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Trophic transfer efficiency (TTE) is usually calculated as the ratio of production rates between two consecutive trophic levels. Although seemingly simple, TTE estimates from lakes are rare. In our review, we explore the processes and structures that must be understood for a proper lake TTE estimate.
We briefly discuss measurements of production rates and trophic positions and mention how ecological efficiencies, nutrients (N, P) and other compounds (fatty acids) affect energy transfer between trophic levels and hence TTE.
Furthermore, we elucidate how TTE estimates are linked with size-based approaches according to the Metabolic Theory of Ecology, and how food-web models can be applied to study TTE in lakes.
Subsequently, we explore temporal and spatial heterogeneity of production and TTE in lakes, with a particular focus on the links between benthic and pelagic habitats and between the lake and the terrestrial environment.
We provide an overview of TTE estimates from lakes found in the published literature. Finally, we present two alternative approaches to estimating TTE. First, TTE can be seen as a mechanistic quantity informing about the energy and matter flow between producer and consumer groups.
This approach is informative with respect to food-web structure, but requires enormous amounts of data. The greatest uncertainty comes from the proper consideration of basal production to estimate TTE of omnivorous organisms.
An alternative approach is estimating food-chain and food-web efficiencies, by comparing the heterotrophic production of single consumer levels or the total sum of all heterotrophic production including that of heterotrophic bacteria to the total sum of primary production.
We close the review by pointing to a few research questions that would benefit from more frequent and standardized estimates of TTE in lakes.
Methane (CH4) from aquatic ecosystems contributes to about half of total global CH4 emissions to the atmosphere. Until recently, aquatic biogenic CH4 production was exclusively attributed to methanogenic archaea living under anoxic or suboxic conditions in sediments, bottom waters, and wetlands. However, evidence for oxic CH4 production (OMP) in freshwater, brackish, and marine habitats is increasing. Possible sources were found to be driven by various planktonic organisms supporting different OMP mechanisms. Surprisingly, submerged macrophytes have been fully ignored in studies on OMP, yet they are key components of littoral zones of ponds, lakes, and coastal systems. High CH4 concentrations in these zones have been attributed to organic substrate production promoting classic methanogenesis in the absence of oxygen. Here, we review existing studies and argue that, similar to terrestrial plants and phytoplankton, macroalgae and submerged macrophytes may directly or indirectly contribute to CH4 formation in oxic waters. We propose several potential direct and indirect mechanisms: (1) direct production of CH4; (2) production of CH4 precursors and facilitation of their bacterial breakdown or chemical conversion; (3) facilitation of classic methanogenesis; and (4) facilitation of CH4 ebullition. As submerged macrophytes occur in many freshwater and marine habitats, they are important in global carbon budgets and can strongly vary in their abundance due to seasonal and boom-bust dynamics. Knowledge on their contribution to OMP is therefore essential to gain a better understanding of spatial and temporal dynamics of CH4 emissions and thus to substantially reduce current uncertainties when estimating global CH4 emissions from aquatic ecosystems.
Periphyton is a major contributor to aquatic primary production and often competes with phytoplankton and submerged macrophytes for resources. In nutrient-limited environments, mobilization of sediment nutrients by groundwater can significantly affect periphyton (including epiphyton) development in shallow littoral zones and may affect other lake primary producers. We hypothesized that epiphyton growth in the littoral zone of temperate oligomesotrophic hard-water lakes could be stimulated by nutrient (especially P) supply via lacustrine groundwater discharge (LGD). We compared the dry mass, chlorophyll a (chl a), and nutrient content of epiphyton grown on artificial substrates at different sites in a groundwater-fed lake and in experimental chambers with and without LGD. During the spring-summer periods, epiphyton accumulated more biomass, especially algae, in littoral LGD sites and in experimental chambers with LGD compared to controls without LGD. Epiphyton chl a accumulation reached up to 46 mg chl a/m(2) after 4 wk when exposed to LGD, compared to a maximum of 23 mg chl a/m(2) at control (C) sites. In the field survey, differences in epiphyton biomass between LGD and C sites were most pronounced at the end of summer, when epilimnetic P concentrations were lowest and epiphyton C:P ratios indicated P limitation. Groundwater-borne P may have facilitated epiphyton growth on macrophytes and periphyton growth on littoral sediments. Epiphyton stored up to 35 mg P/m(2) in 4 wk (which corresponds to 13% of the total P content of the littoral waters), preventing its use by phytoplankton, and possibly contributing to the stabilization of a clear-water state. However, promotion of epiphyton growth by LGD may have contributed to an observed decline in macrophyte abundance caused by epiphyton shading and a decreased resilience of small charophytes to drag forces in shallow littoral areas of the studied lake in recent decades.
Groundwater influx can significantly contribute to nutrient budgets of lakes and its influence is strongest in shallow littoral areas. In oligo-or mesotrophic systems, additional nutrient supply by groundwater influx may affect benthic primary producers and their interactions. Potential changes can be expected in community composition, biomass, stoichiometry and interactions between submerged macrophytes and epiphyton.
