Nitrogen stable isotope ratios in surface sediments, epilithon and macrophytes from upland lakes with differing nutrient status

1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were each visited once during June and July 2000. From each lake, samples of surface sediment epilithon, macrophytes and total dissolved nitrogen (TDN) were collected for nitrogen stable isotope analysis. As part of a wider programme, samples were also collected for chemical analysis and bioassays. 2. Considerable variation was found in δ¹⁵N values in all measured nitrogen compartments. Some regional variation was evident but was generally weak. Sediment and epilithon δ¹⁵N were positively correlated with δ¹⁵N of TDN, suggesting that baseline nitrogen isotope ratios influence those in some organic matter compartments in the lakes. 3. Sediment δ¹⁵N was higher when inorganic nitrogen concentration in the water was low, possibly reflecting reduced isotope fractionation under these conditions. However, this was not the case for epilithon or macrophytes. Sediment δ¹⁵N values were also negatively related to annual nitrogen deposition. 4. Sediment, epilithon and macrophyte δ¹⁵N values all showed significant relations to nutrient limitation in the lakes as determined by algal bioassays. We suggest that sediment δ¹⁵N might be developed as a simple integrating measure of the degree of nitrogen limitation in lakes.     

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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Using porewater profiles to assess nutrient availability in seagrass-vegetated carbonate sediments

We measured porewater profiles of inorganic (NH₄ ⁺, NO₃ ^–(+NO₂ ^–), PO₄ ^3– (hereafter referred to as DIP)) and organic (DON, DOP) nutrients in seagrass-vegetated sediments at two sites in a shallow bay in Bermuda within close proximity (200 m) but subject to different nutrient loading. At both sites, total dissolved and inorganic nutrient concentrations were usually 1–2 orders of magnitude higher in the sediments than in the water column, with the exception of NO₃ ^–. Organic N and P were significant components of the total dissolved nutrient pools both in the sediment porewater and in the overlying water column (up to 75% for DON and 40% for DOP), and may be important in meeting plant nutrient demands. We used two approaches to examine how well porewater nutrient concentrations reflected the relative availabilities of N and P for seagrasses: (1) a simple stoichiometric nutrient regeneration model based on the N:P ratio of decomposing organic matter and porewater NH₄ ⁺ concentrations to predict porewater DIP, and (2) fitting of the porewater profiles to estimate rates of net nutrient production (or consumption), which reflects the balance between nutrient sources and sinks in the rhizosphere. The stoichiometric model indicated that sediment porewaters were depleted in P relative to N in the low-nutrient outer bay site, and enriched in P relative to N in the higher-nutrient inner bay site. These results are consistent with the mechanism of carbonate sediments in oligotrophic tropical environments being a strong sink for dissolved inorganic P and our previous work suggesting that nutrient enrichment causes P to become disproportionately more available than N. Net nutrient production rates of porewater P at both sites and N at the inner bay site were low (typically < 2%) relative to the nutrient demands of the seagrasses. The implications of the profile interpretation are two-fold: (1) the low rates of net nutrient production indicate diffusive losses from the root zone were insignificant and that nutrient turnover rates were high, except in the P-limited outer bay where N accumulated in sediment porewaters; and (2) because standing stock nutrient concentrations often represent a small fraction of the total nutrients cycled in the sediments, they are in many cases a poor indicator of nutrient availability. Based on our estimates of losses from the root zone, decomposition, and plant uptake we have constructed a rough budget for the cycling of P and N at our two sites.     

