Plasticity in Resource Allocation and Nitrogen-use Efficiency in Riparian Vegetation: Implications for Nitrogen Retention

In this work, we summarize our current understanding of the function of riparian zones and describe an investigation of changes in the production per unit nitrogen (N) taken up, or nitrogen-use efficiency (NUE) and resource allocation of a riparian shrub in response to changes in N availability. Empirical work included measuring leaf %N and root-to-shoot ratios (R:S) of individual riparian shrubs (Baccharis salicifolia, or seepwillow) growing at a range of N availabilities in the field and growing in fertilized and unfertilized plots in a field fertilization experiment. In both observational and experimental work, N availability was related positively to %N of plant tissues and negatively to R:S. We used a simulation model to investigate feedbacks between seepwillow responses to and effects on N availability. In the model, plasticity in resource allocation and NUE in response to changes in N led to lower productivity at low N supply and higher productivity and lower retention at high N supply than was observed in plants constrained to a constant %N and R:S. Furthermore, uptake became relatively more important as a retention mechanism when plants responded to high N supply. These feedbacks could have significant effects on N retention by riparian zones in watersheds receiving large fertilizer inputs of N or on ecosystems exposed to high rates of atmospheric N deposition.     

Sources of Nitrogen to the Riparian Zone of a Desert Stream: Implications for Riparian Vegetation and Nitrogen Retention

Riparian zones effectively remove nitrogen (N) from water flowing through riparian soils, particularly in agricultural watersheds. The mechanism of N removal is still unclear, especially the role of vegetation. Uptake and denitrification are the two most commonly studied mechanisms. Retention of groundwater N by plant uptake is often inferred from measurements of N in net incremental biomass. However, this assumes other sources of N are not contributing to the N demand of plants. The purpose of this work was to investigate the relative importance of three sources of available N to riparian trees in a desert stream—input in stream water during floods, input during baseflow, and mineralization of N from soil organic matter. Two approaches were used; a mass balance approach in which the mass of available N from each source was estimated, and a correlational approach in which indexes of each source were compared to leaf N for individual willow trees. Total N from all sources was 396 kg ha⁻¹ y⁻¹, with 172 kg ha⁻¹ y⁻¹ from mineralization, 214 kg ha⁻¹ y⁻¹ from the stream during baseflow, and 9.6 kg ha⁻¹ y⁻¹ from floods. Leaf N was significantly related to N mineralization rates and flood inputs; it was not related to baseflow inputs. We conclude that mineralization is a major source of available N for willow trees, subsidized by input of N from floods. Baseflow inputs are most likely removed by rapid denitrification at the stream–riparian edge, while higher rates of flood supply exceed the capacity of this “filter.” Received 18 January 2001; accepted 15 June 2001.     

Retention of nitrogen and phosphorus in a Danish lowland river system: implications for the export from the watershed

In a Danish lowland river system intensive measurements were made, in four 80 m reaches, of the nitrogen (N) and phosphorus (P) stored in the stream sediment. The results were used for calculation of the total retention in the river system during two summers (June to August). In addition, the mobilization of nutrients from the stream bottom in autumn 1987 was compared with the export from the watershed.During the study period (June 1987 to September 1988) the amounts of N and P stored in stream reaches were determined fortnightly using a core-sample technique. In reaches dominated by submersed macrophytes, 25–40 g N m^–2 and 20–30 g P m^–2 were stored during two summers, against only 10–15g N and P m^–2 for sandy and gravely reaches. In riparian zones with emergent macrophytes the retention was even higher than in the submersed macrophytes. Gross retention exceeded net retention by a factor of two to three.Net retention of P in the river system during the summer of 1987 was equal to the summer export from the watershed. On an annual basis, retention in the summer constituted 20% of the P export. In contrast, retention in the summer of 1988 amounted to 60% of the total P export during the same period (38% reduction) and 22% in comparison with the annual export. The corresponding figures for N were lower, showing reductions of 16% and 12% of the export of total N in the two summer periods, and about 1% of the annual exports.In September 1987 6.4% of the total N export and 65% of the total P export from the watershed consisted of resuspended material. In 1987 the N and P retained during the summer was almost completely resuspended during storm events during September to November.     

