Nutrient Enrichment Treatments Research Articles

Artificial night‐time lighting and nutrient enrichment synergistically favour the growth of alien ornamental plant species over co‐occurring native plants

Abstract Insights into ecological drivers of alien plant invasions can be gained through comparative studies of growth and fecundity of invasive alien plants versus those of co‐occurring non‐invasive alien plants and native plants across environmental conditions in common garden settings. Habitats that harbour alien plant species in many ecosystems globally are presently experiencing light pollution resulting from artificial light at night (ALAN) and increased rates of nutrient enrichment of the soil. However, the potential interactive effects of ALAN and nutrient enrichment on invasiveness of alien plant species remain unknown. Here, we performed a common‐garden experiment to test the interactive effects of ALAN and soil nutrient enrichment on the growth of a random set of ten alien (five invasive and five non‐invasive naturalized) and seven co‐occurring native ornamental plant species that are commonly cultivated within urban and peri‐urban areas of Nairobi city in Kenya. We predicted that a simultaneous increase in photoperiod via ALAN and nutrient enrichment will favour growth of invasive alien plant species over that of non‐invasive alien and native plant species. We grew the 17 plant species under natural daylight (ALAN−) versus natural daylight followed by ALAN (ALAN+) and fully crossed with two levels of nutrient enrichment (low vs. high) and competition (competition vs. no‐competition against a native plant Ocimum gratissimum) treatments. Under simultaneous high‐nutrient and no‐competition treatments, ALAN enhanced mean total biomass of invasive and non‐invasive naturalized alien species by 61.1% and 131.4%, respectively but decreased that of native plant species by 34%. In contrast, under simultaneous high‐nutrient and competition treatments, ALAN enhanced mean total biomass of invasive alien plant species by 68.6% and that of non‐invasive naturalized alien species by 51.9% and native species by 35.4%. High‐nutrient treatment enhanced flower formation more strongly in invasive and non‐invasive naturalized alien plants than in native plants. The invasive and non‐invasive naturalized alien species grew taller than native species across the light, nutrient, and competition treatments. Synthesis: The present findings suggest that light pollution and nutrient enrichment may jointly confer growth advantage to invasive alien plant species over that of co‐occurring native plant species and enhance invasiveness of alien plant species.

Nutrient enrichment promotes invasion success of alien plants via increased growth and suppression of chemical defenses.

In support of the prediction of the enemy release hypothesis regarding a growth-defense trade-off, invasive alien plants often exhibit greater growth and lower anti-herbivory defenses than native plants. However, it remains unclear how nutrient enrichment of invaded habitats may influence competitive interactions between invasive alien and co-occurring native plants, as well as production of anti-herbivore defense compounds, growth-promoting hormones, and defense-regulating hormones by the two groups of plants. Here, we tested whether: (i) nutrient enrichment causes invasive alien plants to produce greater biomass and lower concentrations of the defense compounds flavonoids and tannins than native plants; and (ii) invasive alien plants produce lower concentrations of a defense-regulating hormone jasmonic acid (JA) and higher concentrations of a growth-promoting hormone gibberellic acid (GA3). In a greenhouse experiment, we grew five congeneric pairs of invasive alien and native plant species under two levels each of nutrient enrichment (low vs. high), simulated herbivory (leaf clipping vs. no-clipping), and competition (alone vs. competition) in 2.5-L pots. In the absence of competition, high-nutrient treatment induced a greater increase in total biomass of invasive alien species than that of native species, whereas the reverse was true under competition as native species benefitted more from nutrient enrichment than invasive alien species. Moreover, high-nutrient treatment caused a greater increase in total biomass of invasive alien species than that of native species in the presence of simulated herbivory. Competition induced higher production of flavonoids and tannins. Simulated herbivory induced higher flavonoid expression in invasive alien plants under low-nutrient than high-nutrient treatments. However, flavonoid concentrations of native plants did not change under nutrient enrichment and simulated herbivory treatments. Invasive alien plants produced higher concentrations of GA3 than native plants. Taken together, these results suggest that impact of nutrient enrichment on growth of invasive alien and co-occurring native plants may depend on the level of competition that they experience. Moreover, invasive alien plants might adjust their flavonoid-based defense more efficiently than native plants in response to variation in soil nutrient availability and herbivory pressure. Our findings suggest that large-scale efforts to reduce nutrient enrichment of invaded habitats may help to control future invasiveness of target alien plant species.