Assessment of nutrient availability and limitation using macroalgae

Discs of the macroalga,Ulva lactuca L., were transplanted around an ocean outfall and at a reference site in Køge Bay, Denmark, to assess the influence of the outfall on the nutrient availability. At 2-wk intervals, samples were collected and analyzed for growth, nitrogen, and phosphorus content.The tissue concentrations of nitrogen and phosphorus decreased with distance to the outfall, showing that the tissue concentrations are suitable for monitoring nutrient availability in coastal areas and provide a time-integrated measure of the nutrient availability. The lowest tissue concentrations of nitrogen were recorded at the reference station, where the internal concentrations generally were below the critical concentration level, showing that nitrogen limited the growth. At the station located close to the outfall, the flux of nitrogen was sufficient to maintain the maximum growth rate. The tissue concentrations of phosphorus were only below the critical concentration level on one occasion, and the result showed a net uptake throughout the study period.It was concluded that in the Køge Bay, nitrogen was the main limiting factor for macroalgae growth during the summer. The applicability of tissue concentrations for assessment of nutrient availability is discussed and it is considered that the method, when evaluated against established critical concentrations, provides a valuable tool for assessing ecosystem health with regard to eutrophication.     

Lacustrine sediment/groundwater nutrient dynamics

A small pond in southwestern peninsular Florida was sampled to determine the areal and depth distribution of total phosphorus, total nitrogen, and organic matter. Concentrations of these constituents decreased from the center to the edge of the pond and from the top to the bottom of the sediments. At the center of the pond total phosphorus showed a secondary concentration peak at a depth of 20 to 40 cm in the sediments indicative of the downward transfer of soluble phosphorus during times of low water.A system of monitoring wells was installed at the pond to measure groundwater input and output nutrient concentrations. There was no difference in total groundwater nitrogen concentration across the pond but the mean increase in total phosphorus concentration between input and output wells, ca. 200 g1^–1, was statistically highly significant. The results suggest that whereas nitrogen is recycled from sediments to the pond and the atmosphere, the pond-sediment system loses some phosphorus to groundwater throughflow.     

Variable rates of phosphate uptake by shallow marine carbonate sediments: Mechanisms and ecological significance

We determined phosphate uptake by calcareous sediments at two locations within a shallow lagoon in Bermuda that varied in trophic status, with one site being mesotrophic and the other being more eutrophic. Phosphate adsorption over a six hour period was significantly faster in sediments from the mesotrophic site. Uptake at both sites was significantly less than that reported for a similar experiment on calcareous sediments in an oligotrophic lagoon in the Bahamas. The difference in phosphorus adsorption between our sites did not appear to be related to sediment characteristics often cited as important, such as differences in surface area (as inferred from grain size distributions), total organic matter content, or iron content. However, the sediment total phosphorus contents were inversely related to phosphorus uptake at our sites in Bermuda, and at the previously studied Bahamas site.We hypothesize that phosphate uptake in these calcareous sediments is a multi-step process, as previously described for fluvial sediments or pure calcium carbonate solids, with rapid initial surface chemisorption followed by a slower incorporation into the carbonate solid-phase matrix. Accordingly, sediments already richer in solid phase phosphorus take up additional phosphate more slowly since the slower incorporation of surface-adsorbed phosphate into the carbonate matrix limits the rate of renewal of surface-reactive adsorption sites.Although carbonate sediments are a sink for phosphate, and thereby reduce the availability of phosphorus for benthic macrophytes and phytoplankton in the shallow overlying water, phosphate uptake by these sediments appears to decrease along a gradient from oligotrophic to eutrophic sites. If our result is general, it implies a positive feedback in phosphorus availability, with a proportionately greater percentage of phosphorus loading being biologically available longer as phosphorus loading increases. This pattern is supported by the significantly higher tissue phosphorus content of the seagrass,Thalassia testudinum, collected from the eutrophic inner bay site. Over time, this effect may tend to cause a shift from phosphorus to nitrogen limitation in some calcareous marine environments.     

