Responses of seagrass communities to fertilization along a gradient of relative availability of nitrogen and phosphorus in a carbonate environment

Patterns of relative nutrient availability in south Florida suggest spatial differences regarding the importance of nitrogen (N) and phosphorus (P) to benthic primary producers. We did a 14-month in situ fertilization experiment to test predictions of N and P limitation in the subtropical nearshore marine waters of the upper Florida Keys. Six sites were divided into two groups (nearshore, offshore) representing the endpoints of an N: P stoichiometric gradient. Twenty-four plots were established at each site with six replicates of each treatment (1N, 1P, 1N1P, control), for a total of 144 experimental plots. The responses of benthic communities to N and P enrichment varied appreciably between nearshore and offshore habitats. Offshore seagrass beds were strongly limited by nitrogen, and nearshore beds were affected by nitrogen and phosphorus. Nutrient addition at offshore sites increased the length and aboveground standing crop of the two seagrasses, Thalassia testudinum and Syringodium filiforme, and growth rates of T. testudinum. Nutrient addition at nearshore sites increased the relative abundance of macroalgae, epiphytes, and sediment microalgae. N limitation of seagrass in this carbonate system was clearly demonstrated. However, added phosphorus was retained in the system more effectively than N, suggesting that phosphorus might have important long-term effects on these benthic communities. The observed species-specific responses to nutrient enrichment underscores the need to monitor all primary producers when addressing questions of nutrient limitation and eutrophication in seagrass communities.

Carbon (d13c) and Nitrogen (d15n) Isotopic Discrimination in Mangroves in Florida Coastal Everglades as a Function of Environmental Stress

Isotope signatures of mangrove leaves can vary depending on discrimination associated with plant response to environmental stressors defined by gra­dients of resources (such as water and nutrient limitation) and regulators (such as salinity and sul­fide toxicity). We tested the variability of man­grove isotopic signatures (d13C and d15N) across a stress gradient in south Florida, using green leaves from four mangrove species collected at six sites. Mangroves across the landscape studied are stressed by resource and regulator gradients repre­sented by limited phosphorus concentrations com­bined with high sulfide concentrations, respec­tively. Foliar d13C ratios exhibited a range from ­ 24.6 to –32.7‰, and multiple regression analysis showed that 46% of the variability in mangrove d13C composition could be explained by the differ­ences in dissolved inorganic nitrogen, soluble reac­tive phosphorus, and sulfide porewater concentra­tions. 15N discrimination in mangrove species ranged from –0.1 to 7.7‰, and porewater N, salin­ity, and leaf N:Pa ratios accounted for 41% of this variability in mangrove leaves. The increase in soil P availability reduced 15N discrimination due to higher N demand. Scrub mangroves (<1.5 m tall) are more water-use efficient, as indicated by higher d13C; and have greater nutrient use efficiency ratios of P than do tall mangroves (5 to 10 m tall) existing in sites with greater soil P concentrations. The high variability of mangrove d13C and d15N across these resource and regulator gradients could be a con­founding factor obscuring the linkages between mangrove wetlands and estuarine food webs. These results support the hypothesis that landscape fac­tors may control mangrove structure and function, so that nutrient biogeochemistry and mangrove-based food webs in adjacent estuaries should ac­count for watershed-specific organic inputs.

The effects of manipulation of sedimentary iron and organic matter on sediment biogeochemistry and seagrasses in a subtropical carbonate environment