Fluctuations in resource availability shape the competitive balance among non-native plant species.

Fluctuating resource availability plays a critical role in determining non-native plant invasions through mediating the competitive balance between non-native and native species. However, the impact of fluctuating resource availability on interactions among non-native species remains largely unknown. This represents a barrier to understanding invasion mechanisms, particularly in habitats that harbor multiple non-native species with different responses to fluctuating resource availability. To examine the responses of non-native plant species to nutrient fluctuations, we compared the growth of each of 12 non-native species found to be common in local natural areas to nutrients supplied at a constant rate or supplied as a single large pulse in a pot experiment. We found that seven species produced more biomass with pulsed nutrients compared to constant nutrients (hereafter "benefitting species"), while the other five species did not differ between nutrient enrichment treatments (hereafter "non-benefitting species"). To investigate how nutrient fluctuations influence the interactions among non-native plant species, we established experimental non-native communities in the field with two benefitting and two non-benefitting non-native species. Compared with constant nutrient supply, the single large pulse of nutrient did not influence community biomass, but strongly increased the biomass and cover of the benefitting species and decreased those of the non-benefitting species. Furthermore, the benefitting species had higher leaf N content and greater plant height when nutrients were supplied as a single large pulse than at a constant rate, whereas the non-benefitting species showed no differences in leaf N content and were shorter when nutrients were supplied as a single large pulse than at a constant rate. Our results add to the growing evidence that the individual responses of non-native species to nutrient fluctuation are species-specific. More importantly, benefitting species were favored by nutrients coming in a pulse, while non-benefitting ones were favored by nutrients coming constantly when they grew together. This suggests that nutrient fluctuations can mediate the competitive balance among non-native plants and may thus determine their invasion success in a community harboring multiple non-native plant species.

Activity and abundance of methane-oxidizing bacteria on plants in experimental lakes subjected to different nutrient and warming treatments

Shallow lakes produce and emit substantial amounts of methane (CH4). Part of the CH4 produced in lakes is consumed by methane-oxidizing bacteria (MOB) present in the sediment and water column, thus reducing the overall CH4 emissions. However, the role of aquatic plants as habitat for CH4 oxidation by MOB is poorly understood. In this study, we compared CH4 oxidation rates and MOB abundance associated with different types of aquatic plants (periphyton, filamentous algae, and both above-ground macrophytes and their rhizosphere). The plants were collected from shallow lake mesocosms exposed to experimental nutrient enrichment and warming treatments for 17 years prior to this study. Incubations of all sampled plants showed CH4 oxidation, with above-ground macrophyte tissue and filamentous algae having the highest rates of up to 0.25 µmol CH4 h−1 g−1 dw. Oxidation rates associated with macrophytes were species dependent, with consumption rates on rhizospheres of Potamogeton crispus higher than those on Elodea canadensis. The increase in nutrients and dissolved CH4 in the water tended to increase MOB abundance and activity for all plant types, while no effect of long term warming was detectable. Our results showed that MOB associated with periphyton, filamentous algae and macrophytes oxidize CH4 in shallow lakes at different rates across species or plant types. We also found that high macrophyte biomass is associated with reduced CH4 concentration in the water. This study shows that CH4 oxidation occurs on many plant surfaces but that oxidation rates alone cannot explain the reduced CH4 emissions at higher plant biomass.

Plant community re-organization and increased productivity due to multi-year nutrient enrichment of a coastal grassland.

Nutrient enrichment alters plant community structure and function at a global scale. Coastal plant systems are expected to experience increased rates of nitrogen and phosphorus deposition by 2100, caused mostly by anthropogenic activity. Despite high density of studies investigating connections between plant community structure and ecosystem function in response to nutrient addition, inconsistencies in system response based on the ecosystem in question calls for more detailed analyses of nutrient impacts on community organization and resulting productivity response. Here, we focus on nutrient addition impacts on community structure and organization as well as productivity of different lifeforms in a coastal grassland. We established long-term nutrient enrichment plots in 2015 consisting of control (C), nitrogen (N), phosphorus (P), and nitrogen + phosphorus (NP) treatments. In 2017 we collected graminoid and forb productivity, root productivity, and community composition for each plot. We found no N x P interaction, but N enrichment was a significant main effect on productivity, highlighting N limitation in coastal systems. Importantly, nutrient enrichment treatments did not alter root productivity. However, all treatments caused significant differences in community composition. Using rank abundance curves, we determined that community composition differences were driven by increased dominance of nitrophilous graminoids, re-organization of subordinate species, and species absences in N and NP plots. Results of this study highlight how coastal grassland communities are impacted by nutrient enrichment. We show that community re-organization, increased dominance, and absence of critical species are all important mechanisms that reflect community-level impacts of nutrient enrichment in our coastal grassland site.

