The importance of nutrient co‐limitation in regulating algal community composition,productivity and algal‐derived DOC in an oligotrophic marsh in interior Alaska

1. Compared to lakes and streams, we know relatively little about the factors that regulate algae in freshwater wetlands. This discrepancy is particularly acute in boreal regions, where wetlands are abundant and processes related to climate change (i.e. increased permafrost collapse and soil weathering) are expected to increase nutrient inputs into aquatic systems. To investigate how accelerated nutrient inputs might affect algal structure and function in northern boreal wetlands, we added nitrogen, phosphorus and silica to mesocosms in an oligotrophic marsh in interior Alaska. 2. We conducted two in situ mesocosm enrichment experiments during consecutive summer growing seasons, each lasting 24 days. In 2007, we investigated the effects of +N, +P, +Si and +N+P+Si enrichment on benthic algal biomass (chlorophyll‐a, ash‐free dry mass, biovolume), chemistry (N : P ratio) and community composition. In 2008, we expanded our first experiment to investigate the effects +N+P, +N+Si, +P+Si and +N+P+Si on the same algal parameters as well as productivity (mg C m^?2 h^?1). 3. In both experiments, we measured water‐column dissolved organic carbon (DOC) inside treatment enclosures and related changes in DOC to standing algal biomass. 4. Benthic algal accrual did not increase following 24 days of enrichment with any nutrient alone or with P and Si together (+P+Si), but increased significantly with the addition of N in any combination with P and Si (+N+P, +N+Si, +N+P+Si). 5. Algal productivity (20 mg C m^?2 h^?1) increased between three‐ and seven‐fold (57–127 mg C m^?2 h^?1) with the addition of N in combination with any other nutrient (+N+P, +N+Si, +N+P+Si). Water‐column DOC concentration was significantly higher inside N‐combination treatments compared to the control during each season, and DOC increased linearly with benthic algal biomass in 2007 (r² = 0.89, P < 0.0001) and 2008 (r² = 0.74, P < 0.0001). 6. Taxonomic composition of the wetland algal community responded most strongly to N‐combination treatments in both seasons. In 2007, there was a significant shift from Euglena and Mougeotia in the control treatment to Chroococcus and Gloeocystis with +N+P+Si enrichment, and in 2008, a Mougeotia‐dominated community was replaced by Gloeocystis in the +N+P treatment and by Nitzschia in +N+Si and +N+P+Si treatments. 7. Together, these data provide several lines of evidence for co‐limitation, and the central importance of N as a co‐limiting nutrient for the wetland algal community. Changes in algal dynamics with increased nutrient concentrations could have important implications for wetland food webs and suggest that algae may provide a functional link between increasing nutrient inputs and altered wetland carbon cycling in this region.     

Interactive Effects of Fire, Soil Climate, and Moss on CO2 Fluxes in Black Spruce Ecosystems of Interior Alaska

Fire is an important control on the carbon (C) balance of the boreal forest region. Here, we present findings from two complementary studies that examine how fire modifies soil organic matter properties, and how these modifications influence rates of decomposition and C exchange in black spruce (Picea mariana) ecosystems of interior Alaska. First, we used laboratory incubations to explore soil temperature, moisture, and vegetation effects on CO₂ and DOC production rates in burned and unburned soils from three study regions in interior Alaska. Second, at one of the study regions used in the incubation experiments, we conducted intensive field measurements of net ecosystem exchange (NEE) and ecosystem respiration (ER) across an unreplicated factorial design of burning (2 year post-fire versus unburned sites) and drainage class (upland forest versus peatland sites). Our laboratory study showed that burning reduced the sensitivity of decomposition to increased temperature, most likely by inducing moisture or substrate quality limitations on decomposition rates. Burning also reduced the decomposability of Sphagnum-derived organic matter, increased the hydrophobicity of feather moss-derived organic matter, and increased the ratio of dissolved organic carbon (DOC) to total dissolved nitrogen (TDN) in both the upland and peatland sites. At the ecosystem scale, our field measurements indicate that the surface organic soil was generally wetter in burned than in unburned sites, whereas soil temperature was not different between the burned and unburned sites. Analysis of variance results showed that ER varied with soil drainage class but not by burn status, averaging 0.9 ± 0.1 and 1.4 ± 0.1 g C m⁻² d⁻¹ in the upland and peatland sites, respectively. However, a more complex general linear model showed that ER was controlled by an interaction between soil temperature, moisture, and burn status, and in general was less variable over time in the burned than in the unburned sites. Together, findings from these studies across different spatial scales suggest that although fire can create some soil climate conditions more conducive to rapid decomposition, rates of C release from soils may be constrained following fire by changes in moisture and/or substrate quality that impede rates of decomposition. Author contributions: JAO: performed research, analyzed data, contributed new methods, wrote the paper; MRT: designed laboratory study, performed research, analyzed data; JWH: designed field study, performed research; KLM: performed research; LEP: performed research, contributed new method; GS: performed research; JCN: performed research.     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.     