The microbial metabolism of organic matter (OM) in seagrass beds can create sulfidic conditions detrimental to seagrass growth; iron (Fe) potentially has ameliorating effects through titration of the sulfides and the precipitation of iron-sulfide minerals into the sediment. In this study, the biogeochemical effects of Fe availability and its interplay with sulfur and OM on sulfide toxicity, phosphorous (P) availability, seagrass growth and community structure were tested. The availability of Fe and OM was manipulated in a 2 × 2 factorial experiment arranged in a Latin square, with four replicates per treatment. The treatments included the addition of Fe, the addition of OM, the addition of both Fe and OM as well as no addition. The experiment was conducted in an oligotrophic, iron-deficient seagrass bed. Fe had an 84.5% retention efficiency in the sediments with the concentration of Fe increasing in the seagrass leaves over the course of the experiment. Porewater chemistry was significantly altered with a dramatic decrease in sulfide levels in Fe addition plots while sulfide levels increased in the OM addition treatments. Phosphorus increased in seagrass leaves collected in the Fe addition plots. Decreased sulfide stress was evidenced by heavier δ34S in leaves and rhizomes from plots to which Fe was added. The OM addition negatively affected seagrass growth but increased P availability; the reduced sulfide stress in Fe added plots resulted in elevated productivity. Fe availability may be an important determinant of the impact that OM has on seagrass vitality in carbonate sediments vegetated with seagrasses.

Patterns of phosphorus, nitrogen and δ15N along a peat development gradient in a coastal mire, Panama

Differentiation of limiting nutrients within small spatial scales has been observed in coastal mangrove forests, but research on other tropical peatlands suggests it is a more widespread phenomenon. In the Changuinola mire of coastal Panama, oligotrophy was hypothesized to increase along a gradient of peat development (peat doming). Nutrient and carbon concentration of leaf tissue, soil, and soil porewater were characterised over a successive sequence of plant communities along the gradient. Soil phosphorus (P) and nitrogen (N) concentrations decreased from 1200 μg P g−1 and 27 mg N g−1 to 377 μg P g−1 and 22 mg N g−1 within 2.7 km into the mire interior. These changes coincided with an increase in soil and average leaf N:P molar ratios from 52–128 and 24–41, respectively. Soil P was strongly related to leaf P and soil N:P to foliar N:P. There was a wide range in δ15N values for canopy (4.0 to −9.4‰), Campnosperma panamense (4.0 to −7.8‰) and understorey (4.8 to −3.1‰) species. Foliar δ15N values of canopy species were strongly related to soil N:P, soil P and leaf P. The depleted foliar δ15N values appeared to be an effect of both the N atmospheric source and P limitation. Here, P limitation is likely associated with ombrotrophic conditions that developed as hydrologic inputs became dominated by precipitation.

Top down control in a relatively pristine seagrass ecosystem

The loss of large-bodied herbivores and/or top predators has been associated with large-scale changes in terrestrial, freshwater, and marine ecosystems around the world. Understanding the consequences of these declines has been hampered by a lack of studies in relatively pristine systems. To fill this gap, I investigated the dynamics of the relatively pristine seagrass ecosystem of Shark Bay, Australia. I began by examining the seagrass species distributions, stoichiometry, and patterns of nutrient limitation across the whole of Shark Bay. Large areas were N-limited, P-limited, or limited by factors other than nutrients. Phosphorus-limitation was centered in areas of restricted water exchange with the ocean. Nutrient content of seagrasses varied seasonally, but the strength of seasonal responses were species-specific. Using a cafeteria-style experiment, I found that fast-growing seagrass species, which had higher nutrient content experienced higher rates of herbivory than slow-growing species that are dominant in the bay but have low nutrient content. Although removal rates correlated well with nutrient content at a broad scale, within fast-growing species removal rates were not closely tied to N or P content. Using a combination of stable isotope analysis and animal borne video, I found that green turtles (Chelonia mydas)—one of the most abundant large-bodied herbivores in Shark Bay—appear to assimilate little energy from seagrasses at the population level. There was, however, evidence of individual specialization in turtle diets with some individuals foraging largely on seagrasses and others feeding primarily on macroalgae and gelatinous macroplankton. Finally, I used exclusion cages, to examine whether predation-sensitive habitat shifts by megagrazers (green turtles, dugongs) transmitted a behavior-mediated trophic cascade (BMTC) between sharks and seagrasses. In general, data were consistent with predictions of a behavior-mediated trophic cascade. Megaherbivore impacts on seagrasses were large only in the microhabitat where megaherbivores congregate to reduce predation risk. My study highlights the importance of large herbivores in structuring seagrass communities and, more generally, suggests that roving top predators likely are important in structuring communities—and possibly ecosystems—through non-consumptive pathways.