Responses of Enterococcus faecalis resistance and cytolysin up-regulation to nutrients in constructed mesocosms

Spreading of multidrug-resistant bacteria has become a growing and significant threat to environmental and public health. Our understanding of environmental variables such as nutrients which contribute to the dissemination of antibiotic resistance and pathogenesis is still limited. In this study, we operated outdoor mesocosm experiments to evaluate how nitrate and phosphate ions with different concentrations influence on an adaptation of Enterococcus faecalis to aquatic environments. E. faecalis were isolated from the mesocosms at 8 sampling events for 96 days to evaluate biofilm production, quorum-sensing-related genes expression, and sensitivity to oxytetracycline, erythromycin, ciprofloxacin, ampicillin, vancomycin, and chloramphenicol. Quantitative real-time PCR was used to compare mRNA levels of E. faecalis quorum-sensing-related genes. E. faecalis isolates exhibited resistance to oxytetracycline, ampicillin, and ciprofloxacin, respectively. We observed that the biofilm production of E. faecalis isolates on day 60 and 96 was significantly increased (p < 0.01). The expressions of quorum-sensing-related genes were significantly up-regulated (∼tenfold) at the transcriptional level in nutrient-enriched treatments. Our findings indicate that nitrate and phosphate ions facilitate resistance to commonly used antibiotics, increase biofilm production and intra-species communication, which could be a major reviving strategy of E. faecalis in the aquatic environments.

The decomposition process and nutrient release of invasive plant litter regulated by nutrient enrichment and water level change.

Wetlands are vulnerable to plant invasions and the decomposition of invasive plant litter could make impacts on the ecosystem services of wetlands including nutrient cycle and carbon sequestration. However, few studies have explored the effects of nutrient enrichment and water level change on the decomposition of invasive plant litter. In this study, we conducted a control experiment using the litterbag method to compare the decomposition rates and nutrient release in the litter of an invasive plant Alternanthera philoxeroides in three water levels and two nutrient enrichment treatments. This study found that the water level change and nutrient enrichment showed significant effects on the litter decomposition and nutrient dynamic of A. philoxeroides. The increase of water level significantly reduced the decomposition rate and nutrient release of litter in the nutrient control treatment, whereas no clear relationship was observed in the nutrient enrichment treatment, indicating that the effect of water level change on litter decomposition might be affected by nutrient enrichment. At the late stage of decomposition, the increase of phosphorus (P) concentration and the decrease of the ratio of carbon to P suggested that the decomposition of invasive plant litter was limited by P. Our results suggest that controlling P enrichment in water bodies is essential for the management of invasive plant and carbon sequestration of wetlands. In addition, the new index we proposed could provide a basis for quantifying the impact of invasive plant litter decomposition on carbon cycle in wetlands.

Phytoplankton responses to solar UVR and its combination with nutrient enrichment in a plateau oligotrophic Lake Fuxian: a mesocosm experiment.

Multiple stressors associated with global change are influencing the phytoplankton taxonomic composition and biomass in plateau lakes, such as higher levels of ultraviolet radiation (UVR, 280-400nm) and the risk of eutrophication. Although the restrictive effects of high UVR on phytoplankton are generally recognized, the effects will be impacted by the UVR levels of seasonal changes and the nutrient status. In this study, in situ phytoplankton productivity was measured seasonally under full solar radiationand only visible light (photosynthetically active radiation, PAR) conditionsin the plateau oligotrophic Lake Fuxian, Southwest China. To determine the single effects of UVR and nutrient enrichment and their combined impact on phytoplankton communities, a mesocosm experiment (1month) was conducted outdoors during late summer (rainy season). The interactive experiment was designed with two radiation treatments (UVR + PAR and PAR) and four nutrient treatments (raw water, nitrogen addition, phosphorus addition, nitrogen and phosphorus addition). Our results suggested that the response of phytoplankton to UVR depended largely on nutrient availability, which was embodied in the no significant responses of phytoplankton productivity and total biomass to UVR in the oligotrophic raw water from Lake Fuxian, but the positive responses of total biomass to UVR in the nutrient enrichment treatment. Furthermore, the response of phytoplankton to UVR was also taxa dependent. Chlorophytes and dinoflagellates showed positive response to UVR, whereas chrysophytes were quite sensitive to UVR; diatoms had no significant response to UVR under all nutrient conditions, while the response of cyanobacteria to UVR was highly dependent on nutrient status. This study highlights that the increase in nutrient concentrations in oligotrophic lake not only directly promotes the growth of phytoplankton but also combines with low UVR during summer to benefit the growth of eutrophic taxa based on the non-negative effect of UVR on total phytoplankton biomass and the positive effect on eutrophic taxa.