Denitrification Potential, Root Biomass, and Organic Matter in Degraded and Restored Urban Riparian Zones

Hydrologic changes associated with urbanization often lead to lower water tables and drier, more aerobic soils in riparian zones. These changes reduce the potential for denitrification, an anaerobic microbial process that converts nitrate, a common water pollutant, into nitrogen gas. In addition to oxygen, denitrification is controlled by soil organic matter and nitrate. Geomorphic stream restorations are common in urban areas, but their effects on riparian soil conditions and denitrification have not been evaluated. We measured root biomass, soil organic matter, and denitrification potential (anaerobic slurry assay) at four depths in duplicate degraded, restored, and reference riparian zones in the Baltimore, Maryland, U.S.A., metropolitan area. There were three main findings in this study. First, although reference sites were wet and had high soil organic matter, they had low levels of nitrate relative to degraded and restored sites and therefore there were few differences in denitrification potential among sites. Evaluations of riparian restorations that have nitrate removal by denitrification as a goal should consider the complex controls of this process and how they vary between sites. Second, all variables declined markedly with depth in the soil. Restorations that increase riparian water tables will thus foster interaction of groundwater nitrate with near-surface soils with higher denitrification potential. Third, we observed strong positive relationships between root biomass and soil organic matter and between soil organic matter and denitrification potential, which suggest that establishment of deep-rooted vegetation may be particularly important for increasing the depth of the active denitrification zone in restored riparian zones.     

Effects of nutrients and arbuscular mycorrhizal colonization on the growth of Salix gracilistyla seedlings in a nutrient-poor fluvial bar

A field survey and a pot culture experiment were conducted to examine the effects of nutrients (N and P) and arbuscular mycorrhizal (AM) fungi on the growth of Salix gracilistyla, a pioneer plant in riparian habitats. The plants growing in the field were colonized by AM and/or ectomycorrhizal fungi. However, the direct effect of AM colonization on seedling growth was not detected in the pot culture experiment. In contrast, N application significantly promoted plant growth, suggesting that the growth of S. gracilistyla seedlings is largely limited by the availability of N in the field.     

河岸带农田不同恢复年限对土壤碳氮磷生态化学计量特征的影响——以温榆河为例

弃耕地撂荒是土壤与植被向自然方向进行的次生演替,研究河岸带土壤撂荒后碳氮磷生态化学计量特征,是恢复和重建由农田干扰导致的退化河岸带生态系统的重要科学基础之一。以河岸带农地为对照,不同撂荒年限(撂荒2年、撂荒8年、撂荒10年)的土壤为研究对象,探索不同撂荒年限对土壤碳、氮、磷含量及相互关系的影响。结果表明:(1)土壤有机碳、氮的含量均呈现撂荒10年>撂荒8年>农田>撂荒2年;土壤中磷含量呈现撂荒10年>撂荒8年>撂荒2年>农田;农田和各撂荒年限的土壤碳、氮、磷含量,均随着土层深度的增加而呈降低的规律,但土壤碳和氮差异的显著性比磷明显。(2)河岸带土壤中C/N、C/P的均值均呈现:撂荒10年>农田>撂荒8年>撂荒2年趋势。N/P的均值呈现:撂荒10年(0.78)>农田(0.77)>撂荒8年(0.77)>撂荒2年(0.67),表明N是本研究区河岸带植被恢复的限制性营养元素。(3)河岸带农田和不同撂荒年限土壤碳、氮含量均存在极显著的耦合线性关系,而碳与磷、氮与磷之间的线性拟合程度相对较低。(4)在农田撂荒演替的初期阶段(2...     

Drainage history and land use pattern of a Swedish river system — their importance for understanding nitrogen and phosphorus load

During the 19th and the first half of the 20th century, approximately 300 km² of lakes and wetlands, representing 29% of the River Kavlingean catchment in Southern Sweden, were drained to make land available for agriculture. Published accounts of nutrient loads from the catchment indicated that until the mid 20th century, factories and urban point sources were the major contributors of both nitrogen and phosphorus. By the middle of the 20th century, the construction of sewage treatment plants had effectively reduced phosphorus pollution. Concurrently, the land drained in the previous century underwent a more intense cultivation, with productivity being maintained by commercial fertilizers. Subsequently, net nutrient loads from agriculture continued to increase, reaching an annual load of 2652 tons total-nitrogen and 70 tons total-phosphorus for the River Kävlingeån. Whilst high nutrient leakage from agricultural watersheds may be a problem which is only recently recognized, it had its origins in nearly a hundred years of commonly accepted agricultural policy.To assess the importance of agriculture as the major source of nutrients to the River Kävlingeån system, three tributary catchment areas, differing in terms of their land use patterns (high, medium and low intensity of agricultural use), were studied and compared with literature figures. Results indicated that agricultural nutrient loss areal coefficients were substantially higher than the literature figures, demonstrating the role of agriculture as source of nutrients to the River Kävlingeån system. The agricultural land use policies of the last fifty years were revealed to be most important with regard to this role. Of such land use policies, the cultivation of the last 10–15% of land employed for agricultural use (primarily riparian ecotones) may be of most significance. The literature indicates that intense agricultural use of this final 10–15% may account for a ca. 50% increase in nitrogen loss. This suggests that one solution to the problem of agricultural diffuse pollution may lie in the restoration and sustainable management of riparian ecotones of agricultural streams.     

