Nitrogen subsidies to pelagic food webs through profundal methane-oxidising bacteria in oligotrophic fresh water

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The significance of partial migration for food web and ecosystem dynamics

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.     

Nutrient Enrichment and Planktonic Biomass Ratios in Lakes

Phosphorus (P) to chlorophyll ratios and zooplankton–phytoplankton (Z:P) biomass ratios were assessed in 400 temperate lakes over a gradient of phosphorus (P) and with different fish communities. Most of the lakes in this survey were oligotrophic, with a median total P of 7.3 μg P L⁻¹. Thus, the survey provided information on food web effects during the early phase of eutrophication. There was no tendency toward a reduced yield of autotrophs per unit of P over the gradient covered in this survey. The zooplankton yield per unit of P or chlorophyll a decreased slightly with increased nutrient concentrations, and Z:P biomass ratios decreased with fish community classes, reflecting increased fish predation pressure. However, the variability in biomass ratios within a given range of P and fish class was some 100 times higher than the difference over the gradients. This finding suggests that lake-specific properties, community composition, and food quality are by far the most important determinants of biomass ratios and probably also trophic efficiency in lakes; it further suggests that these factors are superimposed on the general effect of eutrophication, at least up to 30 μg P L⁻¹.     

Zooplankton biomass dynamics in oligotrophic versus eutrophic conditions: a test of the PEG model

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Roles of food web and heterotrophic microbial processes in upper ocean biogeochemistry: Global patterns and processes

The growth and dynamics of plankton in the ocean vary with natural cycles, global climate change and the long-term evolution of ecosystems. The ocean is a large reservoir for CO₂ and the food webs in the upper ocean play critical roles in regulating the global carbon cycle, changes in atmospheric CO₂ and associated global warming. Microheterotrophs are a key component of the upper ocean food webs. Here, we report on the results of an analysis of the distribution of bacteria and related properties in the World Ocean. We found that, for the data set as a whole, there is a significant latitudinal gradient in all field-measured and computed bacterial properties, except growth rate. Gradients were, for the most part, driven by an equator-ward increase in the Southern Hemisphere. The biomass, rates of production and respiration and dissolved organic carbon concentrations were significantly higher in the Northern than the Southern hemispheres. In contrast, growth rates were the same in the two hemispheres. We conclude that the lower biomass and production in the Southern Hemisphere reflects greater top-down control by microbial grazers, which would be due to a lower abundance or activity of omnivorous zooplankton in the Southern than Northern Hemispheres. These large spatial differences in dynamics, structure and activity of the bacterial community and the microbial food web will be reflected in different patterns of carbon cycling, export and air–sea exchange of CO₂ and the potential ability of the ocean to sequester carbon.     

Carbon:phosphorus stoichiometry and food chain production

Incident light was manipulated in large plankton towers containing algae, microbes, and herbivores. Paradoxically, food chain production was lower with greater light energy input. This apparent paradox is resolved by recognizing stoichiometric constraints to food chain production. At high light, elevated algal biomass was achieved mainly by increases in cellular carbon. Consumers have a high phosphorus demand for growth, and thus a large excess of carbon inhibited, rather than stimulated, their growth. These experiments may help us predict the consequences of anthropogenic perturbations in nutrients, carbon, and solar energy. They also may help us to understand the wide range of consumer biomass and production at a given level of primary productivity in ecosystems.     

Effect of young-of-the-year walleye (Percidae: Stizostedion vitreum) on plankton dynamics and water quality in ponds

To understand the impact of young-of-the-year (YOY) fish on food web dynamics and water quality, we stocked larval walleye (9 mm TL) (Stizostedion vitreum) in six experimental ponds using two fish densities (10 and 50 fish m^–3) with three replicates. At high fish density, the average abundances of cladocerans and copepods and the Secchi depth were lower whereas abundances of rotifers and algae, gross primary productivity (GPP), pH and total phosphorus concentration were higher than at low fish density. Fish impact on bacterial abundance, dissolved oxygen, nitrogen and phosphorus concentrations, however, was not significant. The within treatment measurements of all variables except GPP were significantly different over time. Our results indicate that YOY walleye predation at high density can affect plankton community by reducing large zooplankton biomass and water clarity, and increasing phytoplankton abundance. The impact of YOY piscivorous fish on plankton should be considered when biomanipulation is applied for improvement of water quality.     