The density of organisms declines with size, because larger organisms need more energy than smaller ones and energetic losses occur when larger organisms feed on smaller ones. A potential expression of density-size distributions are Normalized Biomass Size Spectra (NBSS), which plot the logarithm of biomass independent of taxonomy within bins of logarithmic organismal size, divided by the bin width. Theoretically, the NBSS slope of multi-trophic communities is exactly - 1.0 if the trophic transfer efficiency (TTE, ratio of production rates between adjacent trophic levels) is 10% and the predator-prey mass ratio (PPMR) is fixed at 10(4). Here we provide evidence from four multi-trophic lake food webs that empirically estimated TTEs correspond to empirically estimated slopes of the respective community NBSS. Each of the NBSS considered pelagic and benthic organisms spanning size ranges from bacteria to fish, all sampled over three seasons in 1 yr. The four NBSS slopes were significantly steeper than -1.0 (range -1.14 to -1.19, with 95% CIs excluding -1). The corresponding average TTEs were substantially lower than 10% in each of the four food webs (range 1.0% to 3.6%, mean 1.85%). The overall slope merging all biomass-size data pairs from the four systems (-1.17) was almost identical to the slope predicted from the arithmetic mean TTE of the four food webs (-1.18) assuming a constant PPMR of 10(4). Accordingly, our empirical data confirm the theoretically predicted quantitative relationship between TTE and the slope of the biomass-size distribution. Furthermore, we show that benthic and pelagic organisms can be merged into a community NBSS, but future studies have yet to explore potential differences in habitat-specific TTEs and PPMRs. We suggest that community NBSS may provide valuable information on the structure of food webs and their energetic pathways, and can result in improved accuracy of TTE-estimates.
Across a landscape, aquatic-terrestrial interfaces within and between ecosystems are hotspots of organic matter (OM) mineralization. These interfaces are characterized by sharp spatio-temporal changes in environmental conditions, which affect OM properties and thus control OM mineralization and other transformation processes. Consequently, the extent of OM movement at and across aquatic-terrestrial interfaces is crucial in determining OM turnover and carbon (C) cycling at the landscape scale. Here, we propose expanding current concepts in aquatic and terrestrial ecosystem sciences to comprehensively evaluate OM turnover at the landscape scale. We focus on three main concepts toward explaining OM turnover at the landscape scale: the landscape spatiotemporal context, OM turnover described by priming and ecological stoichiometry, and anthropogenic effects as a disruptor of natural OM transfer magnitudes and pathways. A conceptual framework is introduced that allows for discussing the disparities in spatial and temporal scales of OM transfer, changes in environmental conditions, ecosystem connectivity, and microbial-substrate interactions. The potential relevance of priming effects in both terrestrial and aquatic systems is addressed. For terrestrial systems, we hypothesize that the interplay between the influx of OM, its corresponding elemental composition, and the elemental demand of the microbial communities may alleviate spatial and metabolic thresholds. In comparison, substrate level OM dynamics may be substantially different in aquatic systems due to matrix effects that accentuate the role of abiotic conditions, substrate quality, and microbial community dynamics. We highlight the disproportionate impact anthropogenic activities can have on OM cycling across the landscape. This includes reversing natural OM flows through the landscape, disrupting ecosystem connectivity, and nutrient additions that cascade across the landscape. This knowledge is crucial for a better understanding of OM cycling in a landscape context, in particular since terrestrial and aquatic compartments may respond differently to the ongoing changes in climate, land use, and other anthropogenic interferences.
Most studies of aquatic plankton focus on either macroscopic or microbial communities, and on either eukaryotes or prokaryotes. This separation is primarily for methodological reasons, but can overlook potential interactions among groups. Here we tested whether DNA metabarcoding of unfractionated water samples with universal primers could be used to qualitatively and quantitatively study the temporal dynamics of the total plankton community in a shallow temperate lake. Significant changes in the relative proportions of normalized sequence reads of eukaryotic and prokaryotic plankton communities over a 3-month period in spring were found. Patterns followed the same trend as plankton estimates measured using traditional microscopic methods. The bloom of a conditionally rare bacterial taxon belonging to Arcicella was characterized, which rapidly came to dominate the whole lake ecosystem and would have remained unnoticed without metabarcoding. The data demonstrate the potential of universal DNA metabarcoding applied to unfractionated samples for providing a more holistic view of plankton communities.