Lake restoration using a reed swamp to remove nutrients from non-point sources

The reed swamp adjacent to Lake Glumsø was partly separated from the lake by a dam and the inflow to the reed swamp controlled by pumping from the tributary. An investigation of the nutrient balances for the reed swamp showed an average daily denitrification rate of 252 mg/m2 from August 1988 to March 1989. Mineralization of the peat took place simultaneously, in sufficient amounts to supply carbon for the denitrification process. Ammonia and phosphorus were released in the ratio 7:1, corresponding to 111 mg N and 16 mg P/m²/ day. Most of the ammonia was nitrified and denitrified (included in the above mentioned denitrification rate). The net release of phosphorus implies that the method only can be applied satisfactorily in situations where nitrogen is the limiting nutrient. This is the case for Lake Glumso during the summer period, but not during the spring.This paper was presented at the INTECOL IV International Wetlands Conference in Columbus, Ohio, 1992, as part of a session organized by Prof. S. E. Jørgensen and sponsored by the International Lake Environment Committee.Corresponding Editor: Prof K. R. Reddy     

Nitrogen retention in wetlands,lakes and rivers

As human activities continue to alter the global nitrogen cycle, the ability to predict the impact of increased nitrogen loading to freshwater systems is becoming more and more important. Nitrogen retention is of particular interest because it is through its combined processes (denitrification, nitrogen sedimentation and uptake by aquatic plants) that local and downstream nitrogen concentrations are reduced. Here, we compare the magnitude of nitrogen retention and its components in wetlands, lakes and rivers. We show that wetlands retain the highest proportion of total nitrogen loading, followed by lakes and then rivers. The differences in the proportion of N retained among systems is explained almost entirely by differences in water discharge. Denitrification is the primary mechanism of nitrogen retention, followed by nitrogen sedimentation and uptake by aquatic plants.     

The Influence of Land Use on Lake Nutrients Varies with Watershed Transport Capacity

Nutrient loading to lakes depends on both the availability of nutrients in a watershed and their potential for movement to a lake. Many studies have demonstrated that variation in watershed land use can translate to differences in lake water quality by affecting nutrient availability. There have been few attempts, however, to understand how loading to surface waters is affected by land use when there are differences in watershed transport capacity. We compared the relationship between land use/cover and lake nutrients in lakes draining watersheds that exhibited high and low transport capacity using a 5 year (2001–2005) dataset describing the chemistry of 101 lakes and reservoirs in a region of intensive agriculture. We measured watershed transport capacity by compositing the hydrologic, geologic, and topographic variables correlated with interannual variability in lake total nitrogen (TN) or phosphorus (TP) because the hydrologic permeability of watersheds amplifies downstream responses to rainfall events. Factors describing watershed transport capacity differed for TN and TP, consistent with differences in nutrient mobility and biogeochemistry. Partial least squares regression revealed that watershed transport capacity influenced the nature of the association between land use/cover and lake chemistry. In watersheds with low transport capacity, in-lake processes and near-shore land use/cover tended to be more influential, whereas, in watersheds with high transport capacity, land use/cover across the entire watershed was important for explaining lake chemistry. Thus, although land use is a key driver of nutrient loading to lakes, the extent to which it influences water quality can vary with watershed transport capacity. JMF conceived the study and analyzed the data. JAD collected the data. JMF and JAD wrote the paper.     

Particulate phosphorus sedimentation at the river inflow to a lake

Sedimentation at the Krutynia River inflow to Lake Kujno was closely related to hydrological regime. The highest sedimentation rates, recorded in spring, decreased during summer by two orders of magnitude. Granulometric segregation of settling seston along the inflow zone was related to differentiation of phoshorus content. Smaller particles were richer in P, producing a gradient of increasing P concentrations in the settling material. A substantial loss of P from polyphosphate and various organic fractions was found after settlement of river suspensoids. Mechanisms of P losses are discussed and possibilities of P retention within the inflow zone are considered.     