Phytoplankton-bacteria coupling under elevated CO2 levels: a stable isotope labelling study, 2010

The potential impact of rising carbon dioxide (CO2) on carbon transfer from phytoplankton to bacteria was investigated during the 2005 PeECE III mesocosm study in Bergen, Norway. Sets of mesocosms, in which a phytoplankton bloom was induced by nutrient addition, were incubated under 1x (~350 µatm), 2x (~700 µatm), and 3x present day CO2 (~1050 µatm) initial seawater and sustained atmospheric CO2 levels for 3 weeks. 13C labelled bicarbonate was added to all mesocosms to follow the transfer of carbon from dissolved inorganic carbon (DIC) into phytoplankton and subsequently heterotrophic bacteria, and settling particles. Isotope ratios of polar-lipid-derived fatty acids (PLFA) were used to infer the biomass and production of phytoplankton and bacteria. Phytoplankton PLFA were enriched within one day after label addition, whilst it took another 3 days before bacteria showed substantial enrichment. Group-specific primary production measurements revealed that coccolithophores showed higher primary production than green algae and diatoms. Elevated CO2 had a significant positive effect on post-bloom biomass of green algae, diatoms, and bacteria. A simple model based on measured isotope ratios of phytoplankton and bacteria revealed that CO2 had no significant effect on the carbon transfer efficiency from phytoplankton to bacteria during the bloom. There was no indication of CO2 effects on enhanced settling based on isotope mixing models during the phytoplankton bloom, but this could not be determined in the post-bloom phase. Our results suggest that CO2effects are most pronounced in the post-bloom phase, under nutrient limitation.

Responses of the Emiliania huxleyi Proteome to Ocean Acidification

Ocean acidification due to rising atmospheric CO2 is expected to affect the physiology of important calcifying marine organisms, but the nature and magnitude of change is yet to be established. In coccolithophores, different species and strains display varying calcification responses to ocean acidification, but the underlying biochemical properties remain unknown. We employed an approach combining tandem mass-spectrometry with isobaric tagging (iTRAQ) and multiple database searching to identify proteins that were differentially expressed in cells of the marine coccolithophore species Emiliania huxleyi (strain NZEH) between two CO2 conditions: 395 (~current day) and ~1340 p.p.m.v. CO2. Cells exposed to the higher CO2 condition contained more cellular particulate inorganic carbon (CaCO3) and particulate organic nitrogen and carbon than those maintained in present-day conditions. These results are linked with the observation that cells grew slower under elevated CO2, indicating cell cycle disruption. Under high CO2 conditions, coccospheres were larger and cells possessed bigger coccoliths that did not show any signs of malformation compared to those from cells grown under present-day CO2 levels. No differences in calcification rate, particulate organic carbon production or cellular organic carbon: nitrogen ratios were observed. Results were not related to nutrient limitation or acclimation status of cells. At least 46 homologous protein groups from a variety of functional processes were quantified in these experiments, of which four (histones H2A, H3, H4 and a chloroplastic 30S ribosomal protein S7) showed down-regulation in all replicates exposed to high CO2, perhaps reflecting the decrease in growth rate. We present evidence of cellular stress responses but proteins associated with many key metabolic processes remained unaltered. Our results therefore suggest that this E. huxleyi strain possesses some acclimation mechanisms to tolerate future CO2 scenarios, although the observed decline in growth rate may be an overriding factor affecting the success of this ecotype in future oceans.