Pelagic food web interactions among benthic invertebrates and trout in mountain lakes

1. Benthic chironomid larvae and the amphipod Gammarus lacustris have been observed in the pelagic habitats of many mountain lakes. The main goal of this study was to determine if chironomid larvae and gammarids potentially affect predator–prey and nutrient dynamics in pelagic food webs of mountain lakes. 2. Eighty‐six mountain lakes were surveyed in Alberta and eastern British Columbia during the years 1965–1984, 1991–2004 and 2005–2007. Pelagic chironomid larvae were found in 86% of these lakes, and pelagic gammarids were found in 29% of lakes. Densities of pelagic chironomid larvae were 92% lower in lakes with pelagic gammarids and 76% lower in lakes with trout (P < 0.05). Intraguild predation of trout on gammarids appeared to reduce predation pressure on chironomid larvae. Gammarids consumed in vitro about 1 chironomid per gammarid per day or about 20% of their body mass in chironomid biomass per day. 3. Concentrations of total dissolved P and N, particulate C, and chlorophyll‐a increased with increasing densities of pelagic gammarids and chironomid larvae in situ (R² = 0.14 ± 0.19 SD, P < 0.1) and in vitro (P < 0.001). 4. Our findings suggest that gammarids and chironomid larvae are linked as predators and prey in pelagic food webs, possibly stimulating phytoplankton abundance via nutrient release.     

Planktonic food web in marine mesocosms in the Eastern Mediterranean: bottom-up or top-down regulation?

A mesocosm experiment was conducted in order to studythe structure of the planktonic food web. The dynamicsof pico-, nano- and microplankton populations werefollowed during 40 days in four large (40 m³)enclosures. In three tanks a gradient of addednutrients (nitrogen and phosphorus) was applied, whilea fourth tank was used as a control. On day 14, thetop predator (sea bream Sparus aurata larvae)was introduced into the tanks and part of the watercolumn in each tank was isolated in a plastic bagwithout fish larvae, to act as a control forpredation. Physical parameters, chlorophyll aand nutrient concentrations, as well as planktonconcentrations were monitored. A diatom bloom wasobserved in all four tanks, in the first phase endingwith silicate depletion. Flagellate and dinoflagellateabundance subsequently increased, these organismsbeing limited by zooplankton grazing. The zooplanktonpopulations were controlled by both resources (mostlyflagellates) and predation (by fish larvae) asindicated by the results of the control experiments. This revised version was published online in July 2006 with corrections to the Cover Date.     

Growth and survival of Daphnia in epilimnetic and metalimnetic water from oligotrophic lakes: the effects of food and temperature

SUMMARY 1. In oligotrophic lakes, phytoplankton and bacteria growing in the deep chlorophyll maximum in the cool metalimnion of lakes often dominate biomass and production, but the importance of this source of food for zooplankton is unknown.
2. During much of the day, Daphnia rosea in two mountain lakes inhabited deep chlorophyll layers where food availability was at least equal to that in the epilimnion.
3. To determine the importance of the two strata (epilimnion and metalimnion) for Daphnia , we used a cross-classified factorial experiment to measure how epilimnetic and metalimnetic food and temperature (10 and 16 °C) influenced survival, growth and reproduction.
4. Daphnia survived and grew better when fed seston from the epilimnion of one lake, although chlorophyll, particulate nitrogen and particulate carbon were 2–2.5 times greater in the metalimnion.
5. Temperature had no significant influence on Daphnia survival or growth. Similar results were obtained with food from the second lake, with Daphnia surviving and reproducing better when provided with epilimnetic, rather than metalimnetic food, although the quantities of chlorophyll and carbon in the two strata were similar.
6. Food quality, rather than quantity or temperature, appeared to be the most important determinant influencing survival, growth and reproduction, and the greater food quantity in the metalimnia was not used effectively by the Daphnia .     