Impact of nutrient availability on the trophic strategies of the planktonic protist communities in a disturbed Mediterranean coastal lagoon

The impact of changes in nitrogen (N) and phosphorus (P) availability on the trophic strategies of planktonic protists was evaluated in a disturbed Mediterranean lagoon (Biguglia lagoon, France) using short-term bioassays. Natural communities were collected in three periods, i.e., autumn, spring and summer, to address the influence of the different environmental conditions. The responses of autotrophic plankton communities to experimentally induced N and/or P limitations were assessed as changes in chlorophyll a (Chl a) concentrations and in the abundances of potentially mixotrophic protists taxa. We observed blooms (> 105 cells l−1) of nanoflagellates in autumn, and of phycocyanin-rich picocyanobacteria in summer. Communities showed a co-limitation by N and P at the three sampling periods, despite high N:P ratios in autumn and spring. The high abundances of potentially mixotrophic dinoflagellates during these periods suggest the involvement of alternative trophic pathways for their maintenance in the lagoon. After bioassay incubations using different nutrient enrichment treatments, we often observed reduced abundances of mixotrophic protists containing Chl a with a concomitant increased abundance of protists without Chl a. This indicates a loss of chloroplasts and photoautotrophic abilities in protists cells, possibly reflecting a shift towards heterotrophy that could be sustained by phagotrophy.

Effects of two wood-based biochars on the fate of added fertilizer nitrogen\u2014a 15N tracing study

A 15N tracing pot experiment was conducted using two types of wood-based biochars: a regular biochar and a Kon-Tiki-produced nutrient-enriched biochar, at two application rates (1% and 5% (w/w)), in addition to a fertilizer only and a control treatment. Ryegrass was sown in pots, all of which except controls received 15N-labelled fertilizer as either 15NH4NO3 or NH415NO3. We quantified the effect of biochar application on soil N2O emissions, as well as the fate of fertilizer-derived ammonium (NH4+) and nitrate (NO3−) in terms of their leaching from the soil, uptake into plant biomass, and recovery in the soil. We found that application of biochars reduced soil mineral N leaching and N2O emissions. Similarly, the higher biochar application rate of 5% significantly increased aboveground ryegrass biomass yield. However, no differences in N2O emissions and ryegrass biomass yields were observed between regular and nutrient-enriched biochar treatments, although mineral N leaching tended to be lower in the nutrient-enriched biochar treatment than in the regular biochar treatment. The 15N analysis revealed that biochar application increased the plant uptake of added nitrate, but reduced the plant uptake of added ammonium compared to the fertilizer only treatment. Thus, the uptake of total N derived from added NH4NO3 fertilizer was not affected by the biochar addition, and cannot explain the increase in plant biomass in biochar treatments. Instead, the increased plant biomass at the higher biochar application rate was attributed to the enhanced uptake of N derived from soil. This suggests that the interactions between biochar and native soil organic N may be important determinants of the availability of soil N to plant growth.

Phytoplankton growth and microzooplankton grazing in the central and northern South China Sea in the spring intermonsoon season of 2017