The effect of nitrate and phosphate availability on Emiliania huxleyi(NZEH) physiology under different CO2 scenarios

Growth and calcification of the marine coccolithophorid Emiliania huxleyi is affected by ocean acidification and macronutrients limitation and its response varies between strains. Here we investigated the physiological performance of a highly calcified E. huxleyi strain, NZEH, in a multiparametric experiment. Cells were exposed to different CO2 levels (ranging from 250 to 1314 µatm) under three nutrient conditions [nutrient replete (R), nitrate limited (-N), and phosphate limited (-P)]. We focused on calcite and organic carbon quotas and on nitrate and phosphate utilization by analyzing the activity of nitrate reductase (NRase) and alkaline phosphatase (APase), respectively. Particulate inorganic (PIC) and organic (POC) carbon quotas increased with increasing CO2 under R conditions but a different pattern was observed under nutrient limitation. The PIC:POC ratio decreased with increasing CO2 in nutrient limited cultures. Coccolith length increased with CO2 under all nutrient conditions but the coccosphere volume varied depending on the nutrient treatment. Maximum APase activity was found at 561 ?atm of CO2 (pH 7.92) in -P cultures and in R conditions, NRase activity increased linearly with CO2. These results suggest that E. huxleyi's competitive ability for nutrient uptake might be altered in future high-CO2 oceans. The combined dataset will be useful in model parameterizations of the carbon cycle and ocean acidification.

Taxonomic and Functional Diversity of Soil and Hypolithic Microbial Communities in Miers Valley, McMurdo Dry Valleys, Antarctica

The McMurdo Dry Valleys of Antarctica are an extreme polar desert. Mineral soils support subsurface microbial communities and translucent rocks support development of hypolithic communities on ventral surfaces in soil contact. Despite significant research attention, relatively little is known about taxonomic and functional diversity or their inter-relationships. Here we report a combined diversity and functional interrogation for soil and hypoliths of the Miers Valley in the McMurdo Dry Valleys of Antarctica. The study employed 16S rRNA fingerprinting and high throughput sequencing combined with the GeoChip functional microarray. The soil community was revealed as a highly diverse reservoir of bacterial diversity dominated by actinobacteria. Hypolithic communities were less diverse and dominated by cyanobacteria. Major differences in putative functionality were that soil communities displayed greater diversity in stress tolerance and recalcitrant substrate utilization pathways, whilst hypolithic communities supported greater diversity of nutrient limitation adaptation pathways. A relatively high level of functional redundancy in both soil and hypoliths may indicate adaptation of these communities to fluctuating environmental conditions.

Estimates of global sources and sinks of carbon from land cover and land use changes

The distribution of sources and sinks of carbon over the land surface is dominated by changes in land use such as deforestation, reforestation, and agricultural management. Despite, the importance of land-use change in dominating long-term net terrestrial fluxes of carbon, estimates of the annual flux are uncertain relative to other terms in the global carbon budget. The interaction of the nitrogen cycle via atmospheric N inputs and N limitation with the carbon cycle contributes to the uncertain effect of land use change on terrestrial carbon uptake. This study uses two different land use datasets to force the geographically explicit terrestrial carbon-nitrogen coupled component of the Integrated Science Assessment Model (ISAM) to examine the response of terrestrial carbon stocks to historical LCLUC (cropland, pastureland and wood harvest) while accounting for changes in N deposition, atmospheric CO2 and climate. One of the land use datasets is based on satellite data (SAGE) while the other uses population density maps (HYDE), which allows this study to investigate how global LCLUC data construction can affect model estimated emissions. The timeline chosen for this study starts before the Industrial Revolution in 1765 to the year 2000 because of the influence of rising population and economic development on regional LCLUC. Additionally, this study evaluates the impact that resulting secondary forests may have on terrestrial carbon uptake. The ISAM model simulations indicate that uncertainties in net terrestrial carbon fluxes during the 1990s are largely due to uncertainties in regional LCLUC data. Also results show that secondary forests increase the terrestrial carbon sink but secondary tropical forests carbon uptake are constrained due to nutrient limitation.

