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Global warming and excess nitrogen deposition can exert strong impacts on aquatic populations, communities, and ecosystems. However, experimental data to establish clear cause-and-effect relationships in naturally complex field conditions are scarce in aquatic environments. Here, we describe the design and performance of a unique outdoor enclosure facility used to simulate warming, increased nitrogen supply, and both factors combined in a littoral freshwater wetland dominated by common reed, Phragmites australis. The experimental system effectively simulated a 2.8 °C climate warming scenario over an extended period, capturing the natural temperature variations in the wetland at diel and seasonal scales with only small deviations. Excess nitrogen supply enhanced nitrate concentrations especially in winter when it was associated with increased concentration of ammonium and dissolved organic carbon. Nitrogen also reduced dissolved oxygen concentrations, particularly in the summer. Importantly, by stimulating biological activity, warming enhanced the nitrogen uptake capacity of the wetland during the winter, emphasizing the need for multifactorial global change experiments that examine both warming and nitrogen loading in concert. Establishing similar experiments across broad environmental gradients holds great potential to provide robust assessments of the impacts of climate change on shallow aquatic ecosystems.

Drought-induced events of massive tree mortality appear to be increasing worldwide. Species-specific vulnerability to drought mortality may alter patterns of species diversity and affect future forest composition. We have explored the consequences of the extreme drought of 2005, which caused high sapling mortality (approx. 50 %) among 10-year-old saplings of two coexisting pine species in the Mediterranean mountains of Sierra Nevada (Spain): boreo-alpine Pinus sylvestris and Mediterranean P. nigra. Sapling height growth, leaf δ(13)C and δ(18)O, and foliar nitrogen concentration in the four most recent leaf cohorts were measured in dead and surviving saplings. The foliar isotopic composition of dead saplings (which reflects time-integrated leaf gas-exchange until mortality) displayed sharp increases in both δ(13)C and δ(18)O during the extreme drought of 2005, suggesting an important role of stomatal conductance (g(s)) reduction and diffusional limitations to photosynthesis in mortality. While P. nigra showed decreased growth in 2005 compared to the previous wetter year, P. sylvestris maintained similar growth levels in both years. Decreased growth, coupled with a sharper increase in foliar δ(18)O during extreme drought in dead saplings, indicate a more conservative water use strategy for P. nigra. The different physiological behavior of the two pine species in response to drought (further supported by data from surviving saplings) may have influenced 2005 mortality rates, which contributed to 2.4-fold greater survival for P. nigra over the lifespan of the saplings. This species-specific vulnerability to extreme drought could lead to changes in dominance and distribution of pine species in Mediterranean mountain forests.

The importance of rock-derived mineral nutrients (P, K, Mn, Mg, and Ca) in plant physiological function is well established. However, one important and relatively unexplored question is whether or not the same rules of plant nutrient use efficiency apply to these essential elements even if they are not limiting to primary production. We examined conifer growth and nutrient use dynamics across sites with contrasting geologies (sedimentary and volcanic) that vary in both rock-derived mineral nutrient and N availability. Differences in bedrock geochemistry generally corresponded to differences in available soil nutrients, such that the volcanic site tended to have greater available nutrients. Foliar nutrient concentrations reflected both differences in bedrock chemistry and indices of available soil nutrients for P, K, and Mn. Aboveground biomass production did not follow expected patterns and was greater for trees growing on low nutrient sites, but only with respect to the annual woody increment. Fine litter production did not differ between sites. Finally, we found evidence for trade-offs between two commonly examined components of nutrient use efficiency (NUE): nutrient productivity (A n) and mean residence time of nutrients. However, we did not find evidence for higher plant NUE in soils with lower nutrient availability for N or rock-derived nutrients.

Estimating the nutritional value of a herbivore's diet is difficult because it requires knowing what the animal eats, the relative quality of each component and how these components interact in relation to animal physiology. Current methods are cumbersome and rely on many assumptions that are hard to evaluate. We describe a new method for estimating relative diet quality directly from faeces that avoids the problems inherent in other methods. We combine this method with near infrared reflectance spectroscopy (NIRS) to analyse many samples and thus provide a technique with immense value in ecological studies. The method stems from the correlation between the concentrations of dietary and faecal nitrogen in herbivores eating a tannin-free diet, but a weaker relationship in browsers that ingest substantial amounts of tannins, which form complexes with proteins. These complexes reduce the availability of nitrogen and may increase faecal nitrogen concentrations. Using the tannin-binding compound, polyethylene glycol, we showed that tannin-bound nitrogen is a significant and variable part of faecal nitrogen in wild common brushtail possums (Trichosurus vulpecula). We developed a technique to measure faecal available nitrogen and found that it predicted the reproductive success of female brushtail possums in northern Australia. Faecal available nitrogen combined with NIRS provides a powerful tool for estimating the relative nutritional value of the diets of browsing herbivores in many ecological systems. It is a better indicator of diet quality than other commonly used single-nutrient measures such as faecal nitrogen and foliage analysis paired with observed feeding behaviour.

Both atmospheric [CO2] and average surface temperatures are predicted to increase with potentially different, additive or opposing, effects on leaf quality and insect herbivore activity. Few studies have directly measured the interactive effects of concurrent changes in [CO2] and temperature on insect herbivores. None have done so over the entire developmental period of a tree-feeding insect, and none have compared responses to low pre-industrial [CO2] and present day [CO2] to estimate responses to future increases. Eucalypt herbivores may be particularly sensitive to climate-driven shifts in plant chemistry, as eucalypt foliage is naturally low in [N]. In this study, we assessed the development of the eucalypt herbivore Doratifera quadriguttata exposed concurrently to variable [CO2] (290, 400, 650 μmol mol(-1)) and temperature (ambient, ambient +4 °C) on glasshouse-grown Eucalyptus tereticornis. Overall, insects performed best on foliage grown at pre-industrial [CO2], indicating that modern insect herbivores have already experienced nutritional shifts since industrialisation. Rising [CO2] increased specific leaf mass and leaf carbohydrate concentration, subsequently reducing leaf [N]. Lower leaf [N] induced compensatory feeding and impeded insect performance, particularly by prolonging larval development. Importantly, elevated temperature dampened the negative effects of rising [CO2] on larval performance. Therefore, rising [CO2] over the past 200 years may have reduced forage quality for eucalypt insects, but concurrent temperature increases may have partially compensated for this, and may continue to do so in the future. These results highlight the importance of assessing plant-insect interactions within the context of multiple climate-change factors because of the interactive and potentially opposing effects of different factors within and between trophic levels.

We examined geographic variation in the structure and function of salt marsh communities along the Atlantic and Gulf coasts of the United States. Focusing on the arthropod community in the dominant salt marsh plant Spartina alterniflora, we tested two hypotheses: first, that marsh community structure varies geographically, and second, that two aspects of marsh function (response to eutrophication and addition of dead plant material) also vary geographically. We worked at eleven sites on the Gulf Coast and eleven sites on the Atlantic Coast, dividing each coast up into two geographic areas. Abiotic conditions (tidal range, soil organic content, and water content, but not soil salinity), plant variables (Spartina nitrogen content, height, cover of dead plant material, but not live Spartina percent cover or light interception), and arthropod variables (proportional abundances of predators, sucking herbivores, stem-boring herbivores, parasitoids, and detritivores, but not total arthropod numbers) varied among the four geographic regions. Latitude and mean tidal range explained much of this geographic variation. Nutrient enrichment increased all arthropod functional groups in the community, consistent with previous experimental results, and had similar effects in all geographic regions, contrary to our hypothesis, suggesting widespread consistency in this aspect of ecosystem function. The addition of dead plant material had surprisingly little effect on the arthropod community. Our results caution against the uncritical extrapolation of work done in one geographic region to another, but indicate that some aspects of marsh function may operate in similar ways in different geographic regions, despite spatial variation in community structure.