Phytoplankton growth rates and mortality rates were experimentally examined at 21 stations during the 2017 spring intermonsoon (April to early May) in the northern and central South China Sea (SCS) using the dilution technique, with emphasis on a comparison between the northern and central SCS areas which had different environmental factors. There had been higher temperature but lower nutrients and chlorophyll a concentrations in the central SCS than those in the northern SCS. The mean rates of phytoplankton growth (μ0) and microzooplankton grazing (m) were (0.88±0.33) d−1 and (0.55±0.22) d−1 in the central SCS, and both higher than those in the northern SCS with the values of μ0 ((0.81±0.16) d−1) and m ((0.30±0.09) d−1), respectively. Phytoplankton growth and microzooplankton grazing rates were significantly coupled in both areas. The microzooplankton grazing impact (m/μ0) on phytoplankton was also higher in the central SCS (0.63±0.12) than that in the northern SCS (0.37±0.06). The microzooplankton abundance was significantly correlated with temperature in the surface. Temperature might more effectively promote the microzooplankton grazing rate than phytoplankton growth rate, which might contribute to higher m and m/μ0 in the central SCS. Compared with temperature, nutrients mainly affected the growth rate of phytoplankton. In the nutrient enrichment treatment, the phytoplankton growth rate (μn) was higher than μ0 in the central SCS, suggesting phytoplankton growth in the central SCS was nutrient limited. The ratio of μ0/μn was significantly correlated with nutrients concentrations in the both areas, indicating the limitation of nutrients was related to the concentrations of background nutrients in the study stations.

Long-term nutrient addition increases respiration and nitrous oxide emissions in a New England salt marsh.

Salt marshes may act either as greenhouse gas (GHG) sources or sinks depending on hydrological conditions, vegetation communities, and nutrient availability. In recent decades, eutrophication has emerged as a major driver of change in salt marsh ecosystems. An ongoing fertilization experiment at the Great Sippewissett Marsh (Cape Cod, USA) allows for observation of the results of over four decades of nutrient addition. Here, nutrient enrichment stimulated changes to vegetation communities that, over time, have resulted in increased elevation of the marsh platform. In this study, we measured fluxes of carbon dioxide (CO 2), methane (CH 4) and nitrous oxide (N2O) in dominant vegetation zones along elevation gradients of chronically fertilized (1,572 kg N ha−1 year−1) and unfertilized (12 kg N ha−1 year−1) experimental plots at Great Sippewissett Marsh. Flux measurements were performed using darkened chambers to focus on community respiration and excluded photosynthetic CO 2 uptake. We hypothesized that N‐replete conditions in fertilized plots would result in larger N2O emissions relative to control plots and that higher elevations caused by nutrient enrichment would support increased CO 2 and N2O and decreased CH 4 emissions due to the potential for more oxygen diffusion into sediment. Patterns of GHG emission supported our hypotheses. Fertilized plots were substantially larger sources of N2O and had higher community respiration rates relative to control plots, due to large emissions of these GHGs at higher elevations. While CH 4 emissions displayed a negative relationship with elevation, they were generally small across elevation gradients and nutrient enrichment treatments. Our results demonstrate that at decadal scales, vegetation community shifts and associated elevation changes driven by chronic eutrophication affect GHG emission from salt marshes. Results demonstrate the necessity of long‐term fertilization experiments to understand impacts of eutrophication on ecosystem function and have implications for how chronic eutrophication may impact the role that salt marshes play in sequestering C and N.

Wetland Plant Community Responses to the Interactive Effects of Simulated Nutrient and Sediment Loading: Implications for Coastal Restoration Using Mississippi River Diversions

The Mississippi River Delta Complex (MRDC) has experienced extensive wetland loss in the last century due, in part, to flood control levees that have isolated the lower Mississippi River and its sediment resource from adjacent wetlands. Reconnecting the River to these wetlands through diversions is being used and proposed on a larger scale for the future, to reduce wetland loss rates. However, some currently operating diversions (e.g., Caernarvon and Davis Pond) have been implicated in causing negative impacts on wetland ecosystem structure and function due to increased nutrient loads in diverted Mississippi River water combined with insufficient sediment delivery. Initial assessments of these concerns were carried out in a greenhouse setting where six nutrient enrichment treatment levels (control, NO3, NH4, PO4, SO4, and Combo [NO3 + NH4 + PO4 + SO4]) were applied with and without sediment addition to 60 marsh sods from a Sagittaria lancifolia-dominated oligohaline wetland at rates simulating the Davis Pond Diversion of the Mississippi River. After 25 months, independent enrichment with N (regardless of form) and sediment was generally beneficial to wetland structure and function, while SO4 enrichment had the opposite effect, regardless of sediment addition. Simultaneous application of N and P (i.e., the Combo treatment level) ameliorated the negative impacts of SO4-loading, but the concurrent application of sediment did not, likely because the loading rate was based on a diversion that was designed to deliver water and not to maximize sediment input. Nonetheless, sediment input is critical to the sustainability of MRDC wetlands experiencing high rates of deterioration. Thus, optimizing future diversions to maximize sediment delivery, along with continued surveillance of negative nutrient effects, are recommended management decisions.

