Cascading effects of induced terrestrial plant defences on aquatic and terrestrial ecosystem function

Herbivores induce plants to undergo diverse processes that minimize costs to the plant, such as producing defences to deter herbivory or reallocating limited resources to inaccessible portions of the plant. Yet most plant tissue is consumed by decomposers, not herbivores, and these defensive processes aimed to deter herbivores may alter plant tissue even after detachment from the plant. All consumers value nutrients, but plants also require these nutrients for primary functions and defensive processes. We experimentally simulated herbivory with and without nutrient additions on red alder (Alnus rubra), which supplies the majority of leaf litter for many rivers in western North America. Simulated herbivory induced a defence response with cascading effects: terrestrial herbivores and aquatic decomposers fed less on leaves from stressed trees. This effect was context dependent: leaves from fertilized-only trees decomposed most rapidly while leaves from fertilized trees receiving the herbivory treatment decomposed least, suggesting plants funnelled a nutritionally valuable resource into enhanced defence. One component of the defence response was a decrease in leaf nitrogen leading to elevated carbon : nitrogen. Aquatic decomposers prefer leaves naturally low in C : N and this altered nutrient profile largely explains the lower rate of aquatic decomposition. Furthermore, terrestrial soil decomposers were unaffected by either treatment but did show a preference for local and nitrogen-rich leaves. Our study illustrates the ecological implications of terrestrial herbivory and these findings demonstrate that the effects of selection caused by terrestrial herbivory in one ecosystem can indirectly shape the structure of other ecosystems through ecological fluxes across boundaries.     

RESPONSE OF THE ANNUAL ABUTILON THEOPHRASTI MEDIC. (MALVACEAE) TO TIMING OF NUTRIENT AVAILABILITY

Because resource variability is likely to be high in disturbed environments, colonizing species should be opportunistic in their abilities to use resources at any time during the growing season. We examined the effects of timing of nutrient availability on seed quantity and quality in the colonizing annual Abutilon theophrasti. Plants were grown under six different nutrient treatments involving additions of different size, frequency and timing but having the same total amount of nutrients in all treatments. The plants showed an opportunistic response, being able to use nutrients supplied at different times for reproduction. However, when nutrient availability was delayed until fruit maturation had begun, seed number and total seed weight were reduced, production of mature fruits was delayed, and seed P concentrations were reduced. Seeds from the first two fruits of the plants had higher N concentration (mg/g), higher N content (mg/seed), lower P concentration, higher K content, and higher mean seed weight.     

Effects of Land Use on Lake Nutrients: The Importance of Scale,Hydrologic Connectivity,and Region

Catchment land uses, particularly agriculture and urban uses, have long been recognized as major drivers of nutrient concentrations in surface waters. However, few simple models have been developed that relate the amount of catchment land use to downstream freshwater nutrients. Nor are existing models applicable to large numbers of freshwaters across broad spatial extents such as regions or continents. This research aims to increase model performance by exploring three factors that affect the relationship between land use and downstream nutrients in freshwater: the spatial extent for measuring land use, hydrologic connectivity, and the regional differences in both the amount of nutrients and effects of land use on them. We quantified the effects of these three factors that relate land use to lake total phosphorus (TP) and total nitrogen (TN) in 346 north temperate lakes in 7 regions in Michigan, USA. We used a linear mixed modeling framework to examine the importance of spatial extent, lake hydrologic class, and region on models with individual lake nutrients as the response variable, and individual land use types as the predictor variables. Our modeling approach was chosen to avoid problems of multi-collinearity among predictor variables and a lack of independence of lakes within regions, both of which are common problems in broad-scale analyses of freshwaters. We found that all three factors influence land use-lake nutrient relationships. The strongest evidence was for the effect of lake hydrologic connectivity, followed by region, and finally, the spatial extent of land use measurements. Incorporating these three factors into relatively simple models of land use effects on lake nutrients should help to improve predictions and understanding of land use-lake nutrient interactions at broad scales.     