The sum of benthic autotrophic and bacterial production often exceeds the sum of pelagic autotrophic and bacterial production, and hence may contribute substantially to whole-lake carbon fluxes, especially in shallow lakes. Furthermore, both benthic and pelagic autotrophic and bacterial production are highly edible and of sufficient nutritional quality for animal consumers. We thus hypothesised that pelagic and benthic transfer efficiencies (ratios of production at adjacent trophic levels) in shallow lakes should be similar. We performed whole ecosystem studies in two shallow lakes (3.5ha, mean depth 2m), one with and one without submerged macrophytes, and quantified pelagic and benthic biomass, production and transfer efficiencies for bacteria, phytoplankton, epipelon, epiphyton, macrophytes, zooplankton, macrozoobenthos and fish. We expected higher transfer efficiencies in the lake with macrophytes, because these provide shelter and food for macrozoobenthos and may thus enable a more efficient conversion of basal production to consumer production. In both lakes, the majority of the whole-lake autotrophic and bacterial production was provided by benthic organisms, but whole-lake primary consumer production mostly relied on pelagic autotrophic and bacterial production. Consequently, transfer efficiency of benthic autotrophic and bacterial production to macrozoobenthos production was an order of magnitude lower than the transfer efficiency of pelagic autotrophic and bacterial production to rotifer and crustacean production. Between-lake differences in transfer efficiencies were minor. We discuss several aspects potentially causing the unexpectedly low benthic transfer efficiencies, such as the food quality of producers, pelagic-benthic links, oxygen concentrations in the deeper lake areas and additional unaccounted consumer production by pelagic and benthic protozoa and meiobenthos at intermediate or top trophic levels. None of these processes convincingly explain the large differences between benthic and pelagic transfer efficiencies. Our data indicate that shallow eutrophic lakes, even with a major share of autotrophic and bacterial production in the benthic zone, can function as pelagic systems with respect to primary consumer production. We suggest that the benthic autotrophic production was mostly transferred to benthic bacterial production, which remained in the sediments, potentially cycling internally in a similar way to what has previously been described for the microbial loop in pelagic habitats. Understanding the energetics of whole-lake food webs, including the fate of the substantial benthic bacterial production, which is either mineralised at the sediment surface or permanently buried, has important implications for regional and global carbon cycling.
Sedimentary lipid biomarkers have become widely used tools for reconstructing past climatic and ecological changes due to their ubiquitous occurrence in lake sediments. In particular, the hydrogen isotopic composition (expressed as delta D values) of leaf wax lipids derived from terrestrial plants has been a focus of research during the last two decades and the understanding of competing environmental and plant physiological factors influencing the delta D values has greatly improved. Comparatively less attention has been paid to lipid biomarkers derived from aquatic plants, although these compounds are abundant in many lacustrine sediments. We therefore conducted a field and laboratory experiment to study the effect of salinity and groundwater discharge on the isotopic composition of aquatic plant biomarkers. We analyzed samples of the common submerged plant species, Potamogeton pectinatus (sago pondweed), which has a wide geographic distribution and can tolerate high salinity. We tested the effect of groundwater discharge (characterized by more negative delta D values relative to lake water) and salinity on the delta D values of n-alkanes from P. pectinatus by comparing plants (i) collected from the oligotrophic freshwater Lake Stechlin (Germany) at shallow littoral depth from locations with and without groundwater discharge, and (ii) plants grown from tubers collected from the eutrophic Lake Muggelsee in nutrient solution at four salinity levels. Isotopically depleted groundwater did not have a significant influence on the delta D values of n-alkanes in Lake Stechlin P. pectinatus and calculated isotopic fractionation factors epsilon(l/w) between lake water and n-alkanes averaged -137 +/- 9%(n-C-23), -136 +/- 7%(n-C-25) and -131 +/- 6%(n-C-27), respectively. Similar epsilon values were calculated for plants from Lake Muggelsee grown in freshwater nutrient solution (-134 +/- 11% for n-C-23), while greater fractionation was observed at increased salinity values of 10 (163 +/- 12%) and 15(-172 +/- 15%). We therefore suggest an average e value of -136 +/- 9% between source water and the major n-alkanes in P. pectinatus grown under freshwater conditions. Our results demonstrate that isotopic fractionation can increase by 30-40% at salinity values 10 and 15. These results could be explained either by inhibited plant growth at higher salinity, or by metabolic adaptation to salt stress that remain to be elucidated. A potential salinity effect on dD values of aquatic lipids requires further examination, since this would impact on the interpretation of downcore isotopic data in paleohydrologic studies. (C) 2017 Elsevier Ltd. All rights reserved.
The importance of ciliates as herbivores and in biogeochemical cycles is increasingly recognized. An opportunity to observe the potential consequences of zooplankton dominated by ciliates arose when winter fish kills resulted in strong suppression of crustaceans by young planktivorous fish in two shallow lakes. On an annual average, ciliates made up 38-76% of the total zooplankton biomass in both lakes during two subsequent years. Consequently, ciliate biomass and their estimated grazing potential were extremely high compared with other lakes of various trophic states and depths. Grazing estimates based on abundance and size suggest that ciliates should have cleared the water column of small (<5 mu m) and intermediate (5-50 mu m) sized phytoplankton more than once a day. Especially, small feeders within the ciliates were important, likely exerting a strong top-down control on small phytoplankton. Particle-attached bacteria were presumably strongly suppressed by intermediate-sized ciliate feeders. In contrast to other lakes, large phytoplankton was proportionately very abundant. The phytoplankton community had a high evenness, which may be attributed to the feeding by numerous fast growing and selective ciliate species. Our study highlights ciliates as an important trophic link and adds to the growing awareness of the role of winter processes for plankton dynamics.