Top Down Control in a Relatively Pristine Seagrass Ecosystem

The loss of large-bodied herbivores and/or top predators has been associated with large-scale changes in terrestrial, freshwater, and marine ecosystems around the world. Understanding the consequences of these declines has been hampered by a lack of studies in relatively pristine systems. To fill this gap, I investigated the dynamics of the relatively pristine seagrass ecosystem of Shark Bay, Australia. I began by examining the seagrass species distributions, stoichiometry, and patterns of nutrient limitation across the whole of Shark Bay. Large areas were N-limited, P-limited, or limited by factors other than nutrients. Phosphorus-limitation was centered in areas of restricted water exchange with the ocean. Nutrient content of seagrasses varied seasonally, but the strength of seasonal responses were species-specific. Using a cafeteria-style experiment, I found that fast-growing seagrass species, which had higher nutrient content experienced higher rates of herbivory than slow-growing species that are dominant in the bay but have low nutrient content. Although removal rates correlated well with nutrient content at a broad scale, within fast-growing species removal rates were not closely tied to N or P content. Using a combination of stable isotope analysis and animal borne video, I found that green turtles (Chelonia mydas) – one of the most abundant large-bodied herbivores in Shark Bay – appear to assimilate little energy from seagrasses at the population level. There was, however, evidence of individual specialization in turtle diets with some individuals foraging largely on seagrasses and others feeding primarily on macroalgae and gelatinous macroplankton. Finally, I used exclusion cages, to examine whether predation-sensitive habitat shifts by megagrazers (green turtles, dugongs) transmitted a behavior-mediated trophic cascade (BMTC) between sharks and seagrasses. In general, data were consistent with predictions of a behavior-mediated trophic cascade. Megaherbivore impacts on seagrasses were large only in the microhabitat where megaherbivores congregate to reduce predation risk. My study highlights the importance of large herbivores in structuring seagrass communities and, more generally, suggests that roving top predators likely are important in structuring communities - and possibly ecosystems - through non-consumptive pathways.

Constraints of nutrient availability on primary production in two alpine tundra communities

A nutrient amendment experiment was conducted for two growing seasons in two alpine tundra communities to test the hypotheses that: (1) primary production is limited by nutrient availability, and (2) physiological and developmental constraints act to limit the responses of plants from a nutrient-poor community more than plants from a more nutrient-rich community to increases in nutrient availability. Experimental treatments consisted of N, P, and N+P amendments applied to plots in two physiognomically similar communities, dry and wet meadows. Extractable N and P from soils in nonfertilized control plots indicated that the wet meadow had higher N and P availability. Photosynthetic, nutrient uptake, and growth responses of the dominants in the two communities showed little difference in the relative capacity of these plants to respond to the nutrient additions. Aboveground production responses of the communities to the treatments indicated N availability was limiting to production in the dry meadow community while N and P availability colimited production in the wet meadow community. There was a greater production response to the N and N+P amendments in the dry meadow relative to the wet meadow, despite equivalent functional responses of the dominant species of both communities. The greater production response in the dry meadow was in part related to changes in community structure, with an increase in the proportion of graminoid and forb biomass, and a decrease in the proportion of community biomass made up by the dominant sedge Kobresia myosuroides. Species richness increased significantly in response to the N+P treatment in the dry meadow. Graminoid biomass increased significantly in the wet meadow N and N+P plots, while forb biomass decreased significantly, suggesting a competitive interaction for light. Thus, the difference in community response to nutrient amendments was not the result of functional changes at the leaf level of the dominant species, but rather was related to changes in community structure in the dry meadow, and to a shift from a nutrient to a light limitation of production in the wet meadow.