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China's West Development.

Partial mycoheterotrophy, a newly discovered form of mixotrophy in plants, has been described in at least two major lineages of angiosperms, the orchids and ericaceous plants in the tribe Pyroleae. Partial mycoheterotrophy entails carbon gains both directly from photosynthesis and via symbiotic mycorrhizal fungi, but determining the degree of plant dependence on fungal carbon is challenging. The purpose of this study was to determine if two chlorophyllous species of Pyroleae, Chimaphila umbellata and Pyrola picta, were receiving carbon via mycorrhizal networks and, if so, if their proportional dependency on fungal carbon gains increased under reduced light conditions. This was accomplished by a field experiment that manipulated light and plants' access to mycorrhizal networks, and by using the stable carbon isotope composition (δ(13)C) of leaf soluble sugars as a marker for the level of mycoheterotrophy. Based on leaf soluble sugars δ(13)C values, we calculated a site-independent isotope enrichment factor as a measure of fungal contributions to plant C. We found that, under each treatment and over time, the two test species demonstrated different isotopic responses caused by their different intrinsic physiologies. Our data, along with previously published studies, suggest that Chimaphila umbellata is primarily an autotrophic understory plant, while Pyrola picta may be capable of partial mycoheterotrophy. However, in this study, a 50% decrease in light availability did not significantly change the relative dependency of P. picta on carbon gains via mycoheterotrophy.

Ecologists want to explain why populations of animals are not evenly distributed across landscapes and often turn to nutritional explanations. In seeking to link population attributes with food quality, they often contrast nutritionally positive traits, such as the concentration of nitrogen, against negative ones, such as fibre concentration, by using a ratio of these traits. This specific ratio has attracted attention because it sometimes correlates with the biomass of colobine primates across sites in Asia and Africa. Although empirically successful, we have identified problems with the ratio that may explain why it fails under some conditions to predict colobine biomass. First, available nitrogen, rather than total nitrogen, is nutritionally important, while the presence of tannins is the major factor reducing the availability of nitrogen in browse plant species. Second, tannin complexes inflate measures of fibre. Finally, simple ratios may be unsound statistically because they implicitly assume isometric relationships between variables. We used data on the chemical composition of plants from three continents to examine the relationships between the concentrations of nitrogen, available nitrogen, fibre and tannins in foliage and to evaluate the nitrogen to fibre ratio. Our results suggest that the ratio of the concentration of nitrogen to fibre in leaves does sometimes closely correlate with the concentration of available nitrogen. However, the ratio may give misleading results when leaves contain high concentrations of tannins. The concentration of available nitrogen, which incorporates measures of total nitrogen, dry matter digestibility and tannins, should give a better indication of the nutritional value of leaves for herbivorous mammals that can readily be extrapolated to habitats.

This study evaluates acclimation of photosynthesis and stomatal conductance in three evolutionary lineages of C(3), C(3)-C(4) intermediate, and C(4) species grown in the low CO(2) and hot conditions proposed to favo r the evolution of C(4) photosynthesis. Closely related C(3), C(3)-C(4), and C(4) species in the genera Flaveria, Heliotropium, and Alternanthera were grown near 380 and 180 μmol CO(2) mol(-1) air and day/night temperatures of 37/29°C. Growth CO(2) had no effect on photosynthetic capacity or nitrogen allocation to Rubisco and electron transport in any of the species. There was also no effect of growth CO(2) on photosynthetic and stomatal responses to intercellular CO(2) concentration. These results demonstrate little ability to acclimate to low CO(2) growth conditions in closely related C(3) and C(3)-C(4) species, indicating that, during past episodes of low CO(2), individual C(3) plants had little ability to adjust their photosynthetic physiology to compensate for carbon starvation. This deficiency could have favored selection for more efficient modes of carbon assimilation, such as C(3)-C(4) intermediacy. The C(3)-C(4) species had approximately 50% greater rates of net CO(2) assimilation than the C(3) species when measured at the growth conditions of 180 μmol mol(-1) and 37°C, demonstrating the superiority of the C(3)-C(4) pathway in low atmospheric CO(2) and hot climates of recent geological time.

Seasonal oscillations in the carbon (δ(13)C) and nitrogen (δ(15)N) isotope signatures of aquatic algae can cause seasonal enrichment-depletion cycles in the isotopic composition of planktonic invertebrates (e.g., copepods). Yet, there is growing evidence that seasonal enrichment-depletion cycles also occur in the isotope signatures of larger invertebrate consumers, taxa used to define reference points in isotope-based trophic models (e.g., trophic baselines). To evaluate the general assumption of temporal stability in non-zooplankton aquatic invertebrates, δ(13)C and δ(15)N time series data from the literature were analyzed for seasonality and the influence of biotic (feeding group) and abiotic (trophic state, climate regime) factors on isotope temporal patterns. The amplitude of δ(13)C and δ(15)N enrichment-depletion cycles was negatively related to body size, although all size-classes of invertebrates displayed a winter-to-summer enrichment in δ(13)C and depletion in δ(15)N. Among feeding groups, periphytic grazers were more variable and displayed larger temporal changes in δ(13)C than detritivores. For nitrogen, temporal variability and magnitude of directional change of δ(15)N was most strongly related to ecosystem trophic state (eutrophic > mesotrophic, oligotrophic). This study provides evidence of seasonality in the isotopic composition of aquatic invertebrates across very broad geographical and ecological gradients as well as identifying factors that are likely to modulate the strength and variability of seasonality. These results emphasize the need for researchers to recognize the likelihood of temporal changes in non-zooplankton aquatic invertebrate consumers at time scales relevant to seasonal studies and, if present, to account for temporal dynamics in isotope trophic models.

Although the productivity and nitrogen (N)-use traits of mire plants differ dramatically between fens and bogs, soil N richness does not necessarily differ, whereas the soil-water pH is distinctly lower in bogs than in fens. The ecophysiological mechanisms underlying these relations are unclear. To assess the relative availability of N forms in relation to soil-water pH, we focused on the net N uptake rate per unit root weight (NNUR), glutamine synthetase activity and nitrate reductase activity, and performed reciprocal transplant experiments with the seedlings of fen (Carex lyngbyei) and bog (C. middendorffii) sedge species in intact habitat sites. The soil-water pH was clearly lower at the bog site, but the NH(4) (+), NO(3) (-) or dissolved organic-N concentrations did not differ between the fen and bog sites. The activity of both enzymes for inorganic-N assimilation did not differ among the sites and species. However, the fen species grown at bog sites showed a drastic decrease in the NNUR, suggesting a suppression of organic-N uptake. The bog species showed no NNUR difference between the sites. These results indicate that inorganic-N availability does not differ between the two habitats, but organic-N availability is lowered in a low-pH bog, particularly in the case of fen species. Therefore, the relative availability of N forms shows species-specific variations that depend on the differences in the soil-water pH of root zone, even at similar N richness, which would play a key role in plant distribution strategies in relation to the fen-bog gradient.

Global climate change is expected to affect terrestrial ecosystems in a variety of ways. Some of the more well-studied effects include the biogeochemical feedbacks to the climate system that can either increase or decrease the atmospheric load of greenhouse gases such as carbon dioxide and nitrous oxide. Less well-studied are the effects of climate change on the linkages between soil and plant processes. Here, we report the effects of soil warming on these linkages observed in a large field manipulation of a deciduous forest in southern New England, USA, where soil was continuously warmed 5°C above ambient for 7 years. Over this period, we have observed significant changes to the nitrogen cycle that have the potential to affect tree species composition in the long term. Since the start of the experiment, we have documented a 45% average annual increase in net nitrogen mineralization and a three-fold increase in nitrification such that in years 5 through 7, 25% of the nitrogen mineralized is then nitrified. The warming-induced increase of available nitrogen resulted in increases in the foliar nitrogen content and the relative growth rate of trees in the warmed area. Acer rubrum (red maple) trees have responded the most after 7 years of warming, with the greatest increases in both foliar nitrogen content and relative growth rates. Our study suggests that considering species-specific responses to increases in nitrogen availability and changes in nitrogen form is important in predicting future forest composition and feedbacks to the climate system.