Phosphatase activity and nitrogen fixation reflect species differences,not nutrient trading or nutrient balance,across tropical rainforest trees

A fundamental biogeochemical paradox is that nitrogen‐rich tropical forests contain abundant nitrogen‐fixing trees, which support a globally significant tropical carbon sink. One explanation for this pattern holds that nitrogen‐fixing trees can overcome phosphorus limitation in tropical forests by synthesizing phosphatase enzymes to acquire soil organic phosphorus, but empirical evidence remains scarce. We evaluated whether nitrogen fixation and phosphatase activity are linked across 97 trees from seven species, and tested two hypotheses for explaining investment in nutrient strategies: trading nitrogen‐for‐phosphorus or balancing nutrient demand. Both strategies varied across species but were not explained by nitrogen‐for‐phosphorus trading or nutrient balance. This indicates that (1) studies of these nutrient strategies require broad sampling within and across species, (2) factors other than nutrient trading must be invoked to resolve the paradox of tropical nitrogen fixation, and (3) nitrogen‐fixing trees cannot provide a positive nitrogen‐phosphorus‐carbon feedback to alleviate nutrient limitation of the tropical carbon sink.     

Effects of nutrient additions on litter production and nutrient return in a nutrient-poor Cape fynbos ecosystem

Litter production and N and P return were determined at bimonthly intervals for two years in 10×5 m plots, amended with a complete factorial fertilizer addition of N as NH₄NO₃(N_a), P as Ca₃(PO₄)₂(P_a) and a mixture of all essential nutrients excluding N and P (M_a) in a 4–7-year-old post-fire sand-plain lowland fynbos ecosystem, South Africa. Litter production increased with vegetation age, was highly seasonal and peaked from late spring to mid-summer (November to January). No significant differences in annual litter production and N return were found in response to the nutrient treatments, although both tended to increase during the second year in response to N_a and M_a. Phosphorus return increased significantly with P_a, and to a lesser extend, N₃, during the first year, whereas it increased in response to N_a and M_a and decreased in the P_a amended plots during the second year. The nutrient treatments did not result in a change in the timing of the annual peak litter production period or in the plant growth form composition of the litter. The litter layer dry mass and N and P contents increased in response to N_a and M_a, while P_a resulted in an increased P content. The evidence from this study indicates that the vegetative growth of the evergreen sclerophyllous shrubs and hemicryptophytes of sand-plain lowland fynbos is not only limited by N, as shown by other studies on shoot growth and vegetation cover, but also by one or more other nutrients excluding P.     

Release rates and potential fates of nitrogen and phosphorus from sediments in a eutrophic reservoir

1. Nutrients released from lake sediments can influence water column nutrient concentrations and planktonic productivity. We examined sediment nutrient release [soluble reactive phosphorus (SRP) and ammonia (NH)] at two sites in a eutrophic reservoir (Acton Lake, OH, U.S.A.) that differed in physical mixing conditions (a thermally stratified and an unstratified site). 2. Sediment nutrient release rates were estimated with three methods: sediment core incubations, seasonal in situ hypolimnetic accumulation and a published regression model that predicted sediment phosphorous (P) release rate from sediment P concentration. All three methods were applied to the deeper stratified site in the reservoir; however, we used only sediment core incubations to estimate SRP and NH release rates at the shallow unstratified site because of the lack of thermal stratification. We also compared the total P concentration (TP_S) of sediments and the concentration of P in various sediment fractions at both sites. 3. Anoxic sediments at the stratified site released SRP at rates more than an order of magnitude greater than oxic sediments at the shallow unstratified site. However, P accumulated in the hypolimnion at much lower rates than predicted by sediment core incubations. In contrast, NH was released at similar rates at both sites and accumulated in the hypolimnion at close to the expected rate, indicating that P was ‘lost’ from the hypolimnion through biogeochemical pathways for P, such as precipitation with inorganic material or biological uptake and sedimentation. 4. TP_S was significantly greater at the deeper stratified site and organically bound P accounted for >50% of TP_S at both sites. 5. We examined the magnitude of SRP fluxes into the study reservoir in 1996 by comparing the mean summer daily SRP fluxes from anaerobic sediments, aerobic sediments, stream inflows and gizzard shad excretion. While the SRP release from anaerobic sediments was high, we hypothesise that little of this SRP gained access to the epilimnion in mid‐summer. SRP flux to the reservoir from aerobic sediments was less than from gizzard shad excretion and streams. Large interannual variability in thermocline stability, gizzard shad biomass and stream discharge volumes, will affect SRP loading rates from different sources in different years. Therefore, construction of P budgets for different years should account for interannual variation in these parameters.     