Nitrogen Retention, Removal, and Saturation in Lotic Ecosystems

Increased nitrogen (N) loading to lotic ecosystems may cause fundamental changes in the ability of streams and rivers to retain or remove N due to the potential for N saturation. Lotic ecosystems will saturate with sustained increases in the N load, but it is unclear at what point saturation will occur. Rates of N transformation in lotic ecosystems will vary depending on the total N load and whether it is an acute or chronic N load. Nitrogen saturation may not occur with only pulsed or short-term increases in N. Overall, saturation of microbial uptake will occur prior to saturation of denitrification of N and denitrification will become saturated prior to nitrification, exacerbating increases in nitrate concentrations and in N export downstream. The rate of N export to downstream ecosystems will increase proportionally to the N load once saturation occurs. Long term data sets showed that smaller lotic ecosystems have a greater capacity to remove in-stream N loads, relative to larger systems. Thus, denitrification is likely to become less important as a N loss mechanism as the stream size increases. There is a great need for long-term studies of N additions in lotic ecosystems and clear distinctions need to be made between ecosystem responses to short-term or periodic increases in N loading and alterations in ecosystem functions due to chronic N loading.     

The influence of leaf litter on zooplankton in floodplain wetlands: changes resulting from river regulation

1. The loss of input of leaf litter through clearing of riparian vegetation may result in significant changes to aquatic ecosystems. River red gums (Eucalyptus camaldulensis) surrounding floodplain wetlands in the Murray–Darling Basin, Australia, contribute large quantities of leaf litter, but the quality of this resource may change depending on the timing of inundation. 2. We used experimental mesocosms to test the hypotheses that zooplankton would have a greater abundance with an input of leaf litter and that fewer zooplankton would emerge from egg banks in cleared than forested wetlands. The experiment was carried out in summer/autumn and in spring to test a third hypothesis that zooplankton would respond to changes in the timing of wetland inundation as a result of river regulation. 3. In summer/autumn, leaf litter reduced zooplankton abundance by 89% at the beginning of the experiment through its influence on water quality. Only a few taxa (Polyarthra spp., Colurella spp. and the cladoceran Family Moinidae) responded positively to leaf litter when water quality improved later in the experiment, indicating a switch in the role of leaf litter from a non‐trophic to a trophic pathway. 4. In spring, microcrustaceans emerged in smaller numbers from sediment sourced from cleared compared to forested wetlands, reflecting different communities in these two wetland types and/or disturbances to the sediment that interfere with emergence. 5. Although leaf litter appears not to be an important resource for zooplankton in floodplain wetlands, riparian clearing may have lasting effects on future emerging zooplankton communities. Additionally, river regulation may have considerable impacts on the influence of leaf litter on zooplankton, which has implications for the management of floodplain river systems.     

The use of Bayesian networks to guide investments in flow and catchment restoration for impaired river ecosystems

1.  The provision of environmental flows and the removal of barriers to water flow are high priorities for restoration where changes to flow regimes have caused degradation of riverine ecosystems. Nevertheless, flow regulation is often accompanied by changes in catchment and riparian land-use, which also can have major impacts on river health via local habitat degradation or modification of stream energy regimes.
2.  The challenges are determining the relative importance of flow, land-use and other impacts as well as deciding where to focus restoration effort. As a consequence, flow, catchment and riparian restoration efforts are often addressed in isolation. River managers need decision support tools to assess which flow and catchment interventions are most likely to succeed and, importantly, which are cost-effective.
3.  Bayesian networks (BNs) can be used as a decision support tool for considering the influence of multiple stressors on aquatic ecosystems and the relative benefits of various restoration options. We provide simple illustrative examples of how BNs can address specific river restoration goals and assist with the prioritisation of flow and catchment restoration options. This includes the use of cost and utility functions to assist decision makers in their choice of potential management interventions.
4.  A BN approach facilitates the development of conceptual models of likely cause and effect relationships between flow regime, land-use and river conditions and provides an interactive tool to explore the relative benefits of various restoration options. When combined with information on the costs and expected benefits of intervention, one can derive recommendations about the best restoration option to adopt given the network structure and the associated cost and utility functions.     