We investigated how photosynthesis by understory seedlings of the lowland tropical tree species Alseis blackiana responded to 10 years of soil nutrient fertilization with N, P and K. We ask whether nutrients are limiting to light and CO(2) acquisition in a low light understory environment. We measured foliar nutrient concentrations of N, P and K, isotopic composition of carbon (δ(13)C) and nitrogen (δ(15)N), and light response curves of photosynthesis and chlorophyll fluorescence. Canopy openness was measured above each study seedling and included in statistical analyses to account for variation in light availability. Foliar N concentration increased by 20% with N addition. Foliar P concentration increased by 78% with P addition and decreased by 14% with N addition. Foliar K increased by 8% with K addition. Foliar δ(13)C showed no significant responses, and foliar δ(15)N decreased strongly with N addition, matching the low δ(15)N values of applied fertilizer. Canopy openness ranged from 0.01 to 6.71% with a mean of 1.76 ± 0.14 (± 1SE). Maximum photosynthetic CO(2) assimilation rate increased by 9% with N addition. Stomatal conductance increased with P addition and with P and K in combination. Chlorophyll fluorescence measurements revealed that quantum yield of photosystem II increased with K addition, maximum electron transport rate trended 9% greater with N addition (p = 0.07), and saturating photosynthetically active radiation increased with N addition. The results demonstrate that nutrient addition can enhance photosynthetic processes, even under low light availability.

Changing environments can have divergent effects on biodiversity-ecosystem function relationships at alternating trophic levels. Freshwater mussels fertilize stream foodwebs through nutrient excretion, and mussel species-specific excretion rates depend on environmental conditions. We asked how differences in mussel diversity in varying environments influence the dynamics between primary producers and consumers. We conducted field experiments manipulating mussel richness under summer (low flow, high temperature) and fall (moderate flow and temperature) conditions, measured nutrient limitation, algal biomass and grazing chironomid abundance, and analyzed the data with non-transgressive overyielding and tripartite biodiversity partitioning analyses. Algal biomass and chironomid abundance were best explained by trait-independent complementarity among mussel species, but the relationship between biodiversity effects across trophic levels (algae and grazers) depended on seasonal differences in mussel species' trait expression (nutrient excretion and activity level). Both species identity and overall diversity effects were related to the magnitude of nutrient limitation. Our results demonstrate that biodiversity of a resource-provisioning (nutrients and habitat) group of species influences foodweb dynamics and that understanding species traits and environmental context are important for interpreting biodiversity experiments.

In semi-arid ecosystems, heterogeneous resources can lead to variable seedling recruitment. Existing vegetation can influence seedling establishment by modifying the resource and physical environment. We asked how a native legume, Lupinus argenteus, modifies microenvironments in unburned and burned sagebrush steppe, and if L. argenteus presence facilitates seedling establishment of native species and the non-native annual grass, Bromus tectorum. Field treatments examined mechanisms by which L. argenteus likely influences establishment: (1) live L. argenteus; (2) dead L. argenteus; (3) no L. argenteus; (4) no L. argenteus with L. argenteus litter; (5) no L. argenteus with inert litter; and (6) mock L. argenteus. Response variables included soil nitrogen, moisture, temperature, solar radiation, and seedling establishment of the natives Elymus multisetus and Eriogonum umbellatum, and non-native B. tectorum. In both unburned and burned communities, there was higher spring soil moisture, increased shade and reduced maximum temperatures under L. argenteus canopies. Adult L. argenteus resulted in greater amounts of soil nitrogen (N) only in burned sagebrush steppe, but L. argenteus litter increased soil N under both unburned and burned conditions. Although L. argenteus negatively affected emergence and survival of B. tectorum overall, its presence increased B. tectorum biomass and reproduction in unburned plots. However, L. argenteus had positive facilitative effects on size and survival of E. multisetus in both unburned and burned plots. Our study indicates that L. argenteus can facilitate seedling establishment in semi-arid systems, but net effects depend on the species examined, traits measured, and level of abiotic stress.

We measured structural and chemical traits of the leaves of native, broad-leaved trees in two temperate localities [southern Ontario, Canada (34 species), and Missouri (36 species)] and one tropical locality [central Panama (samples of 21 and 23 species)] to test the hypothesis that the greater diversity of tree species and herbivore species in the tropics is associated with greater resource niche space for herbivores. Variables were leaf toughness, water content, dry mass per unit area, several structural and nutritional carbohydrates, common mineral elements, including nitrogen and phosphorus, and several defensive compounds, including tannins and alkaloids. The four samples were almost fully separable by discriminant analysis on the basis of these leaf traits. Variance in log-transformed trait values among species was lowest in the most northern sample, but did not differ significantly between Missouri and Panama. Niche space, estimated as the square root of the total variance in the log-transformed variables within each locality, varied approximately as Panama = 1, Missouri = 0.8, Ontario = 0.5. Although niche space decreases towards higher latitudes, the change does not match the ca. sixfold decrease in tree species richness or the ca. fourfold decrease in Lepidopteran species richness over the latitude range of our samples. Accordingly, tropical folivore diversity is associated with greater resource niche overlap, greater niche specialization, and/or more completely filled niches, or with variation in niche dimensions not measured in this study.

Anthropogenic nitrogen (N) deposition affects many natural processes, including forest litter decomposition. Saprotrophic fungi are the only organisms capable of completely decomposing lignocellulosic (woody) litter in temperate ecosystems, and therefore the responses of fungi to N deposition are critical in understanding the effects of global change on the forest carbon cycle. Plant litter decomposition under elevated N has been intensively studied, with varying results. The complexity of forest floor biota and variability in litter quality have obscured N-elevation effects on decomposers. Field experiments often utilize standardized substrates and N-levels, but few studies have controlled the decay organisms. Decomposition of beech (Fagus sylvatica) blocks inoculated with two cord-forming basidiomycete fungi, Hypholoma fasciculare and Phanerochaete velutina, was compared experimentally under realistic levels of simulated N deposition at Wytham Wood, Oxfordshire, UK. Mass loss was greater with P. velutina than with H. fasciculare, and with N treatment than in the control. Decomposition was accompanied by growth of the fungal mycelium and increasing N concentration in the remaining wood. We attribute the N effect on wood decay to the response of cord-forming wood decay fungi to N availability. Previous studies demonstrated the capacity of these fungi to scavenge and import N to decaying wood via a translocating network of mycelium. This study shows that small increases in N availability can increase wood decomposition by these organisms. Dead wood is an important carbon store and habitat. The responses of wood decomposers to anthropogenic N deposition should be considered in models of forest carbon dynamics.

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.

The relationship between inorganic nitrogen (N) cycling and plant productivity is well established. However, recent research has demonstrated the ability of plants to take up low molecular weight organic N compounds (i.e., amino acids) at rates that often rival those of inorganic N forms. In this study, we hypothesize that temperate forest tree species characteristic of low-fertility habitats will prefer amino acids over species characteristic of high-fertility habitats. We measured the uptake of (15)N-labeled amino acids (glycine, glutamine, arginine, serine), ammonium (NH(4)(+)), and nitrate (NO(3)(-)) by four tree species that commonly occur in eastern North America, where their abundances have been correlated with inorganic N availability. Specific uptake rates of amino acids were largely similar for all tree species; however, high-fertility species took up NH(4)(+) at rates more than double those of low-fertility species, rendering amino acid N relatively more important to the N nutrition of low-fertility species. Low-fertility species acquired over four times more total N from arginine compared to NH(4)(+) and NO(3)(-); high-fertility species acquired the most N from NH(4)(+). Arginine had the highest uptake rates of any amino acid by all species; there were no significant differences in uptake rates of the remaining amino acids. Our results support the idea that the dominant species in a particular habitat are those best able to utilize the most available N resources.