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.     

Benthic algal response to N and P enrichment along a pH gradient

Nutrient enrichment and its effect on benthic algal growth, community composition, and average cell size was assessed across two sites of differing pH within a single habitat. Nutrients were added using in situ substrata, which released either N, P, or no additional nutrients (controls) at each site for 21 days. Upon collection, chlorophyll and biovolume standing stocks of the attached algal microflora were measured. Chlorophyll concentration was different among all treatments, accumulating greatest on P, followed by N, and the least on C substrata (P < 0.001) and was highest at site-2 (P < 0.001), while total algal biovolume was highest on P compared to both N and C substrata (P < 0.05) and did not vary between sites. Increased growth on P substrata was due to the enhanced biovolume of filamentous green algae, although the affected taxa varied between sites. Biovolume to cell density ratios (as a measure of average cell size) were highest on P substrata over both N-enriched and control substrata (P < 0.05) and this pattern was similar between sites. Progression towards a community composed of larger cells following P enrichment observed along this pH gradient, seems to be related to the dominance of larger celled filamentous green algae. Thus, nutrients exhibited greater control on benthic algal growth than did changes in hydrogen ion concentration.Contribution number 581, Great Lakes Environmental Research LaboratoryContribution number 581, Great Lakes Environmental Research Laboratory     

Light availability regulates the response of algae and heterotrophic bacteria to elevated nutrient levels and warming in a northern boreal peatland

相似文献   

Interactive controls of herbivory and fluvial dynamics on landscape vegetation patterns on the Tanana River floodplain, interior Alaska

Aim  We examined the interactive effects of mammalian herbivory and fluvial dynamics on vegetation dynamics and composition along the Tanana River in interior Alaska.
Location  Model parameters were obtained from field studies along the Tanana River, Alaska between Fairbanks (64°50.50' N, 147°43.30' W) and Manley Hot Springs (65°0.0' N, 150°36.0' W).
Methods  We used a spatially explicit model of landscape dynamics (ALFRESCO) to simulate vegetation changes on a 1-year time-step. The model was run for 250 years and was replicated 100 times.
Results  Increases in herbivory decreased the proportion of early successional vegetation and increased the proportion of late successional vegetation on the simulated landscape. Erosion and accretion worked as antagonists to herbivory, increasing the amount of early successional vegetation and decreasing the amount of late successional vegetation. However, the interactive effects of herbivory and erosion/accretion were especially important in determining system response, particularly in early seral vegetation types. High erosion rates, when coupled with low herbivory, greatly increased the proportion of willow on the landscape. When coupled with high herbivory, however, they greatly increased the proportion of alder on the landscape. At low levels of herbivory, alder abundance peaked at intermediate levels of erosion/accretion.
Main conclusions  Neither erosion/accretion nor herbivory produced consistent landscape patterns that could be predicted independently of the other. These findings underscore the importance of the interactive effects of biotic and abiotic disturbances in shaping large-scale landscape vegetation patterns in boreal floodplain ecosystems – systems traditionally thought to be driven primarily by abiotic disturbance alone.     

The response of planktonic and microbenthic algal assemblages to nutrient enrichment in shallow coastal waters,southwest Sweden

Field and laboratory nutrient (nitrogen and phosphorus) enrichment experiments were performed using natural phytoplankton and microphytobenthic assemblages from the brackish water Öresund, S.W. Sweden. The response of algae from a low-nutrient area (Falsterbo Canal) was compared to that of algae from a polluted, nutrient-rich area (Lomma Bay).The biomass (measured as chlorophyll a) of both phytoplankton and microphytobenthos from the Falsterbo Canal increased after the addition of nitrogen. Phytoplankton growth was stimulated by the addition of phosphorus to the nitrogen-rich water of the polluted Lomma Bay. Sediment chlorophyll a showed no significant increase after the addition of nutrients in the Lomma Bay. In containers without sediment, phytoplankton uptake was calculated to account for ≈ 90% of the disappearance of inorganic fixed nitrogen from the water. In the sediment containers the microphytobenthos was estimated to account for ≈20% of the nitrogen uptake. The rest was presumably lost mainly through denitrification.When containers with microphytobenthos from Lomma Bay were kept in the dark, phosphorus was released at a rate of up to ≈ 180 μM · m^?2 · day^?1. We suggest that by producing oxygen microbenthic algae keep the sediment surface oxygenated thereby decreasing phosphorus transport from the sediment to the overlying water.     

