover Date.     

模拟酸沉降对南亚热带季风常绿阔叶林土壤微生物群落结构的长期影响

土壤微生物是生态系统重要的组成成分, 尤其是在土壤风化严重, 养分贫瘠的热带和南亚热带森林生态系统中, 微生物在植物养分的获取、碳循环以及土壤的形成等生态过程中的作用尤为重要。该研究基于鼎湖山南亚热带季风常绿阔叶林长期(10年)的野外模拟酸沉降实验平台, 探究了土壤微生物群落结构对土壤酸化的响应。结果表明, 酸沉降处理显著降低土壤pH (即加剧酸化)。土壤酸化对微生物生物量碳(C)含量的影响不大, 但改变了土壤微生物生物量氮(N)和磷(P)的含量, 导致表层土壤(0-10 cm)微生物生物量C:P和N:P显著提高, 表明土壤酸化可能加剧了微生物P限制。土壤酸化还显著改变了土壤微生物群落结构, 导致次表层土壤(10-20 cm)真菌:细菌显著增加。进一步分析表明, 土壤pH和土壤有效P含量是影响土壤微生物群落最为主要的两个因素。     

Bacteria as a source of phosphorus for zooplankton

The utilization of bacterial phosphorus in zooplankton metabolism was investigated using radio-phosphorus labelled natural bacteria as food source for zooplankton in feeding experiments. Incorporation of labelled bacteria was clearly related to the species' ability to graze on bacteria, with the cladoceran Daphnia reaching the highest biomass-specific activity and the copepod Acanthodiaptomus the lowest. Within Daphnia, juveniles had a higher biomass-specific uptake of phosphorus than adults. This was presumably caused by higher growth rates of the juveniles rather than age-specific differences in the ability to feed on bacteria, supported by the observation that the juveniles had the highest specific P-content. Retention of ingested ³²P from labelled particles exceeded 80%, indicating higher assimilation efficiencies on phosphorus compared with carbon. In the investigated humic lake, approximately 75% of the phosphorus in grazable particles was bound in bacterial cells, making bacteria the most important source of P to the bacterivore zooplankton species.     

Nutrient co-limitation of primary producer communities

Synergistic interactions between multiple limiting resources are common, highlighting the importance of co-limitation as a constraint on primary production. Our concept of resource limitation has shifted over the past two decades from an earlier paradigm of single-resource limitation towards concepts of co-limitation by multiple resources, which are predicted by various theories. Herein, we summarise multiple-resource limitation responses in plant communities using a dataset of 641 studies that applied factorial addition of nitrogen (N) and phosphorus (P) in freshwater, marine and terrestrial systems. We found that more than half of the studies displayed some type of synergistic response to N and P addition. We found support for strict definitions of co-limitation in 28% of the studies: i.e. community biomass responded to only combined N and P addition, or to both N and P when added separately. Our results highlight the importance of interactions between N and P in regulating primary producer community biomass and point to the need for future studies that address the multiple mechanisms that could lead to different types of co-limitation.     

Effects of nutrient (N,P, C) enrichment,grazing and depth upon littoral periphyton of a softwater lake

Three field experiments were performed in Lake Lacawac, PA to determine the importance of potentially limiting nutrients relative to other factors (grazing, depth) in structuring shallow water algal periphyton communities. All three experiments measured periphyton growth (as chlorophyll-a, AFDM or biovolumes of the algal taxa) on artificial clay flower pot substrates which released specified nutrients to their outer surfaces.Control of standing crop by nutrient supply rate vs. grazing was examined in Expt. I. Substrates releasing excess N and P, together with one of 4 levels of C (as bicarbonate) were placed either inside or outside exclosures designed to reduce grazer densities. Chlorophyll-a rose from 1.1–25.6 µg.cm^–2, and some dominant taxa (e.g., Oedogonium, Nostoc, Anacystis) were replaced by others (e.g., Scenedesmus, Cryptomonas) as bicarbonate supply increased. Reductions in invertebrate density did not significantly affect chlorophyll-a at any of the nutrient levels.Reasons for the species shift were further evaluated in Expt. II, using a minielectrode to measure the elevation of pH within the periphyton mat through photosynthetic utilization of bicarbonate. The pH adjacent to pots diffusing N, P and large quantities of bicarbonate, and supporting high chlorophyll-a densities of 32 µg cm^–2, averaged 10.0 compared to 6.3 in the water column. Pots diffusing only N and P supported 0.7 µg chlorophyll-a cm^–2 and elevated pH to 8.2. We suspect that bicarbonate addition favored efficient bicarbonate users (e.g., Scenedesmus), while inhibiting other taxa (e.g., Oedogonium) because of the attendant high pH.Expt. III was designed to test effects of depth (0.1 m vs. 0.5 m) and N (NH₄ ⁺ vs. NO₃ ^–) upon the growth response to bicarbonate observed in Expts. I and II. Similar standing crop and species composition were noted on pots at 0.1 m vs. 0.5 m. Enrichment with NH₄ ⁺ vs. NO₃ ^– also appeared to have little effect upon the periphyton community.Shallow water periphyton communities in Lake Lacawac, when supplied with sufficient N and P, appear to show a distinctive response to increasing bicarbonate concentration and pH which is robust to moderate variation in grazer densities, distance from the water surface, and the form of N enrichment.     

黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征的影响

研究黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征影响有助于深入理解黄土丘陵区不同植被带下土壤和土壤微生物相互作用及养分循环规律.选择黄土丘陵区延河流域3个植被区(森林区、森林草原区、草原区)和5种地形部位(阴/阳沟坡、阴/阳梁峁坡、峁顶)的土壤作为研究对象,利用生态化学计量学理论研究植被和地形对土壤和土壤微生物生物量生态化学计量特征的影响.结果表明: 土壤及土壤微生物生物量碳、氮、磷含量在不同地形之间的差别主要表现在沟坡位置和阴坡高于其他坡位和阳坡.植被类型的变化对两个土层(0～10、10～20 cm)土壤和土壤微生物生物量碳、氮、磷的影响均达到显著水平,坡向对表层(0～10 cm)土壤和土壤微生物生物量碳、氮、磷的影响强于坡位,而在10～20 cm土层,坡位对土壤和土壤微生物生物量碳、氮、磷影响更显著.植被类型显著影响土壤C∶N、C∶P、N∶P和土壤微生物生物量C∶N、C∶P,坡向和坡位仅影响土壤C∶P和N∶P,植被类型的变化是影响土壤C∶N的主要因素.同时,植被类型对土壤养分和微生物生物量碳、氮、磷含量及其生态化学计量特征的影响大于地形因子.标准化主轴分析结果表明,黄土丘陵区不同植被带土壤微生物具有内稳性,特别在草原带,土壤微生物生物量生态化学计量学特征具有更加严格的约束比例.在黄土丘陵区,土壤微生物生物量N∶P或许可以作为判断养分限制的另一个有力工具,若将土壤微生物生物量N∶P与植物叶片N∶P配合使用可能有助于我们更加精确地判断黄土丘陵区的土壤养分限制情况.     

长期施磷对玉米-小麦轮作系统作物产量和磷素吸收及土壤磷积累的影响

本研究利用位于河北省保定市的7年田间定位试验,探讨了 4个磷水平(不施磷、70％优化施磷、优化施磷、130％优化施磷)对玉米-小麦轮作系统作物产量的影响,分析了作物磷素吸收和磷肥利用效率以及土壤磷盈亏的变化.结果表明:长期(7年)施磷能够显著提高玉米和小麦产量及磷素吸收量,且玉米和小麦产量与磷素吸收量随着施磷量的增加均...