Plants in nutrient-poor environments typically have low foliar nitrogen (N) concentrations, long-lived tissues with leaf traits designed to use nutrients efficiently, and low rates of photosynthesis. We postulated that increasing N availability due to atmospheric deposition would increase photosynthetic capacity, foliar N, and specific leaf area (SLA) of bog shrubs. We measured photosynthesis, foliar chemistry and leaf morphology in three ericaceous shrubs (Vaccinium myrtilloides, Ledum groenlandicum and Chamaedaphne calyculata) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada, with a background deposition of 0.8 g N m(-2) a(-1). While biomass and chlorophyll concentrations increased in the highest nutrient treatment for C. calyculata, we found no change in the rates of light-saturated photosynthesis (A(max)), carboxylation (V(cmax)), or SLA with nutrient (N with and without PK) addition, with the exception of a weak positive correlation between foliar N and A(max) for C. calyculata, and higher V(cmax) in L. groenlandicum with low nutrient addition. We found negative correlations between photosynthetic N use efficiency (PNUE) and foliar N, accompanied by a species-specific increase in one or more amino acids, which may be a sign of excess N availability and/or a mechanism to reduce ammonium (NH(4)) toxicity. We also observed a decrease in foliar soluble Ca and Mg concentrations, essential minerals for plant growth, but no change in polyamines, indicators of physiological stress under conditions of high N accumulation. These results suggest that plants adapted to low-nutrient environments do not shift their resource allocation to photosynthetic processes, even after reaching N sufficiency, but instead store the excess N in organic compounds for future use. In the long term, bog species may not be able to take advantage of elevated nutrients, resulting in them being replaced by species that are better adapted to a higher nutrient environment.

Ecosystem processes of northern peatlands are largely governed by the vitality and species composition in the bryophyte layer, and may be affected by global warming and eutrophication. In a factorial experiment in northeast China, we tested the effects of raised levels of nitrogen (0, 1 and 2 g m(-2) year(-1)), phosphorus (0, 0.1 and 0.2 g m(-2) year(-1)) and temperature (ambient and +3°C) on Polytrichum strictum, Sphagnum magellanicum and S. palustre, to see if the effects could be altered by inter-specific interactions. In all species, growth declined with nitrogen addition and increased with phosphorus addition, but only P. strictum responded to raised temperature with increased production of side-shoots (branching). In Sphagnum, growth and branching changed in the same direction, but in Polytrichum, the two responses were uncoupled: with nitrogen addition there was a decrease in growth (smaller than in Sphagnum) but an increase in branching; with phosphorus addition growth increased but branching was unaffected. There were no two-way interactions among the P, N and T treatments. With increasing temperature, our results indicate that S. palustre should decrease relative to P. strictum (Polytrichum increased its branching and had a negative neighbor effect on S. palustre). With a slight increase in phosphorus availability, the increase in length growth and production of side-shoots in P. strictum and S. magellanicum may give them a competitive superiority over S. palustre. The negative response in Sphagnum to nitrogen could favor the expansion of vascular plants, but P. strictum may endure thanks to its increased branching.

Competition between large and small species for the same food is common in a number of ecosystems including aquatic ones. How diversity of larger consumers affects the access of smaller competitors to a limiting resource is not well understood. We tested experimentally how species richness (0-3 spp.) of benthic deposit-feeding macrofauna changes meiofaunal ostracods' incorporation of fresh organic matter from a stable-isotope-labeled cyanobacterial bloom, using fauna from the species-poor Baltic Sea. Presence of macrofauna mostly decreased meiofaunal incorporation of bloom material, depending on the macrofauna species present. As expected, the species identity of macrofauna influenced the incorporation of organic matter by meiofauna. Interestingly, our results show that, in addition, species richness of the macrofauna significantly reduced meiofauna incorporation of freshly settled nitrogen and carbon. With more than one macrofauna species, the reduction was always greater than expected from the single-species treatments. Field data from the Baltic Sea showed a negative correlation between macrofauna diversity and meiofaunal ostracod abundance, as expected from the experimental results. We argue that this is caused by interference competition, due to spatial niche differentiation between macrofauna species reducing the sediment volume in which ostracods can feed undisturbed by larger competitors. Interference from macrofauna significantly reduces organic matter incorporation by meiofauna, indicating that diversity of larger consumers is an important factor controlling the access of smaller competitors to a limiting food resource.

Understanding how food web interactions alter the processing of limiting nutrient elements is an important goal of ecosystem ecology. An experiment manipulating densities of the grasshopper Oedaleus asiaticus was performed to assess top-down effects of grasshoppers on C:N:P stoichiometry of plants and soil in a grassland ecosystem in Inner Mongolia (China). With increased grasshopper feeding, plant biomass declined fourfold, litter abundance increased 30%, and the plant community became dominated by non-host plant taxa. Plant stoichiometric response depended on whether or not the plant was a grasshopper host food species: C:N and C:P ratios increased with increasing grasshopper density (GD) for host plants but decreased in non-host plants. These data suggest either a direct transfer of grasshopper-recycled nutrients from host to non-host plants or a release of non-host plants from nutrient competition with heavily grazed host plants. Litterfall C:N and C:P decreased across moderate levels of grasshopper density but no effects on C:N:P stoichiometry in the surface soil were observed, possibly due to the short experimental period. Our observations of divergent C:N:P stoichiometric response among plant species highlight the important role of grasshopper herbivory in regulating plant community structure and nutrient cycling in grassland ecosystems.

The niche dimension hypothesis predicts that greater numbers of limiting factors can allow greater numbers of species to coexist through species' tradeoffs for different limiting factors. A prediction that follows is that addition of multiple limiting resources to plant communities will increase productivity and simultaneously decrease diversity. Species loss due to limiting resource enrichment might occur through reducing the number of resources that species compete for or by changing the identity of limiting factors. We tested these predictions of the niche dimension hypothesis in an arid annual grassland by adding combinations of nutrients: nitrogen (N), phosphorus (P), and potassium with other elements (O). We found that species number decreased while biomass increased with greater numbers of added resources. In particular, N in combinations with P or O resulted in the greatest species loss, while biomass increased super-additively with N and P together. The addition of greater numbers of added nutrients decreased the availability of light and soil moisture, consistent with a potential shift in the identity of limiting resources. Species also differed in their responses to different combinations of N, P, and O, supporting predictions of resource-ratio tradeoffs. These results are particularly notable because this experiment was conducted during a drought year in an arid grassland (226 mm annual rainfall), which might have been expected to be water-rather than nutrient-limited. Our results support the hypothesis that plant diversity may be maintained by high-dimensional tradeoffs among species in their abilities to compete for multiple limiting factors.