Herbivores and nutrients control grassland plant diversity via light limitation

Human alterations to nutrient cycles1, 2 and herbivore communities3, 4, 5, 6, 7 are affecting global biodiversity dramatically2. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems8, 9. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Nutrient-phytoplankton interrelationships in the Eastern Tropical Pacific Ocean

ENGLISH: Near surface nutrient distributions in the eastern tropical Pacific Ocean, using data from the EASTROPAC Expedition of 1967-68 and pre~EASTROPAC data, are described. Nutrient concentrations were maximal along the equator, in the Peru Current and its offshore extension, and in the Costa Rica Dome and westward tensions of this feature. Nutrient-poor water was found north of the equator well offshore. In this water nitrate was often undetectable (<0.1 µg-at/liter) at the surface, but phosphate and silicic acid concentrations were moderate. Enrichment experiments showed that nitrogen was the primary limiting nutrient in poor water even though large amounts of organic N were found. Half saturation constants (K s ) were determined for ammonium-supported phytoplankton growth. These data were used to calculate near-surface primary productivity values which compared favorably with 14C values. Assimilation ratio measurements indicated that algae were not extremely nitrogen-deficient. Laboratory-determined K, values for phosphate and silicic acid indicated that these nutrients were rarely limiting. In rich water, chlorophyll levels were less than expected from nutrient levels, and this anomaly may be related to limitation by nutrients other than nitrogen (N), phosphorus (P), or silicon (Si), or to grazing. SPANISH: Se describe la distribución subsuperficial de los nutrientes en el Océano Pacífico oriental tropical, empleando los datos de la Expedición EASTROPAC de 1967~68 y datos anteriores a éstos. La concentración de nutrientes fue máxima a lo largo del ecuador, en la Corriente del Perú, en su prolongación mar afuera, en el Domo de Costa Rica y en las prolongaciones occidentales de esta característica. Se encontraron aguas pobres en nutrientes al norte del ecuador y bastante mar adentro. En estas aguas el nitrato era casi imperceptible (<0.1 µg-at/litro) en la superficie, pero las concentraciones de fosfato y ácido silícico fueron moderadas. Los experimentos de enriquecimiento indicaron que el nitrógeno era el principal nutriente limitante en aguas pobres, aun cuando se encontraron grandes cantidades de nitrógeno orgánico. Se determinaron las constantes de saturación media (K s ) para el desarrollo del fitoplancton sostenido por el amonio. Estos datos se emplearon para calcular los valores de la productividad primaria cerca a la superficie que pueden compararse favorablemente con los valores del 14C. Las medidas de la proporción de asimilación indican que las algas no tenían una deficiencia extremada de nitrógeno. Los valores determinados en el laboratorio de K, para el fosfato y ácido silícico indicaron que estos nutrientes limitaron rara vez la producción. En aguas ricas, los niveles de clorofila fueron inferiores a lo esperado según los niveles nutritivos y esta anomalía puede relacionarse a la alimentación o a la escasez de otros nutrientes distintos al nitrógeno (N), fósforo (P) o silicio (Si).

Nutrient dynamics and the eutrophication of shallow lakes Kasumigaura (Japan), Donghu (PR China), and Okeechobee (USA)

We compared the nutrient dynamics of three lakes that have been heavily influenced by point and non-point source pollution and other human activities. The lakes, located in Japan (Lake Kasumigaura), People's Republic of China (Lake Donghu), and the USA (Lake Okeechobee), all are relatively large(>30 km(2)), very shallow (<4 m mean depth), and eutrophic. In all three lakes we found strong interactions among the sediments, water column, and human activities. Important processes affecting nutrient dynamics included nitrogen fixation, light limitation due to resuspended sediments, and intense grazing on algae by cultured fish. As a result of these complex interactions, simple empirical models developed to predict in-lake responses of total phosphorus and algal biomass to external nutrient loads must be used with caution. While published models may provide 'good' results, in terms of model output matching actual data, this may not be due to accurate representation of lake processes in the models. The variable nutrient dynamics that we observed among the three study lakes appears to be typical for shallow lake systems. This indicates that a greater reliance on lake-specific research may be required for effective management, and a lesser role of inter-lake generalization than is possible for deeper, dimictic lake systems. Furthermore, accurate predictions of management impacts in shallow eutrophic lakes may require the use of relatively complex deterministic modeling tools. (C) 2000 Elsevier Science Ltd. All rights reserved.