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Institute
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.
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.
Regime shifts are commonly associated with the loss of submerged macrophytes in shallow lakes; yet, the effects of this on whole-lake primary productivity remain poorly understood. This study compares the annual gross primary production (GPP) of two shallow, eutrophic lakes with different plant community structures but similar nutrient concentrations. Daily GPP rates were substantially higher in the lake containing submerged macrophytes (58623gCm(-2)year(-1)) than in the lake featuring only phytoplankton and periphyton (40823gCm(-2)year(-1); P<0.0001). Comparing lake-centre diel oxygen curves to compartmental estimates of GPP confirmed that single-site oxygen curves may provide unreliable estimates of whole-lake GPP. The discrepancy between approaches was greatest in the macrophyte-dominated lake during the summer, with a high proportion of GPP occurring in the littoral zone. Our empirical results were used to construct a simple conceptual model relating GPP to nutrient availability for these alternative ecological regimes. This model predicted that lakes featuring submerged macrophytes may commonly support higher rates of GPP than phytoplankton-dominated lakes, but only within a moderate range of nutrient availability (total phosphorus ranging from 30 to 100gL(-1)) and with mean lake depths shallower than 3 or 4m. We conclude that shallow lakes with a submerged macrophyte-epiphyton complex may frequently support a higher annual primary production than comparable lakes that contain only phytoplankton and periphyton. We thus suggest that a regime shift involving the loss of submerged macrophytes may decrease the primary productivity of many lakes, with potential consequences for the entire food webs of these ecosystems.
Submerged macrophytes can stabilise clear water conditions in shallow lakes. However, many existing models for deep lakes neglect their impact. Here, we tested the hypothesis that submerged macrophytes can affect the water clarity in deep lakes. A one-dimensional, vertically resolved macrophyte model was developed based on PCLake and coupled to SALMO-1D and GOTM hydrophysics and validated against field data. Validation showed good coherence in dynamic growth patterns and colonisation depths. In our simulations the presence of submerged macrophytes resulted in up to 50% less phytoplankton biomass in the shallowest simulated lake (11 m) and still 15% less phytoplankton was predicted in 100 m deep oligotrophic lakes. Nutrient loading, lake depth, and lake shape had a strong influence on macrophyte effects. Nutrient competition was found to be the strongest biological interaction. Despite a number of limitations, the derived dynamic lake model suggests significant effects of submerged macrophytes on deep lake water quality. (C) 2014 Elsevier Ltd. All rights reserved.
The amount of terrestrial particulate organic matter (t-POM) entering lakes is predicted to increase as a result of climate change. This may especially alter the structure and functioning of ecosystems in small, shallow lakes which can rapidly shift from a clear-water, macrophyte-dominated into a turbid, phytoplankton-dominated state. We used the integrative ecosystem model PCLake to predict how rising t-POM inputs affect the resilience of the clear-water state. PCLake links a pelagic and benthic food chain with abiotic components by a number of direct and indirect effects. We focused on three pathways (zoobenthos, zooplankton, light availability) by which elevated t-POM inputs (with and without additional nutrients) may modify the critical nutrient loading thresholds at which a clear-water lake becomes turbid and vice versa. Our model results show that (1) increased zoobenthos biomass due to the enhanced food availability results in more benthivorous fish which reduce light availability due to bioturbation, (2) zooplankton biomass does not change, but suspended t-POM reduces the consumption of autochthonous particulate organic matter which increases the turbidity, and (3) the suspended t-POM reduces the light availability for submerged macrophytes. Therefore, light availability is the key process that is indirectly or directly changed by t-POM input. This strikingly resembles the deteriorating effect of terrestrial dissolved organic matter on the light climate of lakes. In all scenarios, the resilience of the clear-water state is reduced thus making the turbid state more likely at a given nutrient loading. Therefore, our study suggests that rising t-POM input can add to the effects of climate warming making reductions in nutrient loadings even more urgent.
Groundwater influx can significantly contribute to nutrient and carbon budgets of lakes, and its influence is the strongest in littoral areas dominated by macrophytes and periphyton. We have reviewed the effects of groundwater-borne nitrogen and phosphorus and dissolved inorganic and organic carbon (DIC, DOC) on these benthic primary producers in lakes. We develop a hypothesis for groundwater effects including the less studied impacts of periphyton shading on macrophytes. Groundwater-borne nutrients and DIC promote both macrophytes and periphyton. Direct studies on groundwater-borne DOC effects are lacking, but coloured DOC contributes to light attenuation and thus can restrict the growth of benthic primary producers. We predict that above certain threshold levels of nutrient influx by groundwater, periphyton and macrophyte biomass should decline owing to shading by phytoplankton and periphyton, respectively. However, because of their higher light requirements, those thresholds should be lower for macrophytes. For macrophytes, a threshold level is also predicted for a shift from DIC limitation to light limitation. Differences in light requirements are expected to result in lower thresholds of DOC loading for declines of macrophytes than periphyton.
Contrasting response of two shallow eutrophic cold temperate lakes to a partial winterkill of fish
(2015)
Food-web effects of winterkill are difficult to predict as the enhanced mortality of planktivorous fish may be counterbalanced by an even higher mortality of piscivores. We hypothesised that a winterkill in a clear and a turbid shallow lake would equalise their fish community composition, but seasonal plankton successions would differ between lakes. After a partial winterkill, we observed a reduction of fish biomass by 16 and 43% in a clear-water and a turbid small temperate lake, respectively. Fish biomass and piscivore shares (5% of fish biomass) were similar in both lakes after this winterkill, but young-of-the-year (YOY) abundances were higher in the turbid lake. Top-down control by crustaceans was only partly responsible for low phytoplankton biomass at the end of May following the winterkill in both lakes. Summer phytoplankton biomass remained low in the clear-water lake despite high abundances of YOY fish (mainly roach). In contrast, the crustacean biomass of the turbid lake was reduced in summer by a high YOY abundance (sunbleak and roach), leading to a strong increase in phytoplankton biomass. The YOY abundance of fish in shallow eutrophic lakes may thus be more important for their summer phytoplankton development after winterkill than the relative abundance of piscivores.
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.
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.