Development of an epiphyte indicator of nutrient enrichment: A critical evaluation of observational and experimental studies

An extensive review of the literature describing epiphytes on submerged aquatic vegetation (SAV), especially seagrasses, was conducted in order to evaluate the evidence for response of epiphyte metrics to increased nutrients. Evidence from field observational studies, together with laboratory and field mesocosm experiments, was assembled from the literature and evaluated for a hypothesized positive response to nutrient addition. There was general consistency in the results to confirm that elevated nutrients tended to increase the load of epiphytes on the surface of SAV, in the absence of other limiting factors. In spite of multiple sources of uncontrolled variation, positive relationships of epiphyte load to nutrient concentration or load (either nitrogen or phosphorus) often were observed along strong anthropogenic or natural nutrient gradients in coastal regions. Such response patterns may only be evident for parts of the year. Results from both mesocosm and field experiments also generally support the increase of epiphytes with increased nutrients, although outcomes from field experiments tended to be more variable. Relatively few studies with nutrient addition in mesocosms have been done with tropical or subtropical species, and more such controlled experiments would be helpful. Experimental duration influenced results, with more positive responses of epiphytes to nutrients at shorter durations in mesocosm experiments versus more positive responses at longer durations in field experiments. In the field, response of epiphyte biomass to nutrient additions was independent of climate zone. Mesograzer activity was a critical covariate for epiphyte response under experimental nutrient elevation, but the epiphyte response was highly dependent on factors such as grazer identity and density, as well as nutrient and ambient light levels. The balance of evidence suggests that epiphytes on SAV will be a useful indicator of persistent nutrient enhancement in many situations. Careful selection of appropriate temporal and spatial constraints for data collection, and concurrent evaluation of confounding factors will help increase the signal to noise ratio for this indicator.     

Impact of sediment and nutrient inputs on growth and survival of tadpoles of the Western Toad

1. Sediment and nutrient loading in freshwater systems are leading causes of aquatic habitat degradation globally. We investigated the impacts of fine-sediment and nutrient additions on the growth and survival of western toad ( Bufo boreas ) tadpoles and emergent metamorphs in mesocosm and exclosure experiments.
2. Mesocosm tanks received weekly pulses of fine sediments to create initial concentrations of 0, 130 and 260 mg L⁻¹ of suspended sediment and either bi-weekly additions of nutrients (N = 160 μg L⁻¹, P = 10 μg L⁻¹) or no additions in a factorial design. Within mesocosms, tadpole exclosures allowed for quantification of tadpole grazing pressure on periphyton biomass, chlorophyll- a and sediment deposition.
3. Tadpoles receiving sediment additions experienced slower growth rates and reduced survival to metamorphosis, although no effects of treatment were detected on size at metamorphosis or time to metamorphosis. Nutrient additions also lowered survival, but had no impact on other measured parameters of tadpole fitness. Dissections and gut content analysis revealed that tadpoles ingested sediment in large quantities altering the proportion of the organic content of ingested food.
4. Together these results suggest that although sediment was readily consumed by tadpoles, its presence in the larval environment had an overall negative effect on tadpole growth and survival, although not as severe as predicted.     

Tropical Andean Forests Are Highly Susceptible to Nutrient Inputs—Rapid Effects of Experimental N and P Addition to an Ecuadorian Montane Forest

Tropical regions are facing increasing atmospheric inputs of nutrients, which will have unknown consequences for the structure and functioning of these systems. Here, we show that Neotropical montane rainforests respond rapidly to moderate additions of N (50 kg ha⁻¹ yr⁻¹) and P (10 kg ha⁻¹ yr⁻¹). Monitoring of nutrient fluxes demonstrated that the majority of added nutrients remained in the system, in either soil or vegetation. N and P additions led to not only an increase in foliar N and P concentrations, but also altered soil microbial biomass, standing fine root biomass, stem growth, and litterfall. The different effects suggest that trees are primarily limited by P, whereas some processes—notably aboveground productivity—are limited by both N and P. Highly variable and partly contrasting responses of different tree species suggest marked changes in species composition and diversity of these forests by nutrient inputs in the long term. The unexpectedly fast response of the ecosystem to moderate nutrient additions suggests high vulnerability of tropical montane forests to the expected increase in nutrient inputs.     

The effect of nutrient additions on bacterial activity in seagrass (Posidonia oceanica) sediments

The influence of nutrient additions on benthic bacterial activity under seagrass meadows was tested by enriching five seagrass (Posidonia oceanica) meadows with nutrients over one year. We found a highly significant response of benthic bacterial activity to nutrient additions, which was reflected in greater (about two-fold) ammonification rates and, to a smaller extent, a significant tendency for a greater exoenzymatic activity. Nutrient additions significantly raised bacterial activity, without altering the seasonal changes in bacterial activity. As a result of the increased bacterial activity, the organic content of the sediments declined significantly, by about 33%, after one year of nutrient addition. Hence, nutrient additions to the seagrass meadows enhance seagrass production but also accelerate bacterial decomposition of seagrass carbon, thereby reducing the capacity of the sediments to store organic carbon. These results demonstrate that sediment nutrient availability limits bacterial activity in these Posidonia oceanica meadows, and identify bacteria as important nutrient consumers in these systems.     