Retention of soluble organic nutrients by a forested ecosystem

We document an example of a forested watershed at the Coweeta HydrologicLaboratory with an extraordinary tendency to retain dissolved organic matter(DOM) generated in large quantities within the ecosystem. Our objectives weretodetermine fluxes of dissolved organic C, N, and P (DOC, DON, DOP,respectively),in water draining through each stratum of the ecosystem and synthesizeinformation on the physicochemical, biological and hydrologic factors leadingtoretention of dissolved organic nutrients in this ecosystem. The ecosystemretained 99.3, 97.3, and 99.0% of water soluble organic C, N and P,respectively, produced in litterfall, throughfall, and root exudates. Exportsinstreamwater were 4.1 kg ha^–1yr^–1of DOC, 0.191 kg ha^–1 yr^–1 ofDON, and 0.011 kg ha^–1 yr^–1 ofDOP. Fluxes of DON were greater than those of inorganic N in all strata. MostDOC, DON, and DOP was removed from solution in the A and B horizons, with DOCbeing rapidly adsorbed to Fe and Al oxyhydroxides, most likely by ligandexchange. DON and DOC were released gradually from the forest floor over theyear. Water soluble organic C produced in litterfall and throughfall had adisjoint distribution of half-decay times with very labile and veryrefractory fractions so that most labile DOC was decomposed before beingleachedinto the mineral soil and refractory fractions dominated the DOC transportedthrough the ecosystem. We hypothesize that this watershed retained solubleorganic nutrients to an extraordinary degree because the soils have very highcontents of Fe and Al oxyhydroxides with high adsorption capacities and becausethe predominant hydrologic pathway is downwards as unsaturated flow through astrongly adsorbing A and B horizon. The well recognized retention mechanismsforinorganic nutrients combine with adsorption of DOM and hydrologic pathway toefficiently prevent leaching of both soluble inorganic andorganic nutrients in this watershed.     

The cycling of nutrients in a closed-basin antarctic lake: Lake Vanda

Lake Vanda is a permanently ice covered, meromictic, closed basin lake, located in the Dry Valley region of Southern Victoria Land, Antarctica. A unique feature of the lake water column structure is that the bottom lake waters exist as a natural diffusion cell. The diffusive nature of these waters allows rates of sulfate reduction, nitrification and denitrification to be calculated from nutrient concentration gradients. Calculation reveals that sulfate reduction is by far the most important anoxic process acting to oxidize organic material. In addition, rate calculations reveal that bottom water nutrient profiles are in steady state. One argument in support of this conclusion is that the calculated rate of nitrification balances the flux of ammonia from the anoxic lake waters. The flux of phosphorus from the reducing waters is several times less than would be predicted from the nitrogen and phosphorus content of decomposing lake seston. Solubility calculations show that phosphorus may be actively removed at depth in Lake Vanda by the formation of hydroxyapatite. It is found that estimated rates of nitrogen and phosphorus removal in the bottom lake waters and sediments roughly balance the riverine input flux. This suggests that throughout the lake a nutrient steady state may exist, and that the anoxic zone may be the most important loci for nutrient removal. Finally, the ratio of nitrogen to phosphorus entering Lake Vanda by riverine input is less than the Redfield ratio of 16/1; in contrast to the lake waters which are strongly phosphorus limited at all depths. This curious aspect of the lake's nutrient chemistry is explained by the presence of preformed nitrogen, which has been concentrated in the deep brine due to several episodes of evaporative concentration.     

Influence of soil nutrients on ectomycorrhizal communities in a chronosequence of mixed temperate forests

Many factors associated with forests are collectively responsible for controlling ectomycorrhizal (ECM) fungal community structure, including plant species composition, forest structure, stand age, and soil nutrients. The objective of this study was to examine relationships among ECM fungal community measures, local soil nutrients, and stand age along a chronosequence of mixed forest stands that were similar in vegetation composition and site quality. Six combinations of age class (5-, 26-, 65-, and 100-year-old) and stand initiation type (wildfire and clearcut) were replicated on four sites, each representing critical seral stages of stand development in Interior Cedar-Hemlock (ICH) forests of southern British Columbia. We found significant relationships between ECM fungal diversity and both available and organic P; available P was also positively correlated with the abundance of two ECM taxa (Rhizopogon vinicolor group and Cenoccocum geophilum). By contrast, ECM fungal diversity varied unpredictably with total and mineralizable N or C to N ratio. We also found that soil C, N, available P, and forest floor depth did not exhibit strong patterns across stand ages. Overall, ECM fungal community structure was more strongly influenced by stand age than specific soil nutrients, but better correlations with soil nutrients may occur at broader spatial scales covering a wider range of site qualities.