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.     

Plant allocation of carbon to defense as a function of herbivory,light and nutrient availability

We use modeling to determine the optimal relative plant carbon allocations between foliage, fine roots, anti-herbivore defense, and reproduction to maximize reproductive output. The model treats these plant components and the herbivore compartment as variables. Herbivory is assumed to be purely folivory. Key external factors include nutrient availability, degree of shading, and intensity of herbivory. Three alternative functional responses are used for herbivory, two of which are variations on donor-dependent herbivore (models 1a and 1b) and one of which is a Lotka–Volterra type of interaction (model 2). All three were modified to include the negative effect of chemical defenses on the herbivore. Analysis showed that, for all three models, two stable equilibria could occur, which differs from most common functional responses when no plant defense component is included. Optimal strategies of carbon allocation were defined as the maximum biomass of reproductive propagules produced per unit time, and found to vary with changes in external factors. Increased intensity of herbivory always led to an increase in the fractional allocation of carbon to defense. Decreases in available limiting nutrient generally led to increasing importance of defense. Decreases in available light had little effect on defense but led to increased allocation to foliage. Decreases in limiting nutrient and available light led to decreases in allocation to reproduction in models 1a and 1b but not model 2. Increases in allocation to plant defense were usually accompanied by shifts in carbon allocation away from fine roots, possibly because higher plant defense reduced the loss of nutrients to herbivory.     

Effect of nitrogen and light on nutrient concentrations and associated physiological responses in birch and fir seedlings

We grew seedlings of two co-occurring high elevation tree species in controlled light and nitrogen (N) environments to examine the effect on foliar N and P concentrations and the resulting correlation with photosynthesis and growth. Foliar N concentrations in both heart-leaf paper birch (Betula cordifolia) and balsam fir (Abies balsamea) seedlings were greater in low light treatments than in high light treatments. P concentrations, however, were lower in birch and fir foliage grown in low light than in high light. N-availability had no effect on foliar N in birch but tended to increase N concentration in fir needles at all but 100% ambient light. N-availability had no effect on P concentration in fir seedlings, but high N decreased foliar P in birch. There was a positive relationship between foliar N-concentration (mg g^–1) and mass-based maximum photosynthetic rate (Asat) in birch seedlings and a corresponding growth response to increased N-availability (suggesting N-limitation). Fir photosynthesis exhibited a positive correlation up to 22 mg g^–1 – N and a negative correlation above that point, suggesting that high N concentrations may be detrimental to photosynthesis in the fir seedlings. There was no significant effect of N-treatment on growth.     

The positive net radiative greenhouse gas forcing of increasing methane emissions from a thawing boreal forest‐wetland landscape

At the southern margin of permafrost in North America, climate change causes widespread permafrost thaw. In boreal lowlands, thawing forested permafrost peat plateaus (‘forest’) lead to expansion of permafrost‐free wetlands (‘wetland’). Expanding wetland area with saturated and warmer organic soils is expected to increase landscape methane (CH₄) emissions. Here, we quantify the thaw‐induced increase in CH₄ emissions for a boreal forest‐wetland landscape in the southern Taiga Plains, Canada, and evaluate its impact on net radiative forcing relative to potential long‐term net carbon dioxide (CO₂) exchange. Using nested wetland and landscape eddy covariance net CH₄ flux measurements in combination with flux footprint modeling, we find that landscape CH₄ emissions increase with increasing wetland‐to‐forest ratio. Landscape CH₄ emissions are most sensitive to this ratio during peak emission periods, when wetland soils are up to 10 °C warmer than forest soils. The cumulative growing season (May–October) wetland CH₄ emission of ~13 g CH₄ m^?2 is the dominating contribution to the landscape CH₄ emission of ~7 g CH₄ m^?2. In contrast, forest contributions to landscape CH₄ emissions appear to be negligible. The rapid wetland expansion of 0.26 ± 0.05% yr^?1 in this region causes an estimated growing season increase of 0.034 ± 0.007 g CH₄ m^?2 yr^?1 in landscape CH₄ emissions. A long‐term net CO₂ uptake of >200 g CO₂ m^?2 yr^?1 is required to offset the positive radiative forcing of increasing CH₄ emissions until the end of the 21st century as indicated by an atmospheric CH₄ and CO₂ concentration model. However, long‐term apparent carbon accumulation rates in similar boreal forest‐wetland landscapes and eddy covariance landscape net CO₂ flux measurements suggest a long‐term net CO₂ uptake between 49 and 157 g CO₂ m^?2 yr^?1. Thus, thaw‐induced CH₄ emission increases likely exert a positive net radiative greenhouse gas forcing through the 21st century.     