Synergy between nutrients and warming enhances methane ebullition from experimental lakes

Lakes and ponds are important natural sources of the potent greenhouse gas methane (CH4), with small shallow waters identified as particular hotspots1,2. Ebullition (bubbles) of CH4 makes up a large proportion of total CH4 flux3,4. However, difficulty measuring such episodic events 5 makes prediction of how ebullition responds to nutrient enrichment and rising temperatures challenging. Here, the world’s longest running, mesocosm-based, shallow lake climate change experiment was used to investigate how the combination of warming and eutrophication (that is, nutrient enrichment) affects CH4 ebullition. Eutrophication without heating increased the relative contribution of ebullition from 51% to 75%. More strikingly the combination of nutrient enrichment and experimental warming treatments of +2–3 °C and +4–5 °C had a synergistic effect, increasing mean annual ebullition by at least 1900 mg CH4-C m−2 yr−1. In contrast, diffusive flux showed no response to eutrophication and only a small increase at higher temperatures (average 63 mg CH4–C m−2 yr−1). As shallow lakes are the most common lake type globally, abundant in highly climate sensitive regions 6 and most vulnerable to eutrophication, these results suggest their current and future contributions to atmospheric CH4 concentrations may be significantly underestimated. The combination of nutrient enrichment and warming has a synergistic effect on rates of methane ebullition from experimental lakes. This suggests methane emissions from shallow lakes may be significantly underestimated.

The contrasting effects of nutrient enrichment on growth, biomass allocation and decomposition of plant tissue in coastal wetlands

Eutrophication of coastal waters can have consequences for the growth, function and soil processes of coastal wetlands. Our aims were to assess how nutrient enrichment affects growth, biomass allocation and decomposition of plant tissues of a common and widespread mangrove, Avicennia marina, and how eutrophication drives changes in below-ground carbon sequestration. We assessed this through the measurement of above- and belowground growth and decomposition rates of plants and plant tissue in unenriched or nutrient enriched treatments. Nutrient enrichment increased biomass allocation above-ground compared to below-ground in seedlings but not in fully developed, mature trees where we observed the opposite pattern. Experiments to assess root decomposition found that 40–50% of biomass was lost within six months with little change between 12 and 18 months, indicating a high potential for accumulation of organic matter over time. We estimate root-derived carbon sequestration rates of 53, 250 and 94 g C m−2 year−1 for unenriched control, N and P enriched treatments, respectively. These results show coastal eutrophication can be beneficial and detrimental to ecosystem function of coastal plants. Eutrophication stimulates root growth in fully developed trees, increasing organic matter input to soils. Our data suggests that organic matter accumulation will increase in areas with high nutrient availability where root growth is increased and rates of decomposition are low.

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland, Alaska.

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient-enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer-resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom-up (nutrient limitation) and from the top-down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.

An experimental study on the effects of nutrient enrichment on organic carbon persistence in the western Pacific oligotrophic gyre

Abstract. Carbon sequestration in the ocean is of great concern with respect to the mitigation of global warming. How to hold the fixed organic carbon in the presence of tremendous numbers of heterotrophic microorganisms in marine environments is the central issue. We previously hypothesized that excessive nutrients would ultimately decrease the storage of organic carbon in marine environments. To test this, a series of in situ nutrient enrichment incubation experiments were conducted at a site (17.59° N, 127.00° E) within the western Pacific oligotrophic gyre. Five treatments were employed: glucose (Glu), algal exudation organic material (EOM), nitrate (N) and phosphate (P), N and P in combination with glucose and a control with no added nutrients. The results showed that the dissolved organic carbon consumption rates and bacterial community specific growth rates were enhanced by inorganic nutrient enrichment treatments during the initial 48 h incubation. At the end of 14 days of incubation, about one-third (average 3.3 μmol C kg−1) more organic carbon was respired in the glucose-enriched incubation with the addition of inorganic nutrients compared to that without. In contrast, when nutrients were limiting, glucose could not be efficiently used by the bacteria and thus it remained in the environment. These results suggest that repletion of inorganic nutrients could facilitate microbial consumption of organic carbon and thus has a significant impact on carbon cycling in the environment.