The Effects of Ultraviolet Radiation and Nutrient Additions on Periphyton Biomass and Composition in a Sub‐Alpine Lake (Castle Lake,USA)

Rising levels of ultraviolet radiation (UVR) striking the Earth's surface have led to numerous studies assessing its inhibitory effects on phytoplankton and periphyton in aquatic systems. Mineral nutrients such as nitrogen (N) and phosphorus (P) have been shown to increase aspects of algal metabolism and compensate for UVR inhibition. An in situ substratum enrichment technique and UV shielding was used to assess the effects of nutrient additions on periphyton exposed to different levels of UVR in Castle Lake, California during July‐August, 1997. UV shielding had no effect on total periphyton biomass, but caused shifts in species composition. The dominant periphyton species, Anabaena circinalis RAB., demonstrated sensitivity to ambient levels of UV radiation possibly due to UV inhibition of N2 ‐fixation. Total diatom biovolume decreased when shielded from UVR. Phosphorus additions continually elicited an increase in periphyton biovolume at all levels of analysis. These results suggest an interaction between nutrient status/availability and UV sensitivity.     

Waterfowl feces as a source of nutrients to a prairie wetland: responses of microinvertebrates to experimental additions

We tested the hypothesis that inorganic nutrients released fromwaterfowl feces would stimulate primary production, therebyaffecting microinvertebrate grazers, by making controlledadditions of waterfowl feces to fishless wetland enclosures andmeasuring the response of planktonic and phytophilouscladocerans, copepods, and rotifers. Feces were added in twopulses, four weeks apart, to duplicate enclosures at a ‘high’level (115 g m⁻² wet feces), simulating the total P load(1.6 g m⁻²) applied in an earlier fertilizationexperiment, and a ‘low’ level (11.5 g m⁻²). Density ofmicrocrustacean grazers in the water column increased inresponse to both feces additions, although the response wasmore noticeable after the second feces addition. After eachaddition, cladocerans (predominantly Ceriodaphnia dubia)and copepodites in the water column (and associated withperiphyton on acrylic rods in the water column) were mostabundant in enclosures with high loading. In contrast, densityof microcrustacean grazers associated with macrophytes(predominantly Chydorus spp. and copepodites) increasedin response to the second feces addition only.Microinvertebrate density increased only slightly with lowfeces loading. Community composition showed similar changesover the season in all enclosures, and differences in relativeabundance were not attributable to treatment effects. Given thesmall effects produced by nutrient additions that greatlyexceed natural loadings, nutrients leaching from waterfowlfeces do not appear to have a significant impact on the foodwebof this wetland. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interactive effects of porewater nutrient enrichment, bioturbation and sediment characteristics on benthic assemblages in sandy sediments

Effective management of eutrophic ecosystems requires an understanding of how nutrient input affects the structure and function of benthic communities. The effects of nutrients in soft sediment habitats can be influenced by a variety of factors including sediment characteristics, hydrodynamic exposure, and the presence of bioturbating macroinvertebrates. We used a large scale exclusion experiment (400 m² areas, n = 6) to test if bioturbating lugworms, Arenicola marina mediate the effects of nutrient enrichment. We incorporated small plots (30 × 30 cm) dosed with household garden fertilizer within the lugworm exclusion and corresponding control areas and predicted that the effects of nutrient enrichment would be greater in the absence of lugworms. We found that the increases in nutrient concentrations were higher in the absence of lugworms, but only in the less permeable sediment in the low intertidal zone compared to the more permeable sediment in the high intertidal. Contrary to expectations, the accumulation of nutrients in the plots did not affect the organic matter and chlorophyll levels in the sediment. Interestingly, there were overall negative effects of nutrient additions on some of the most abundant molluscs, Hydrobia ulvae, Retusa obtusa and juvenile Cerastoderma edule. Possible explanations for these adverse effects such as the changes in the sediment chemistry or the physical presence of the fertilizer in the sediment caused by the nutrient additions are discussed. We conclude that the effects of nutrient enrichment in soft sediment habitats on benthic assemblages are determined by the interplay between the presence of bioturbating macroinvertebrates, tidal height and sediment characteristics.     