Zooplankton and bacteria contribution to phosphorus and nitrogen internal cycling in a tropical and eutrophic reservoir: Pampulha Lake,Brazil

Nutrient regeneration and respiration rates of natural zooplankton from a tropical reservoir were experimentally measured. Excretion rates of ammonia (E_a), orthophosphate (E_p) and community respiration rates (R) were estimated considering the variations in the concentrations of ammonia, orthophosphate and dissolved oxygen between control and experimental units. The ranges obtained for these rates from the 2 h assays were E_a = 1.95–4.95 μg N-NH₄ · mg · DW⁻¹ · h⁻¹; E_p = 0.12–0.76 μg P-PO₄ mg DW⁻¹ · h⁻¹. Respiratory rates were quite constant (R = 0.01–0.02 mg O₂ · mg DW⁻¹ · h⁻¹). The uptake of nutrients due to bacteria can affect the experimental determination of excretion rates of zooplankton. Orthophosphate release increased from 0.28 to 0.82 μg P-PO₄ · mg DW⁻¹ · h⁻¹ when bacterial activity was depleted by antibiotic addition in experimental vessels (Exp IV). This demonstrates that free living bacteria are able to consume promptly most phosphorus excreted by zooplankton. Ammonia excretion rates were lower in experimental units containing antibiotics. Lower excretion rates were also obtained with longer exposure times and higher biomass levels in the experimental units. Finally, this study also showed that zooplankton excretion can affect significantly turn over rates of total phosphorus in Pampulha Reservoir. In some periods, specially during the dry season when zooplankton biomass was very high, phosphorus release by zooplankton, during one single day, can be as high as 40% of the total phosphorus content in lake water (Turn over time = 2.5 days).     

Phosphorus cycling and partitioning in an oligotrophic Everglades wetland ecosystem: a radioisotope tracing study

1. Our goal was to quantify short‐term phosphorus (P) partitioning and identify the ecosystem components important to P cycling in wetland ecosystems. To do this, we added P radiotracer to oligotrophic, P‐limited Everglades marshes. ³²PO₄ was added to the water column in six 1‐m² enclosed mesocosms located in long‐hydroperiod marshes of Shark River Slough, Everglades National Park. Ecosystem components were then repeatedly sampled over 18 days. 2. Water column particulates (>0.45 μm) incorporated radiotracer within the first minute after dosing and stored 95–99% of total water column ³²P activity throughout the study. Soluble (<0.45 μm) ³²P in the water column, in contrast, was always <5% of the ³²P in surface water. Periphyton, both floating and attached to emergent macrophytes, had the highest specific activity of ³²P (Bq g^?131P) among the different ecosystem components. Fish and aquatic macroinvertebrates also had high affinity for P, whereas emergent macrophytes, soil and flocculent detrital organic matter (floc) had the lowest specific activities of radiotracer. 3. Within the calcareous, floating periphyton mats, 81% of the initial ³²P uptake was associated with Ca, but most of this ³²P entered and remained within the organic pool (Ca‐associated = 14% of total) after 1 day. In the floc layer, ³²P rapidly entered the microbial pool and the labile fraction was negligible for most of the study. 4. Budgeting of the radiotracer indicated that ³²P moved from particulates in the water column to periphyton and floc and then to the floc and soil over the course of the 18 day incubations. Floc (35% of total) and soil (27%) dominated ³²P storage after 18 days, with floating periphyton (12%) and surface water (10%) holding smaller proportions of total ecosystem ³²P. 5. To summarise, oligotrophic Everglades marshes exhibited rapid uptake and retention of labile ³²P. Components dominated by microbes appear to control short‐term P cycling in this oligotrophic ecosystem.     

The St. Lawrence River in winter: population structure,biomass, and pattern of its primary and secondary food web components

Plankton and benthic invertebrate populations were examined along a 160 km section of the St. Lawrence River between Lake Ontario and Waddington, New York during a period of ice cover. Algal biomass was low in winter with diatoms and cryptomonads the most prominent forms. Zooplankton biomass was about one-tenth the algal standing crop and cyclopoid copepods predominated. Phytoplankton and zooplankton biomass declined downriver providing evidence that most plankton was imported from Lake Ontario. Benthic invertebrate standing crop was greatest near Lake Ontario where molluscs were abundant. The downriver macroinvertebrate community shifted to coarse particle feeders, namely annelids and chironomids. Beds of accumulated detrital material derived primarily from dieback of macrophytes were particularly rich in benthic invertebrates.     