Carbon stable isotopes (δ(13)C) of particulate organic matter (POM) have been used as indicators for energy flow, primary productivity and carbon dioxide concentration in individual lakes. Here, we provide a synthesis of literature data from 32 freshwater lakes around the world to assess the variability of δ(13)C(POM) along latitudinal, morphometric and biogeochemical gradients. Seasonal mean δ(13)C(POM), a temporally integrated measure of the δ(13)C(POM), displayed weak relationships with all trophic state indices [total phosphorus (TP), total nitrogen (TN), and chlorophyll a (Chl a)], but decreased significantly with the increase in latitude, presumably in response to the corresponding decrease in water temperature and increase in CO(2) concentration. The seasonal minimum δ(13)C(POM) also correlated negatively with latitude while seasonal maximum δ(13)C(POM) correlated positively with all trophic state indices, pH, and δ(13)C of dissolved inorganic carbon (DIC). Seasonal amplitude of δ(13)C(POM) (the difference between seasonal maximum and minimum values) correlated significantly with pH, TP and Chl a concentrations and displayed small variations in oligotrophic, mesotrophic and low latitude eutrophic lakes, which is attributed to low primary productivity and abundant non-living POM in the low trophic state lakes and relatively stable environmental conditions in the subtropics. Seasonal amplitude of δ(13)C(POM) was the greatest in high latitude eutrophic lakes. Greater seasonal changes in solar energy and light regime may be responsible for the large seasonal variability in high latitude productive lakes. This synthesis provides new insights on the factors controlling variations in stable carbon isotopes of POM among lakes on the global scale.

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

Invasive alien plants increase both plant N and soil inorganic N pools in many terrestrial ecosystems. This is believed to be the result of altered plant-soil-microbe feedbacks that accelerate N cycling. However, it may also be due to the greater ability of invasive species to uptake lateral N subsidies that can modify ecosystem N dynamics. We conducted manipulative field experiments to determine the impact of smooth cordgrass (Spartina alterniflora) invasion on the N cycling of salt marsh ecosystems in the Yangtze Estuary, China. The results showed that the aboveground plant N and soil inorganic N pools in S. alterniflora marshes, 14.39 and 3.16 g N m(-2), were significantly higher than those in native common reed (Phragmites australis) marshes, 11.61 and 2.29 g N m(-2). These increases after invasion were explained by a significantly higher uptake of dissolved inorganic N (DIN) from tidal subsidies in S. alterniflora marshes (6.59 g N m(-2)) than from those in P. australis marshes (1.61 g N m(-2)), and not by soil organic N mineralization, which was not significantly different between S. alterniflora (6.45 g N m(-2)) and P. australis marshes (6.84 g N m(-2)) during the growing season. Our study indicated that the ecosystem engineering effects of S. alterniflora, which increases the interception of external N input, can be an alternative mechanism that increases plant N and soil inorganic N pools--especially in ecosystems with ample anthropogenic N subsidies, such as the coastal wetlands of China.

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castilleja occidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes.

Survival, growth, aboveground biomass accumulation, sediment surface elevation dynamics and nitrogen accumulation in sediments were studied in experimental treatments planted with four different densities (6.96, 3.26, 1.93 and 0.95 seedlings m(-2)) of the mangrove Rhizophora mucronata in Puttalam Lagoon, Sri Lanka. Measurements were taken over a period of 1,171 days and were compared with those from unplanted controls. Trees at the lowest density showed significantly reduced survival, whilst measures of individual tree growth did not differ among treatments. Rates of surface sediment accretion (means ± SE) were 13.0 (±1.3), 10.5 (±0.9), 8.4 (±0.3), 6.9 (±0.5) and 5.7 (±0.3) mm year(-1) at planting densities of 6.96, 3.26, 1.93, 0.95, and 0 (unplanted control) seedlings m(-2), respectively, showing highly significant differences among treatments. Mean (±SE) rates of surface elevation change were much lower than rates of accretion at 2.8 (±0.2), 1.6 (±0.1), 1.1 (±0.2), 0.6 (±0.2) and -0.3 (±0.1) mm year(-1) for 6.96, 3.26, 1.93, 0.95, and 0 seedlings m(-2), respectively. All planted treatments accumulated greater nitrogen concentrations in the sediment compared to the unplanted control. Sediment %N was significantly different among densities which suggests one potential causal mechanism for the facilitatory effects observed: high densities of plants potentially contribute to the accretion of greater amounts of nutrient rich sediment. While this potential process needs further research, this study demonstrated how higher densities of mangroves enhance rates of sediment accretion and surface elevation processes that may be crucial in mangrove ecosystem adaptation to sea-level rise. There was no evidence that increasing plant density evoked a trade-off with growth and survival of the planted trees. Rather, facilitatory effects enhanced survival at high densities, suggesting that managers may be able to take advantage of high plantation densities to help mitigate sea-level rise effects by encouraging positive sediment surface elevation.

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m(-2) greater than unburned plots for several weeks and increased under shrubs (0.5-1.0 g m(-2)) more than interspaces (0.1-0.2 g m(-2)). Soil NO(3) (-)-N (nitrate-N) increased more and persisted longer than soil NH(4) (+)-N (ammonium-N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.

While climate warming can increase plant N availability over the growing season by increasing rates of N mineralization, increased N mineralization over winter at a time when plant roots are largely inactive, coupled with an increased frequency of soil freeze-thaw cycles, may increase soil N leaching losses. We examined changes in soil net N mineralization and N leaching in response to warming and N addition (6 g m(-2) year(-1)) in a factorial experiment conducted in a temperate old field. We used two warming treatments, year-round and winter-only warming, to isolate the effects of winter warming on soil N dynamics from the year-round warming effects. We estimated net N mineralization using in situ soil cores with resin bags placed at the bottom to catch throughput, and we measured N leaching using lysimeters located below the plant rooting zone at a depth of 50 cm. There were minor effects of warming on changes in soil extractable N and resin N in the soil cores over winter. Nevertheless, the overall effects of both warming and N addition on net N mineralization (the sum of changes in soil extractable N and resin N) were not significant over this period. Likewise, there were no significant treatment effects on the concentration of N in leachate collected below the plant rooting zone. However, in response to winter warming, net N mineralization over summer was approximately double that of both the ambient and year-round warming treatments. This result demonstrates a potentially large and unexpected effect of winter warming on soil N availability in this old field system.

Sphagnum, the main genus which forms boreal peat, is strongly affected by N and S deposition and raised temperature, but the physiological mechanisms behind the responses are largely unknown. We measured maximum photosynthetic rate (NP(max)), maximum efficiency of photosystem II [variable fluorescence (F (v))/maximum fluorescence yield (F (m))] and concentrations of N, C, chlorophyll and carotenoids as responses to N and S addition and increased temperature in Sphagnum balticum (a widespread species in the northern peatlands) in a 12-year factorial experiment. NP(max) did not differ between control (0.2 g N m(-2) year(-1)) and high N (3.0 g N m(-2) year(-1)), but was higher in the mid N treatment (1.5 g N m(-2) year(-1)). N, C, carotenoids and chlorophyll concentration increased in shoot apices after N addition. F (v)/F (m) did not differ between N treatments. Increased temperature (+3.6 degrees C) had a small negative effect on N concentration, but had no significant effect on NP(max) or F (v)/F (m). Addition of 2 g S m(-2) year(-1) showed a weak negative effect on NP(max) and F (v)/F (m). Our results suggest a unimodal response of NP(max) to N addition and tissue N concentration in S. balticum, with an optimum N concentration for photosynthetic rate of ~13 mg N g(-1). In conclusion, high S deposition may reduce photosynthetic capacity in Sphagnum, but the negative effects may be relaxed under high N availability. We suggest that previously reported negative effects on Sphagnum productivity under high N deposition are not related to negative effects on the photosynthetic apparatus, but differences in optimum N concentration among Sphagnum species may affect their competitive ability under different N deposition regimes.