Effects of mineral turbidity on freshwater plankton communities: Three exploratory tank experiments of factorial design

We conducted three exploratory experiments of factorial design on the effects of mineral turbidity in 18 large (7 000 1) fiberglass tanks containing plankton communities derived from Lake Texoma (Oklahoma-Texas), a large reservoir often subject to turbid inflows. Replicate plankton communities developed for 30–45 d in response to 4–9 treatments of planktivorous fish, dead fish, nutrient additions and artificial removal of zooplankton with plankton netting (or combinations of these manipulations). In one experiment we then added three concentrations of kaolin and in the other two experiments we added bentonite or powdered silica to the replicate tank communities. We measured the responses of the diverse plankton communities to the addition of mineral turbidity for an additional period of 30–45 d.Effects of introduced minerals (quality or concentration effects) and food web treatments after mineral addition (fish, nutrients, etc.) were determined by analysis of covariance after observations were adjusted for the value of each response variable at the time of mineral addition. Covariance analysis resulted in a statistical model explaining > 80% of the variance for most response variables. The values of response variables at the time of mineral addition were often the most important source of variation, suggesting the importance of biotic community resiliency to the effects of mineral turbidity. There were few effects of mineral particles on physical or chemical (temperature, conductivity, oxygen, pH), nutrient (nitrate, phosphate, alkalinity) or biotic (algal or zooplankton populations) components of the tank communities. Mineral effects were found for several measures of water clarity (Secchi depth, turbidity, and the concentration of small sestonic particles).     

Limitations to CO2-induced growth enhancement in pot studies

Recently, it has been suggested that small pots may reduce or eliminate plant responses to enriched CO₂ atmospheres due to root restriction. While smaller pot volumes provide less physical space available for root growth, they also provide less nutrients. Reduced nutrient availability alone may reduce growth enhancement under elevated CO₂. To investigate the relative importance of limited physical rooting space separate from and in conjunction with soil nutrients, we grew plants at ambient and double-ambient CO₂ levels in growth containers of varied volume, shape, nutrient concentration, and total nutrient content. Two species (Abutilon theophrasti, a C₃ dicot with a deep tap root andSetaria faberii, a C₄ monocot with a shallow diffuse root system) were selected for their contrasting physiology and root architecture. Shoot demography was determined weekly and biomass was determined after eight and ten weeks of growth. Increasing total nutrients, either by increasing nutrient concentration or by increasing pot size, increased plant growth. Further, increasing pot size while maintaining equal total nutrients per pot resulted in increased total biomass for both species. CO₂-induced growth and reproductive yield enhancements were greatest in pots with high nutrient concentrations, regardless of total nutrient content or pot size, and were also mediated by the shape of the pot. CO₂-induced growth and reproductive yield enhancements were unaffected by pot size (growth) or were greater in small pots (reproductive yield), regardless of total nutrient content, contrary to predictions based on earlier studies. These results suggest that several aspects of growth conditions within pots may influence the CO₂ responses of plants; pot size, pot shape, the concentration and total amount of nutrient additions to pots may lead to over-or underestimates of the CO₂ responses of real-world plants.     

Effects of nutrient supplementation on host‐pathogen dynamics of the amphibian chytrid fungus: a community approach

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Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

Resources drive trade‐off between viral lifestyles in the plankton: evidence from freshwater microbial microcosms

Viruses have evolved different life strategies for coping with environmental challenges and this is a key explanation for their omnipresence in aquatic systems. However, factors that determine the balance between lytic versus lysogenic decision within natural virioplankton are poorly documented, primarily in freshwaters. This study was designed to investigate the experimental short‐term (24 h incubation) effects of added organic and inorganic nutrients on the two viral lifestyles in nutrient‐depleted freshwater microbial (i.e. < 0.8 μm fraction) microcosms, using mitomycin C as prophage inductor agent. In the absence of mitomycin, viral lytic production increased as a functional response to the strong stimulation of bacterial growth rates (0.7–0.8 day^?1) by the added nutrients, primarily the organic nutrients which appeared scarcer than inorganic nutrients and was related to the sampling period and the geomorphological peculiarities of Lake Pavin. In the presence of mitomycin, temperate phage production (frequency of lysogenically infected bacterial cells, FLC = 17–19% of total cells) significantly exceeded lytic production (frequency of lytically infected bacterial cells, FIC = 9–11%) in natural samples (i.e. without nutrient additions) as a result of enhanced prophage induction, which relatively increased with the decreasing contact probability between viruses and their potential hosts. In contrast, addition of nutrients drastically reduced FLC (< 4%) and increased FIC (> 22%). Both variables were antagonistically correlated as was the correlation between FLC and bacterial growth rates, supporting the idea that lysogeny may represent a maintenance strategy for viruses in harsh nutrient/host conditions which appeared as major instigators of the trade‐off between the two viral lifestyles. Overall, at the community level, we reject the hypothesis that nutrients but mitomicyn C stimulate temperate phage induction, and retained the hypotheses that nutrients rather (i) stimulate lytic viruses via enhanced host growth and (ii) when limiting, promote lysogenic conversion in natural waters.     