Integrating classical and microbial food web concepts: evolving views from the open-ocean tropical Pacific

Over the past half-century, and particularly the last two decades, new paradigms, perspectives and technological capabilities have greatly advanced our understanding of open-ocean pelagic ecosystems. Major new insights have come from the microbial loop concept and related discoveries, the iron limitation hypothesis and ocean time series. Focusing mainly on the tropical and subtropical Pacific Ocean, I review the influences of these new perspectives on classical views of food web complexity, phytoplankton regulation and diversity, and temporal dynamics. ``Microorganisms (bacteria, fungi, protozoa) are responsible for about 95% of the CO ₂ evolved into the atmosphere, while animals contribute about 5%. These estimates are based on figures from terrestrial environments, but there is every reason to believe that microorganisms are relatively as important in the oceans'. (Lecture Note Handouts, OCN 434, Winter, 1971)     

Intra‐lake stable isotope ratio variation in selected fish species and their possible carbon sources in Lake Kyoga (Uganda): implications for aquatic food web studies

The stable isotopes of nitrogen (δ¹⁵N) and carbon (δ¹³C) provide powerful tools for quantifying trophic relationships and carbon flow to consumers in food webs; however, the isotopic signatures of organisms vary within a lake. Assessment of carbon and nitrogen isotopic signatures in a suite of plants, invertebrates, and fishes in Lake Kyoga, indicated significant variation between two sites for δ¹³C (paired t = 6.305; df = 14, P < 0.001 and δ¹⁵N paired t = 1.292; df = 14; P < 0.05). The fish fauna in Bukungu was generally more ¹³C enriched (mean δ¹³C = –16.37 ± 1.64‰) than in Iyingo (mean δ¹³C = –20.80 ± 2.41‰) but more δ¹⁵N depleted (mean δ¹⁵N = 5.57 ± 0.71‰) than in Iyingo (mean δ¹⁵N = 6.92 ± 0.83‰). The simultaneous shifts in phytoplankton and consumer signatures confirmed phytoplankton as the major source of carbon for the food chain leading to fish. Limited sampling coverage within lakes may affect lake wide stable isotope signatures, and the same error is transferred into trophic position estimation. Consideration of potential intra‐lake spatial variability in isotope ratios and size is essential in evaluating the spatial and trophic structure of fish assemblages.     

Nutrient dynamics in shallow lakes: effects of changes in loading and role of sediment-water interactions

The transport and cycling of nutrients through the various pools in water, soil and sediment is controlling the long term and short term productivity of water bodies. An understanding of the size of these pools and the fluxes between them is essential for the assessment of the usefulness of management measures resulting in reduced external input and the anticipated resilience of the system towards changes in trophic character. Large pools, such as phosphorus in surficial sediments and nitrate in groundwater have a potential for prolonged stimulation of productivity. Diffuse sources, fluxes towards sinks, competition between biota and adsorbents for sparse nutrients, feedback mechanisms, non-linearities and shifts among prevailing processes are discussed.     

Using food network unfolding to evaluate food–web complexity in terms of biodiversity: theory and applications

Food–web complexity often hinders disentangling functionally relevant aspects of food–web structure and its relationships to biodiversity. Here, we present a theoretical framework to evaluate food–web complexity in terms of biodiversity. Food network unfolding is a theoretical method to transform a complex food web into a linear food chain based on ecosystem processes. Based on this method, we can define three biodiversity indices, horizontal diversity (D_H), vertical diversity (D_V) and range diversity (D_R), which are associated with the species diversity within each trophic level, diversity of trophic levels, and diversity in resource use, respectively. These indices are related to Shannon's diversity index (H′), where H′ = D_H + D_V ? D_R. Application of the framework to three riverine macroinvertebrate communities revealed that D indices, calculated from biomass and stable isotope features, captured well the anthropogenic, seasonal, or other within‐site changes in food–web structures that could not be captured with H′ alone.     