Prolific fine root growth coupled with small accumulations of dead fine roots indicate rapid rates of fine root production, mortality and decay in young tree plantations in lowland Costa Rica. However, published studies indicate that fine roots decay relatively slowly in tropical forests. To resolve this discrepancy, we used the intact-core technique to quantify first-year decay rates of fine roots in four single-species plantations of native tree species. We tested three hypotheses: first, that fine roots from different tree species would decay at different rates; second, that species having rapid fine root growth rates would also have rapid rates of fine root decay; and third, that differences in fine root decay among species could be explained by fine root chemistry variables previously identified as influencing decay rates. Fine roots in Virola koschnyi plantations decayed very slowly (k = 0.29 +/- 0.15 year(-1)); those of Vochysia guatemalensis decayed seven times faster (k = 2.00 +/- 0.13 year(-1)). Decay rates of the remaining two species, Hieronyma alchorneoides and Pentaclethra macroloba, were 1.36 and 1.28 year(-1), respectively. We found a positive, marginally significant correlation between fine root decay rates and the relative growth rates of live fine roots (R = 0.93, n = 4, P = 0.072). There was a highly significant negative correlation between fine root decay and fine root lignin:N (R = 0.99, P = 0.01), which supports the use of lignin:N as a decay-controlling factor within terrestrial ecosystem models. The decay rates that we observed in this single study location encompassed the entire range of fine root decay rates previously observed in moist tropical forests, and thus suggest great potential for individual tree species to alter belowground organic matter and nutrient dynamics within a biotically rich rainforest environment.

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in seasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO(2) assimilation per unit mass (A(mass)), nitrogen concentration (N(mass)), and delta(13)C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO(2) assimilation per unit area (A(area)), phosphorus concentration per unit mass (P(mass)), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree A(area) decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana delta(13)C increased four times more than tree delta(13)C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher A(mass) than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.

Plant communities, soil organic matter and microbial communities are predicted to be interlinked and to exhibit concordant patterns along major environmental gradients. We investigated the relationships between plant functional type composition, soil organic matter quality and decomposer community composition, and how these are related to major environmental variation in non-acid and acid soils derived from calcareous versus siliceous bedrocks, respectively. We analysed vegetation, organic matter and microbial community compositions from five non-acidic and five acidic heath sites in alpine tundra in northern Europe. Sequential organic matter fractionation was used to characterize organic matter quality and phospholipid fatty acid analysis to detect major variation in decomposer communities. Non-acidic and acidic heaths differed substantially in vegetation composition, and these disparities were associated with congruent shifts in soil organic matter and microbial communities. A high proportion of forbs in the vegetation was positively associated with low C:N and high soluble N:phenolics ratios in soil organic matter, and a high proportion of bacteria in the microbial community. On the contrary, dwarf shrub-rich vegetation was associated with high C:N and low soluble N:phenolics ratios, and a high proportion of fungi in the microbial community. Our study demonstrates a strong link between the plant community composition, soil organic matter quality, and microbial community composition, and that differences in one compartment are paralleled by changes in others. Variation in the forb-shrub gradient of vegetation may largely dictate variations in the chemical quality of organic matter and decomposer communities in tundra ecosystems. Soil pH, through its direct and indirect effects on plant and microbial communities, seems to function as an ultimate environmental driver that gives rise to and amplifies the interactions between above- and belowground systems.

Herbivores shape plant communities through selective foraging. However, both herbivore selectivity and the plant's ability to tolerate or resist herbivory may depend on the density of herbivores. In an alpine ecosystem with a long history of grazing, plants are expected to respond to both enhanced and reduced grazing pressures, and the interaction between plant traits and changes in species abundance are expected to differ between the two types of alteration of grazing regime. To understand the mechanisms behind species response, we investigated the relationship between sheep selectivity (measured in situ), plant traits and experimentally derived measures of change in species abundance as a response to the enhancement (from low to high density) or cessation (from low to zero density) of sheep grazing pressure over a six-year time period for 22 abundant herb species in an alpine habitat in south Norway. Sheep selected large, late-flowering herbs with a low leaf C/N ratio. Species that increased in abundance in response to enhanced grazing pressure were generally small and had high root/shoot ratios, thus exhibiting traits that reflect both resistance (through avoidance) and tolerance (through regrowth capacity) strategies. The abundance of selected species remained stable during the study period, and also under the enhanced grazing pressure treatment. There was, however, a tendency for selected species to respond positively to cessation of grazing, although overall responses to cessation of grazing were much less pronounced than responses to enhanced grazing. Avoidance through short stature (probably associated with increased light availability through the removal of tall competitors) as well as a certain amount of regrowth capacity appear to be the main mechanisms behind a positive response to enhanced grazing pressure in this study. The plant trait perspective clearly improves our insight into the mechanisms behind observed changes in species abundance when the disturbance regime is altered.

Atmospheric carbon dioxide (CO(2)) and ozone (O(3)) concentrations are rising, which may have opposing effects on tree C balance and allocation to fine roots. More information is needed on interactive CO(2) and O(3) effects on roots, particularly fine-root life span, a critical demographic parameter and determinant of soil C and N pools and cycling rates. We conducted a study in which ponderosa pine (Pinus ponderosa) seedlings were exposed to two levels of CO(2) and O(3) in sun-lit controlled-environment mesocosms for 3 years. Minirhizotrons were used to monitor individual fine roots in three soil horizons every 28 days. Proportional hazards regression was used to analyze effects of CO(2), O(3), diameter, depth, and season of root initiation on fine-root survivorship. More fine roots were produced in the elevated CO(2) treatment than in ambient CO(2). Elevated CO(2), increasing root diameter, and increasing root depth all significantly increased fine-root survivorship and median life span. Life span was slightly, but not significantly, lower in elevated O(3), and increased O(3) did not reduce the effect of elevated CO(2). Median life spans varied from 140 to 448 days depending on the season of root initiation. These results indicate the potential for elevated CO(2) to increase the number of fine roots and their residence time in the soil, which is also affected by root diameter, root depth, and phenology.

The majority of plants are involved in symbioses with arbuscular mycorrhizal fungi (AMF), and these associations are known to have a strong influence on the performance of both plants and insect herbivores. Little is known about the impact of AMF on complex trophic chains, although such effects are conceivable. In a greenhouse study we examined the effects of two AMF species, Glomus intraradices and G. mosseae on trophic interactions between the grass Phleum pratense, the aphid Rhopalosiphum padi, and the parasitic wasp Aphidius rhopalosiphi. Inoculation with AMF in our study system generally enhanced plant biomass (+5.2%) and decreased aphid population growth (-47%), but there were no fungal species-specific effects. When plants were infested with G. intraradices, the rate of parasitism in aphids increased by 140% relative to the G. mosseae and control treatment. When plants were associated with AMF, the developmental time of the parasitoids decreased by 4.3% and weight at eclosion increased by 23.8%. There were no clear effects of AMF on the concentration of nitrogen and phosphorus in plant foliage. Our study demonstrates that the effects of AMF go beyond a simple amelioration of the plants' nutritional status and involve rather more complex species-specific cascading effects of AMF in the food chain that have a strong impact not only on the performance of plants but also on higher trophic levels, such as herbivores and parasitoids.