Physiological indicators of nutrient deficiency in phytoplankton in southern Chilean lakes

We assessed the nutrient status of phytoplankton in 28 lakes in southern Chile using two types of physiological indicators: specific alkaline phosphatase activity, and the elemental composition (carbon, nitrogen, and phosphorus) of seston. Alkaline phosphatase activity ranged from 0.001 to 0.11 mol P g chl^–1 h^–1, with P-deficiency indicated in about one-half the study lakes. C:N ranged from 3.9 to 24, C:P ranged from 86 to 919, and N:P ranged from 8.7 to 99. C:P and N:P ratios greater than the Redfield ratio were common, suggesting P deficiency in many of the lakes. C:N ratios were not generally indicative of N deficiency. Previous studies have suggested N may be the primary limiting nutrient in southern Chilean lakes, but our results indicate that P should not be discounted as a limiting nutrient.     

Importance of sediments in understanding nutrient cyclings in lakes

Inorganic and organic nutrients are continuously transported to lake bottoms by sedimentation. By various biological, physical, chemical and mechanical processes quantities of certain nutrients can be brought back to the free water again. This cycling between the sediments and water may occur according to various schemes dependent on lake type and bottom conditions. Lake morphology, temperature regimes, trophic level and sediment type can all strongly influence the size of nutrient pools and rates of turnover.The various activities of bacteria, benthic algae, macrophytes, benthic invertebrates and fish, in conjunction with influences of temperature, pH-values, Eh-values, water content, organic matter and elemental sediment composition, lead to the extremely complex nature of nutrient cycling. Three essential components of aquatic ecosystems are discussed, namely carbon, nitrogen and phosphorus.The objective of this paper is to illustrate in condensed form the heterogeneous nature of nutrient cycling processes. In addition, the importance of sediments in understanding nutrient cycling is discussed from a water management perspective.     

Nutrient limitation of phytoplankton in a central Arizona reservoir

The influence of added nutrients nitrogen, phosphorus and carbon on the phytoplankton of a small recreational reservoir in central Arizona was investigated during the summer, 1974. Polyethylene bags were used to isolate lake water and the natural populations for the addition of nitrogen, phosphorus and carbon individually and in combination. A large increase in phytoplankton numbers, extractable chlorophyll, pH and dissolved oxygen occurred only in bags to which both nitrogen and phosphorus were added, suggesting that both nitrogen and phosphorus levels were limiting to the primary producers. Little alteration in species composition resulted from the addition of the above nutrients.     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

Responses of benthic algae to nutrient enrichment in a shallow lake: Linking community production,biomass, and composition

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Nutrient limitation of phytoplankton production in Alaskan Arctic foothill lakes

We used 54 enrichment bioassays to assess nutrient limitation (N, P) of ¹⁴C uptake by natural phytoplankton assemblages in 39 lakes and ponds in the Arctic Foothills region of Alaska. Our purpose was to categorize phytoplankton nutrient status in this under-represented region of North America and to improve our ability to predict the response of primary production to anticipated anthropogenically mediated increases in nutrient loading. Experiments were performed across several watersheds and included assays on terminal lakes and lakes occupying various positions in chains (lakes in series within a watershed and connected by streams). In total, 89% (48 of 54) of the bioassays showed significant stimulation of ¹⁴C primary production by some form of nutrient addition relative to unamended controls. A significant response was observed following enrichment with N and P, N alone and P alone in 83, 35 and 22% of the bioassays, respectively. In experiments where N and P proved stimulatory, the influence of N alone was significantly greater than the influence of P alone. Overall, the data point to a greater importance for N than P in regulating phytoplankton production in this region. The degree of response to N and P enrichment declined as the summer progressed and showed no relationship to irradiance or water temperature, suggesting secondary limitation by some micronutrient such as iron as the summer advanced. Phytoplankton nutrient status was often consistent across lakes within a watershed, suggesting that watershed characteristics influence nutrient availability. Lakes in this region will clearly show increased phytoplankton production in response to anthropogenic activities and anticipated changes in climate that will increase nutrient loading.     