Reduction of transpiration and altered nutrient allocation contribute to nutrient decline of crops grown in elevated CO2 concentrations

Plants grown in elevated [CO₂] have lower protein and mineral concentrations compared with plants grown in ambient [CO₂]. Dilution by enhanced production of carbohydrates is a likely cause, but it cannot explain all of the reductions. Two proposed, but untested, hypotheses are that (1) reduced canopy transpiration reduces mass flow of nutrients to the roots thus reducing nutrient uptake and (2) changes in metabolite or enzyme concentrations caused by physiological changes alter requirements for minerals as protein cofactors or in other organic complexes, shifting allocation between tissues and possibly altering uptake. Here, we use the meta‐analysis of previous studies in crops to test these hypotheses. Nutrients acquired mostly by mass flow were decreased significantly more by elevated [CO₂] than nutrients acquired by diffusion to the roots through the soil, supporting the first hypothesis. Similarly, Mg showed large concentration declines in leaves and wheat stems, but smaller decreases in other tissues. Because chlorophyll requires a large fraction of total plant Mg, and chlorophyll concentration is reduced by growth in elevated [CO₂], this supports the second hypothesis. Understanding these mechanisms may guide efforts to improve nutrient content, and allow modeling of nutrient changes and health impacts under future climate change scenarios.     

Efficacy of algal metrics for assessing nutrient and organic enrichment in flowing waters

1. Algal-community metrics were calculated for periphyton samples collected from 976 streams and rivers by the U.S. Geological Survey's National Water-Quality Assessment (NAWQA) Programme during 1993–2001 to evaluate national and regional relations with water chemistry and to compare whether algal-metric values differ significantly among undeveloped and developed land-use classifications.
2. Algal metrics with significant positive correlations with nutrient concentrations included indicators of trophic condition, organic enrichment, salinity, motility and taxa richness. The relative abundance of nitrogen-fixing algae was negatively correlated with nitrogen concentrations, and the abundance of diatom species associated with high dissolved oxygen concentrations was negatively correlated with both nitrogen and phosphorus concentrations. Median algal-metric values and nutrient concentrations were significantly lower at undeveloped sites than those draining agricultural or urban catchments.
3. Total algal biovolume did not differ significantly among major river catchments or land-use classifications, and was only weakly correlated with nitrate (positive) and suspended-sediment (negative) concentrations. Estimates of periphyton chlorophyll a indicated an oligotrophic–mesotrophic boundary of about 21 mg m⁻² and a mesotrophic–eutrophic boundary of about 55 mg m⁻² based on upper and lower quartiles of the biovolume data distribution.
4. Although algal species tolerance to nutrient and organic enrichment is well documented, additional taxonomic and autecological research on sensitive, endemic algal species would further enhance water-quality assessments.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Analysis of vegetation distribution in Interior Alaska and sensitivity to climate change using a logistic regression approach

Aim To understand drivers of vegetation type distribution and sensitivity to climate change. Location Interior Alaska. Methods A logistic regression model was developed that predicts the potential equilibrium distribution of four major vegetation types: tundra, deciduous forest, black spruce forest and white spruce forest based on elevation, aspect, slope, drainage type, fire interval, average growing season temperature and total growing season precipitation. The model was run in three consecutive steps. The hierarchical logistic regression model was used to evaluate how scenarios of changes in temperature, precipitation and fire interval may influence the distribution of the four major vegetation types found in this region. Results At the first step, tundra was distinguished from forest, which was mostly driven by elevation, precipitation and south to north aspect. At the second step, forest was separated into deciduous and spruce forest, a distinction that was primarily driven by fire interval and elevation. At the third step, the identification of black vs. white spruce was driven mainly by fire interval and elevation. The model was verified for Interior Alaska, the region used to develop the model, where it predicted vegetation distribution among the steps with an accuracy of 60–83%. When the model was independently validated for north‐west Canada, it predicted vegetation distribution among the steps with an accuracy of 53–85%. Black spruce remains the dominant vegetation type under all scenarios, potentially expanding most under warming coupled with increasing fire interval. White spruce is clearly limited by moisture once average growing season temperatures exceeded a critical limit (+2 °C). Deciduous forests expand their range the most when any two of the following scenarios are combined: decreasing fire interval, warming and increasing precipitation. Tundra can be replaced by forest under warming but expands under precipitation increase. Main conclusion The model analyses agree with current knowledge of the responses of vegetation types to climate change and provide further insight into drivers of vegetation change.