Assimilation-based techniques such as stoichiometric analysis, bulk tissue stable isotope analysis (SIA), fatty acid (FA) profiling and compound-specific SIA of FAs can be used to resolve ambiguities in consumer-resource relationships, but comparisons of their effectiveness are lacking. Feeding trials and concurrent application of these techniques were used to investigate the trophic base of two shrimps from Hong Kong streams: the atyid Caridina cantonensis, which is a primary consumer, and predatory Macrobrachium hainanense (Palaemonidae). Leaf litter and periphyton were fed to C. cantonensis (CC-LF and CC-PF, respectively) reared in the laboratory for 2 months, when C. cantonensis, that had fed on a mixed diet (CC-WC), were collected from the field. Atyids from each group (CC-LF, CC-PF and CC-WC) were fed to M. hainanense (MH-L, MH-P and MH-W, respectively) during a further 2-month trial, at the end of which M. hainanense were also collected from the field (MH-R). FA biomarkers present in CC-WC and CC-PF indicated that C. cantonensis depended primarily on autochthonous foods, and FA profiles of CC-WC were distinct from those of CC-LF. Differences in C/N ratios and isotope signatures of leaf litter and periphyton were not reflected in tissues of atyids or palaemonid predators. FA profiles of M. hainanense groups were similar, but FA stable C isotope ratio (delta(13)C) signatures of MH-R and MH-W were distinct from those of MH-L and leaf litter. FA delta(13)C signatures of MH-R were similar to those of MH-P and MH-W, indicating that autochthonous resources constituted the trophic base of production for this predator. This is the first study using compound-specific SIA to study stream food chains. Compound-specific SIA and FA profiling allowed accurate elucidation of consumer-resource relationships that were not revealed by stoichiometry or bulk tissue SIA. Compound-specific SIA was particularly sensitive for detecting rapid shifts in the predator trophic base. This tool will have wide applicability for investigating food webs in a range of ecosystems.

Regional analyses and biogeochemical models predict that ecosystem N pools and N cycling rates must increase from the semi-arid shortgrass steppe to the sub-humid tallgrass prairie of the Central Great Plains, yet few field data exist to evaluate these predictions. In this paper, we measured rates of net N mineralization, N in above- and belowground primary production, total soil organic matter N pools, soil inorganic N pools and capture in resin bags, decomposition rates, foliar (15)N, and N use efficiency (NUE) across a precipitation gradient. We found that net N mineralization did not increase across the gradient, despite more N generally being found in plant production, suggesting higher N uptake, in the wetter areas. NUE of plants increased with precipitation, and delta(15)N foliar values and resin-captured N in soils decreased, all of which are consistent with the hypothesis that N cycling is tighter at the wet end of the gradient. Litter decomposition appeared to play a role in maintaining this regional N cycling trend: litter decomposed more slowly and released less N at the wet end of the gradient. These results suggest that immobilization of N within the plant-soil system increases from semi-arid shortgrass steppe to sub-humid tallgrass prairie. Despite the fact that N pools increase along a bio-climatic gradient from shortgrass steppe to mixed grass and tallgrass prairie, this element becomes relatively more limiting and is therefore more tightly conserved at the wettest end of the gradient. Similar to findings from forested systems, our results suggest that grassland N cycling becomes more open to N loss with increasing aridity.

Fire can alter a multitude of ecosystem properties that have the potential to affect rates of litter decomposition and nitrogen dynamics. In this study, we examined the effect of long-term variation in fire frequency in Minnesota oak savanna on rates of litter mass loss of a common tree species (Quercus ellipsoidalis) to determine how site and intraspecific litter characteristics impacted by variation in fire frequency affect rates of decomposition, litter N dynamics, and litter microbial biomass. Although an increase in fire frequency resulted in higher litter temperatures, lower litter moisture, and decreased soil N and P availability, site characteristics had no net effect on rates of mass loss. Rather, litter C:N ratio, which increased with increased fire frequency, was the dominant predictor of rates of decomposition and litter N dynamics. Increased litter C:N led to decreased rates of decomposition and N immobilization, regardless of the characteristics of the site of decomposition. Therefore, it is the indirect effects of long-term variation in fire frequency on litter characteristics rather than fire's direct effects on site characteristics that determine fire effects on decomposition and N dynamics in this system. Slower rates of decomposition and increased N immobilization of litter produced in frequently burned sites may enhance fire-induced N losses, further decelerating rates of N cycling in frequently burned sites.

Riparian zones are an important transition between terrestrial and aquatic ecosystems, and they function in nutrient cycling and removal. Non-native earthworms invading earthworm-free areas of North America can affect nutrient cycling in upland soils and have the potential to affect it in riparian soils. We examined how the presence of earthworms can affect riparian nutrient cycling and nutrient delivery to streams. Two mesocosm experiments were conducted to determine how (1) the biomass of earthworms and (2) earthworm species can affect nutrient flux from riparian zones to nearby streams and how this flux can affect streamwater nutrients and periphyton growth. In separate experiments, riparian soil cores were amended with one of four mixed earthworm biomasses (0, 4, 10, or 23 g m(-2) ash-free dry mass) or with one of three earthworm species (Aporrectodea caliginosa, Lumbricus terrestris, L. rubellus) or no earthworm species. Riparian soil cores were coupled to artificial streams, and over a 36-day period, we measured nutrient leaching rates, in-stream nutrient concentrations, and periphyton growth. Ammonium leaching increased with increasing biomass and was greatest from the A. caliginosa treatments. Nitrate leaching increased through time and increased at a greater rate with higher biomass and from cores containing A. caliginosa. We suggest that the overall response of increased nitrate leaching [90% of total nitrogen (N)] was due to a combination of ammonium excretion and burrowing by earthworms, which increased nitrification rates. During both experiments, periphyton biomass increased through time but did not differ across treatments despite high in-stream inorganic N. Through time, in-stream phosphorus (P) concentration declined to <5 microg l(-1), and periphyton growth was likely P-limited. We conclude that activities of non-native earthworms (particularly A. caliginosa) can alter biogeochemical cycling in riparian zones, potentially reducing the N-buffering capacity of riparian zones and altering stoichiometric relationships in adjacent aquatic ecosystems.

Most explanations for the positive effect of plant species diversity on productivity have focused on the efficiency of resource use, implicitly assuming that resource supply is constant. To test this assumption, we grew seedlings of Echinacea purpurea in soil collected beneath 10-year-old, experimental plant communities containing one, two, four, eight, or 16 native grassland species. The results of this greenhouse bioassay challenge the assumption of constant resource supply; we found that bioassay seedlings grown in soil collected from experimental communities containing 16 plant species produced 70% more biomass than seedlings grown in soil collected beneath monocultures. This increase was likely attributable to greater soil N availability, which had increased in higher diversity communities over the 10-year-duration of the experiment. In a distinction akin to the selection/complementarity partition commonly made in studies of diversity and productivity, we further determined whether the additive effects of functional groups or the interactive effects of functional groups explained the increase in fertility with diversity. The increase in bioassay seedling biomass with diversity was largely explained by a concomitant increase in N-fixer, C4 grass, forb, and C3 grass biomass with diversity, suggesting that the additive effects of these four functional groups at higher diversity contributed to enhance N availability and retention. Nevertheless, diversity still explained a significant amount of the residual variation in bioassay seedling biomass after functional group biomass was included in a multiple regression, suggesting that interactions also increased fertility in diverse communities. Our results suggest a mechanism, the fertility effect, by which increased plant species diversity may increase community productivity over time by increasing the supply of nutrients via both greater inputs and greater retention.

The invasion of non-native plants can alter the diversity and activity of soil microorganisms and nutrient cycling within forests. We used field studies to analyze the impact of a successful invasive groundcover, Alliaria petiolata, on fungal diversity, soil nutrient availability, and pH in five northeastern US forests. We also used laboratory and greenhouse experiments to test three mechanisms by which A. petiolata may alter soil processes: (1) the release of volatile, cyanogenic glucosides from plant tissue; (2) the exudation of plant secondary compounds from roots; and (3) the decomposition of litter. Fungal community composition was significantly different between invaded and uninvaded soils at one site. Compared to uninvaded plots, plots invaded by A. petiolata were consistently and significantly higher in N, P, Ca and Mg availability, and soil pH. In the laboratory, the release of volatile compounds from the leaves of A. petiolata did not significantly alter soil N availability. Similarly, in the greenhouse, the colonization of native soils by A. petiolata roots did not alter soil nutrient cycling, implying that the exudation of secondary compounds has little effect on soil processes. In a leaf litter decomposition experiment, however, green rosette leaves of A. petiolata significantly increased the rate of decomposition of native tree species. The accelerated decomposition of leaf litter from native trees in the presence of A. petiolata rosette leaves shows that the death of these high-nutrient-content leaves stimulates decomposition to a greater extent than any negative effect that secondary compounds may have on the activity of the microbes decomposing the native litter. The results presented here, integrated with recent related studies, suggest that this invasive plant may change soil nutrient availability in such a way as to create a positive feedback between site occupancy and continued proliferation.