The response of amphibian larvae to exposure to a glyphosate-based herbicide (Roundup WeatherMax) and nutrient enrichment in an ecosystem experiment

Herbicides and fertilizers are widely used throughout the world and pose a threat to aquatic ecosystems. Using a replicated, whole ecosystem experiment in which 24 small wetlands were split in half with an impermeable barrier we tested whether exposure to a glyphosate-based herbicide, Roundup WeatherMax™, alone or in combination with nutrient enrichment has an effect on the survival, growth or development of amphibians. The herbicide was applied at one of two concentrations (low=210μga.e./L, high=2880μga.e./L) alone and in combination with nutrient enrichment to one side of wetlands and the other was left as an untreated control. Each treatment was replicated with six wetlands, and the experiment was repeated over two years. In the high glyphosate and nutrient enrichment treatment the survival of wood frog (Lithobates sylvaticus) larvae was lower in enclosures placed in situ on the treated sides than the control sides of wetlands. However, these results were not replicated in the second year of study and they were not observed in free swimming wood frog larvae in the wetlands. In all treatments, wood frog larvae on the treated sides of wetlands were slightly larger (<10%) than those on the control side, but no effect on development was observed. The most dramatic finding was that the abundance of green frog larvae (Lithobates clamitans) was higher on the treated sides than the control sides of wetlands in the herbicide and nutrient treatments during the second year of the study. The results observed in this field study indicate that caution is necessary when extrapolating results from artificial systems to predict effects in natural systems. In this experiment, the lack of toxicity to amphibian larvae was probably due to the fact the pH of the wetlands was relatively low and the presence of sediments and organic surfaces which would have mitigated the exposure duration.

Modification of a seagrass community by benthic jellyfish blooms and nutrient enrichment

Anthropogenic activities are increasing the number and intensity of disturbances, often acting in concert, in ecosystems across the globe. One result of human activities in many marine ecosystems is an increased abundance of jellyfish (jellyfish blooms), which have garnered recent attention for their detrimental ecological and economic impacts. We conducted a field experiment to determine effects of proliferations of benthic jellyfish, Cassiopea spp., and another common disturbance, anthropogenic nutrient enrichment (via fertilizer additions), on a shallow seagrass community in Abaco, Bahamas. Results suggested a reduction in seagrass abundance and habitat complexity in both jellyfish and nutrient enrichment treatments. Jellyfish additions were associated with reduced faunal densities; nutrient enrichment drove shifts in faunal community composition. Grazing frequency was substantially higher in nutrient-enriched plots, and lower in plots with jellyfish alone or jellyfish combined with nutrients, suggesting that jellyfish may act as a deterrent to grazers. These findings highlight the inherent complexities in predicting ecological changes within shallow seagrass ecosystems.

Distinct responses of Gulf of Mexico phytoplankton communities to crude oil and the dispersant corexit® Ec9500A under different nutrient regimes

This study examines the potential effects of exposure to South Louisiana sweet crude oil (LSC), Corexit(®) EC9500A, and dispersed oil on enclosed phytoplankton communities under different nutrient regimes. Three distinct microcosm experiments were conducted for 10days to assess changes to the structure of natural communities from the Gulf of Mexico as quantified by temporal changes in the biomasses of different phytoplankton groups. Concentration of NO3, Si and PO4 were 0.83, 0.99 and 0.09μM for the unenriched treatments and 14.07, 13.01 and 0.94μM for the enriched treatments, respectively. Overall, the contaminants LSC and Corexit(®) EC9500A led to a decrease in the number of sensitive species and an increase in more resistant species. Phytoplankton communities showed more sensitivity to LSC under nutrient-limited conditions. The addition of nutrients to initially nutrient-limited treatments lessened the inhibitory effect of LSC in the short term. Centric diatoms benefited most from this enrichment, but pennate diatoms demonstrated considerably greater tolerance to crude oil at low crude oil concentrations in nutrient-enriched treatments. Dinoflagellates showed relatively higher tolerance in nutrient-limited treatments and high crude oil concentrations. Corexit(®) EC9500A inputs significantly increased the toxicity of crude oil. Corexit(®) EC9500A alone had a highly inhibitory effect at 63ppm on phytoplankton communities. This study highlights the fact that different nutrient regimes play a major role in determining the shifts of the phytoplankton community in response to exposure to different concentrations of crude oil and dispersant. Determination of the functional equivalence of shifted phytoplankton groups could complement our research and allow for more pertinent extrapolation to real world conditions.