Drivers of Variation in Aboveground Net Primary Productivity and Plant Community Composition Differ Across a Broad Precipitation Gradient

Aboveground net primary production (ANPP) is a key integrator of C uptake and energy flow in many terrestrial ecosystems. As such, ecologists have long sought to understand the factors driving variation in this important ecosystem process. Although total annual precipitation has been shown to be a strong predictor of ANPP in grasslands across broad spatial scales, it is often a poor predictor at local scales. Here we examine the amount of variation in ANPP that can be explained by total annual precipitation versus precipitation during specific periods of the year (precipitation periods) and nutrient availability at three sites representing the major grassland types (shortgrass steppe, mixed-grass prairie, and tallgrass prairie) spanning the broad precipitation gradient of the U.S. Central Great Plains. Using observational data, we found that precipitation periods and nutrient availability were much stronger predictors of site-level ANPP than total annual precipitation. However, the specific nutrients and precipitation periods that best predicted ANPP differed among the three sites. These effects were mirrored experimentally at the shortgrass and tallgrass sites, with precipitation and nutrient availability co-limiting ANPP, but not at the mixed-grass site, where nutrient availability determined ANPP exclusive of precipitation effects. Dominant grasses drove the ANPP response to increased nutrient availability at all three sites. However, the relative responses of rare grasses and forbs were greater than those of the dominant grasses to experimental nutrient additions, thus potentially driving species turnover with chronic nutrient additions. This improved understanding of the factors driving variation in ANPP within ecosystems spanning the broad precipitation gradient of the Great Plains will aid predictions of alterations in ANPP under future global change scenarios.     

PARTICULATE ORGANIC MATTER AS AN ALTERNATIVE NUTRIENT SOURCE FOR TROPICAL SARGASSUM SPECIES (FUCALES, PHAEOPHYCEAE)

Large Fucales are abundant on coastal coral reefs of the Great Barrier Reef, but are often limited by the availability of inorganic nutrients. Particle loads in these reef waters are high, which is generally perceived as detrimental for aquatic plants due to a reduction of light. Here, I provide evidence that several abundant Sargassum species supplement their nutrient supply with nutrients derived from the layer of particulate matter (PM) deposited on their thalli. In experiments involving removal or addition of PM, growth rates of Sargassum spp. were up to 180% higher when particles were present on the thalli. Tissue nitrogen and phosphorus levels of thalli with a surface PM layer were 10%–30% higher than those of thalli without PM. The amount of PM deposited in situ on thalli of five species of brown algae ranged from 0.6 to 0.9 g C_org·g⁻¹ dry weight alga, depending on the species' morphology. I suggest that a nutrient-rich diffusive boundary layer is created on the thallus surface by an epiphytic microbial community that remineralizes the particulate nutrients. When algal growth is nutrient limited, the use of particle-derived nutrients as a source alternative to nutrients in the water column may outweigh any potential adverse effects of the thallus particle layer.     

Tissue nutrient concentrations in freshwater aquatic macrophytes: high inter-taxon differences and low phenotypic response to nutrient supply