The large accumulation of organic matter in peatlands has been partially attributed to litter decomposition rates, which are slowed by a high water table. To test this, we examined whether there were significant differences in the decomposition and N and P dynamics of ten foliar litters and wood blocks at three pairs of upland forest and peatland sites in the transitional grassland, high boreal and low subarctic regions of central Canada, using litterbags collected over a 12-year period. At two of the three pairs, the decomposition rate, as determined by proportion of the original mass remaining after 12 years and by the exponential decay coefficient (k), was faster overall at the upland than at the peatland. In the third pair, there was no significant difference, despite the water table being close to the peat surface; warmer soil temperatures in the peatland than the upland may be the cause. In general, there were small losses or gains of N in the litters after 12 years, compared to the original litter, though there were some differences among litter types and sites, net gains in N likely reflecting the higher exogenous N availability. P was lost from most litters at the two northern pairs of sites, but at the transitional grassland pair, there were large net gains in P and greater variation among litters. The N:P ratio in the original litters ranged from 5 to 26 and after 12 years the ratio narrowed, with the site average of the ten litters ranging from 13 to 22, varying with the soil ratio. Decomposition rates and N and P dynamics after 12 years are different between upland and peatland sites: although the water table is a primary control on these differences, other factors such as temperature and soil nutrient status are also important.

Alien invasive plants are capable of modifying ecosystem function. However, it is difficult to make generalisations because impacts often appear to be species- and site-specific. In this study, we examined the impacts of seven highly invasive plant species in NW Europe (Fallopia japonica, Heracleum mantegazzianum, Impatiens glandulifera, Prunus serotina, Rosa rugosa, Senecio inaequidens, Solidago gigantea) on nutrient pools in the topsoil and the standing biomass. We tested if the impacts follow predictable patterns, across species and sites or, alternatively, if they are entirely idiosyncratic. To that end, we compared invaded and adjacent uninvaded plots in a total of 36 sites with widely divergent soil chemistry and vegetation composition. For all species, invaded plots had increased aboveground biomass and nutrient stocks in standing biomass compared to uninvaded vegetation. This suggests that enhanced nutrient uptake may be a key trait of highly invasive plant species. The magnitude and direction of the impact on topsoil chemical properties were strongly site-specific. A striking finding is that the direction of change in soil properties followed a predictable pattern. Thus, strong positive impacts (higher topsoil nutrient concentrations in invaded plots compared to uninvaded ones) were most often found in sites with initially low nutrient concentrations in the topsoil, while negative impacts were generally found under the opposite conditions. This pattern was significant for potassium, magnesium, phosphorus, manganese and nitrogen. The particular site-specific pattern in the impacts that we observed provides the first evidence that alien invasive species may contribute to a homogenisation of soil conditions in invaded landscapes.

Many regard the concentrations of nitrogen (N), tannins and plant cell wall constituents (fibre) as key indicators of food quality and habitat suitability for browsing herbivores; yet there is no method for measuring their combined effects. We have developed a simple in vitro assay for measuring the effects of tannins and fibre on N availability in browse. We determined the effects of tannins by measuring the polyethylene glycol (PEG)-binding capacity (PEG-BC) of Eucalyptus leaf samples, followed by a two-stage in vitro digestion with pepsin and cellulase to determine the digestibility of dry matter and N. There was a significant relationship between concentrations of digestible N and the PEG-BC of the leaves. Furthermore, adding PEG significantly improved the digestibility of N. Our results concur with in vivo observations from several mammalian species. This suggests that our method is effective for measuring the nutritional quality of browse and the benefits of adding PEG, providing some index of the detrimental effects of tannins. We further simplified the assay by removing the PEG step, allowing us to quickly analyse samples in bulk. Nevertheless, this simplified method is still not practical for analysing the many samples necessary to compare the nutritional values of different tracts of forest. We used near-infrared reflectance spectroscopy to produce calibration equations and predicted total and digestible N in 322 trees at eleven sites. Both within and between sites, we found a wide variation in concentrations of digestible N but a much lower variation in total N, with either no relationship or poor relationships between the two measures. This confirms the variability in the nutritional quality of eucalypt forests, which may explain the distribution and abundance of mammalian herbivores. Thus, our assay provides a useful tool for understanding how food resources influence herbivore populations at different scales.

The performance of individual phytoplankton species is strongly governed by the thermal stratification's impact on vertical mixing within the water column, which alters the position of phytoplankton relative to nutrients and light. The present study documents shifts in phytoplankton structure and vertical positioning that have accompanied intensified long-term stratification in a natural ecosystem. Ordination analysis is used to extract gradients in phytoplankton composition in Lake Tahoe, an extremely nutrient-poor lake, over a 23-year period of records. Community structure in the 1980s was associated most strongly with resource availability (low nitrogen to phosphorus ratios, deeper euphotic zone depth), while intensified stratification dominated the phytoplankton structure since the late 1990s. Within diatoms, small-sized cells increased with reduced mixing, suggesting that suppressed turbulence provides them with a competitive advantage over large-sized cells. Among the morphologically diverse chlorophytes, filamentous and coenobial forms were favored under intensified stratification. The selection for small-sized diatoms is accompanied by a shoaling trend in their vertical position in the water column. In contrast, the motile flagellates displayed a deeper vertical positioning in recent years, indicating that optimal growth conditions shifted likely due to reduced upwelling of nutrients. As the thermal stratification of lakes and oceans is strongly linked to climate variables, the present study confirms that climate warming will alter phytoplankton structure and dynamics largely through effects on nutrient availability and sinking velocities. Intensified stratification should favor the expansion of small-sized species and species with the capability of buoyancy regulation, which may alter primary productivity, nutrient recycling, and higher trophic productivity.

Short-term responses of producers highlight that key nutrients (e.g., N, P)-or combinations of these nutrients-limit primary production in aquatic and terrestrial ecosystems. These discoveries continue to provide highly valuable insights, but it remains important to ask whether nutrients always predominantly limit producers despite wide variation in nutrient supply and herbivory among systems. After all, predictions from simple food chain models (derived here) readily predict that limitation by grazers can exceed that by nutrients, given sufficient enrichment. However, shifts in composition of producers and/or increasing dominance of invulnerable stages of a producer can, in theory, reduce grazer limitation and retain primacy of nutrient limitation along nutrient supply gradients. We observed both mechanisms (inter- and intra-species variation in vulnerability to herbivory) working in a two-part mesocosm experiment. We incubated diverse benthic algal assemblages for several months either in the presence or absence of benthic macro-grazers in mesocosms that spread a broad range of nutrient supply. We then conducted short-term assays of nutrient and grazer limitation on these communities. In the "historically grazed" assemblages, we found shifts from more edible, better competitors to more resistant producers over enrichment gradients (as anticipated by the food web model built with a tradeoff in resistance vs. competitive abilities). However, contrary to our expectations, "historically ungrazed" assemblages became dominated by producers with vulnerable juvenile forms but inedible adult forms (long filaments). Consequently, we observed higher resource limitation rather than grazer limitation over this nutrient supply gradient in both "historically grazed" (expected) and "historically ungrazed" (not initially expected). Thus, via multiple, general mechanisms involving resistance to grazing (changes in species composition or variation in stage-structured vulnerability), producer assemblages should remain more strongly or as strongly limited by nutrients than grazers, even over large enrichment gradients.