Eutrophication and macroalgal blooms in temperate and tropical coastal waters: nutrient enrichment experiments with Ulva spp.

Receiving coastal waters and estuaries are among the most nutrient‐enriched environments on earth, and one of the symptoms of the resulting eutrophication is the proliferation of opportunistic, fast‐growing marine seaweeds. Here, we used a widespread macroalga often involved in blooms, Ulva spp., to investigate how supply of nitrogen (N) and phosphorus (P), the two main potential growth‐limiting nutrients, influence macroalgal growth in temperate and tropical coastal waters ranging from low‐ to high‐nutrient supplies. We carried out N and P enrichment field experiments on Ulva spp. in seven coastal systems, with one of these systems represented by three different subestuaries, for a total of nine sites. We showed that rate of growth of Ulva spp. was directly correlated to annual dissolved inorganic nitrogen (DIN) concentrations, where growth increased with increasing DIN concentration. Internal N pools of macroalgal fronds were also linked to increased DIN supply, and algal growth rates were tightly coupled to these internal N pools. The increases in DIN appeared to be related to greater inputs of wastewater to these coastal waters as indicated by high δ¹⁵N signatures of the algae as DIN increased. N and P enrichment experiments showed that rate of macroalgal growth was controlled by supply of DIN where ambient DIN concentrations were low, and by P where DIN concentrations were higher, regardless of latitude or geographic setting. These results suggest that understanding the basis for macroalgal blooms, and management of these harmful phenomena, will require information as to nutrient sources, and actions to reduce supply of N and P in coastal waters concerned.     

Experimental evidence for weakened tree nutrient use and resorption efficiencies under severe drought in a subtropical monsoon forest

日益频发的干旱严重威胁着全球森林生态系统的功能。由于干旱胁迫抑制了土壤养分可利用性,因此植物的养分利用和重吸收效率对 森林生态系统的功能以及生物地球化学循环至关重要。然而,目前对于植物养分利用和重吸收效率在干旱条件下的动态响应规律的理解十分 有限,且在(亚)热带区域尤为明显。因此,本研究主要探讨了在湿热森林中不同植物物种在不同月份遭受干旱胁迫时,其养分利用和重吸收 效率的重要性是否会发生变化以及如何变化。本研究在2016年10月–2019年5月期间,依托一处位于亚热带常绿阔叶林的隔离70%自然穿透 雨的干旱实验平台,采集了木荷(Schima superba)和石栎(Lithocarpus glaber)两个物种在不同月份的鲜叶及凋落叶,用于分析干旱对两种植物叶片氮磷利用和重吸收效率的影响(NUE和PUE,NRE和PRE)。研究结果表明,干旱对氮磷利用和重吸收效率的作用在不同植物物种和月份之间有差异。基于两年的观测结果显示,干旱对木荷的养分利用和重吸收效率无显著影响,却使石栎的NUE、NRE和PRE分别降低了3.4%、 20.2%和7.1%。另外,2017 年夏季发生的自然干旱进一步加剧了干旱对石栎养分重吸收的负作用。在2017年8月,石栎的NUE和PUE在干旱处理下分别降低了17.2%和58.1%,而NRE和PRE分别下降了56.5%和53.8%。此外,石栎的NRE,PRE和NUE对干旱的响应显著依赖于土壤水分条件,即当土壤湿度降至约9 v/v%时,存在一个阈值使干旱处理的效果从无影响转变为负作用。我们的结果表明,在干旱条件下亚热带常绿阔叶林树木的养分利用呈现出了物种特定的阈值响应。