1. The elemental composition and stoichiometry of aquatic plants has often been suggested to reflect the nutrient enrichment of aquatic habitats. However, the relationship is often weak. Moreover, uncertainties remain in the relevance of laboratory derived critical plant tissue nutrient concentrations to maximum yield or growth rates in the field.
2. Aquatic vascular plants and bryophytes, overlying water and sediment samples were collected to test whether freshwater aquatic macrophytes: (i) show tissue nutrient deficiencies when growing in oligotrophic freshwater habitats, and (ii) have strict homeostatic stoichiometry.
3. Plant nutrient concentrations were significantly related to total inorganic nitrogen (or nitrate), total dissolved phosphorus and sediment total phosphorus. However, these relationships were weak. Virtually all the variance in plant tissue nutrient concentrations, however, could be explained by species (taxon) identity.
4. Critical tissue nutrient concentrations for 95% maximum yield or 95% maximum growth rate in aquatic angiosperms, determined from laboratory bioassays, suggested that nutrients should not limit yield in wild aquatic macrophytes. However, there were a substantial number of samples where potential growth rate limitation was possible, particularly due to phosphorus.
5. Strict C : N : P stoichiometric ratios were found for both vascular plants and bryophytes, suggesting little scope for plants as indicators of nutrient enrichment, but provide robust stoichiometric data for studies on ecosystem metabolism and nutrient cycling.     

The effect of different nutrient concentrations on the growth rate and nitrogen storage of watercress (Nasturtium officinale R. Br.)

The mechanisms that allow broadly distributed aquatic plants to inhabit variable resource environments are unclear, yet understanding these mechanisms is important because broad environmental tolerance is often linked to invasiveness in terrestrial and aquatic plants. In an experimental stream, we examined the effects of different nutrient concentrations on the growth rate, biomass, and foliar nutrient concentrations of a cosmopolitan and potentially invasive aquatic plant, Nasturtium officinale (R. Br.). Nasturtium seedlings were grown under six nutrient treatment levels ranging from 0.64 μm N:0.09 μm P to 1531 μm N:204.13 μm P, for 8 weeks. Absolute and relative growth rates, and biomass of seedlings increased along a gradient of increasing nutrient concentrations but the effect of nutrient concentration was dependent on growing time. Seedling biomass varied among nutrient treatments in weeks 4 through 8 of the experiment, but did not differ in week 2. By week 8, the two highest nutrient treatments had greater biomass than the two lowest nutrient treatments. Foliar nitrogen concentration increased, whereas carbon concentration and C:N ratios decreased in response to increasing nutrients. Nasturtium grows slowly in nutrient-poor conditions but rapidly increases its growth, biomass accrual, and nitrogen storage as conditions become nutrient-rich. The response of Nasturtium to enhanced nutrient conditions may indicate how aquatic nuisance species successfully invade and dominate plant communities in streams, where resources often vary both temporally and spatially.     

Nutrient Limitation to Primary Productivity in a Secondary Savanna in Venezuela1

We examined nutrient limitation to primary productivity in a secondary savanna in the interior branch of the Coastal Range of Venezuela, which was converted from forest to savanna more than 100 years ago. We manipulated soil nutrients by adding nitrogen (+N), phosphorus and potassium (+PK), and nitrogen, phosphorus, and potassium (+NPK) to intact savanna. Eleven months after fertilization, we measured aboveground biomass and belowground biomass as live fine roots in the top 20 cm of soil, and species and functional group composition in response to nutrient additions. Aboveground biomass was highest in the NPK treatment ([mean g/m2]; control = 402, +N = 718, +PK = 490, +NPK = 949). Aboveground production, however, appeared to be limited primarily by N. Aboveground biomass increased 78 percent when N was added alone but did not significantly respond to PK additions when compared to controls. In contrast to aboveground biomass, belowground biomass increased with PK additions but showed no significant increase with N (depth 0–20 cm; [mean g/m2]; control = 685, +N = 443, +PK = 827, +NPK = 832). There was also a 36 percent increase in root length with PK additions when compared to controls. Whole savanna shoot:root ratios were similar for control and +PK (0.6), while those for +N or +NPK fertilization were significantly higher (1.7 and 1.2, respectively). Total biomass response (above + belowground) to nutrient additions showed a strong N and PK co‐limitation ([mean g/m2]; control = 1073, +N = 1111, +PK = 1258, +NPK = 1713). Aboveground biomass of all monocots increased with N additions, whereas dicots showed no response to nutrient additions. Trachypogon spp. (T. plumosus+T. vestitus) and Axonopus canescens, the two dominant grasses, made up more than 89 percent of the total aboveground biomass in these sites. Trachypogon spp. responded to NPK, whereas A. canescens, sedges, and the remaining monocots only responded to N. Even though nutrient additions resulted in higher aboveground biomass in N and NPK fertilized plots, this had little effect on plant community composition. With the exception of sedges, which responded positively to N additions and increased from 4 to 8 percent of die plant community, no changes were observed in plant community composition after 11 months.