Tree Harvest in an Experimental Sand Ecosystem: Plant Effects on Nutrient Dynamics and Solute Generation

The hydrochemical signatures of forested ecosystems are known to be determined by a time-variant combination of physical-hydrologic, geochemical, and biologic processes. We studied subsurface potassium (K), calcium (Ca), and nitrate (NO₃) in an experimental red -pine mesocosm to determine how trees affect the behavior of these nutrients in soil water, both during growth and after a harvest disturbance. Solution chemistry was monitored for 2 years at the end of a 15-year period of tree growth, and then for 3 more years after harvest and removal of aboveground biomass. Concentrations were characterized by three distinct temporal patterns that we ascribe to changes in solute generation mechanisms. Prior to harvest, K soil-water concentrations were relatively uniform with depth, whereas Ca soil-water concentrations doubled with depth. Nitrate concentrations were below detection in soil water and discharge (drainage) water. Plant uptake and water/nutrient cycling exerted strong control during this interval. During the 1st year after harvest, K concentrations tripled in shallow soil water, relative to preharvest levels, and showed a strong seasonal peak in discharge that mimicked soil temperature. Summer soil temperatures and annual water flux also increased. Decomposition of labile litter, with complete nitrogen (N) immobilization, characterized this interval. In the third interval (years 2 and 3 after harvest), decomposition shifted from N to carbon (C) limitation, and Ca and NO₃ concentrations in discharge spiked to nearly 200 and 400 μM, respectively. Relatively stable ionic strength and carbonate chemistry in discharge, throughout the study period, indicate that carbonic-acid weathering was sustained by belowground decomposition long after the harvest. This stable chemical weathering regime, along with the persistence of N limitation for a long period after disturbance, may be characteristic of early-phase primary-successional systems.     

Nutrient content in Quercus ilex canopies: Seasonal and spatial variation within a catchment

Spatial and temporal changes in canopy nutrient content were studied in 1988 and 1989 in a Mediterranean Quercus ilex ssp. ilex L. forest in north-eastern Spain. Sampling was conducted in parallel at two sites which represent endpoints along a slope gradient within a small catchment (ridge top at 975 m and valley bottom at 700 m). Deeper soils resulted in significantly higher N and P concentrations, and N content on a leaf area basis at the valley bottom site. In contrast, K concentration in leaves was significantly higher at the ridge top site, where soil K concentration was also higher. At both sites, N and P content on a leaf area basis was highest at the top of the canopy, where leaf area is highest. N resorption efficiency decreased from top to bottom of the canopy. Results suggested a minor role of shaded leaves as nutrient storage sites. Lower P resorption efficiency was found at the ridge top site. Seasonal changes of P and N concentration on a leaf area basis suggest P replenishment, and to a lesser degree N, during periods of lower growth activity due to low temperatures, but coinciding with higher water availability (autumn-early spring period). Thus, N and P resorption from the remaining foliage in the canopy took place, and to a larger degree at the valley bottom site, coinciding with a slightly higher leaf area index and productivity at this site.     

Climate change and invasion by intracontinental range-expanding exotic plants: the role of biotic interactions

Background and Aims

In this Botanical Briefing we describe how the interactions between plants and their biotic environment can change during range-expansion within a continent and how this may influence plant invasiveness.

Scope

We address how mechanisms explaining intercontinental plant invasions by exotics (such as release from enemies) may also apply to climate-warming-induced range-expanding exotics within the same continent. We focus on above-ground and below-ground interactions of plants, enemies and symbionts, on plant defences, and on nutrient cycling.

Conclusions

Range-expansion by plants may result in above-ground and below-ground enemy release. This enemy release can be due to the higher dispersal capacity of plants than of natural enemies. Moreover, lower-latitudinal plants can have higher defence levels than plants from temperate regions, making them better defended against herbivory. In a world that contains fewer enemies, exotic plants will experience less selection pressure to maintain high levels of defensive secondary metabolites. Range-expanders potentially affect ecosystem processes, such as nutrient cycling. These features are quite comparable with what is known of intercontinental invasive exotic plants. However, intracontinental range-expanding plants will have ongoing gene-flow between the newly established populations and the populations in the native range. This is a major difference from intercontinental invasive exotic plants, which become more severely disconnected from their source populations.