Roger I. Jones

WIDESPREAD CONTRIBUTION OF METHANE‐CYCLE BACTERIA TO THE DIETS OF LAKE PROFUNDAL CHIRONOMID LARVAE

Reports of unexpectedly 13C-depleted chironomid larvae in lakes have led to an hypothesis that significant transfer of detrital organic matter to chironomid larvae may occur via methane-cycle bacteria. However, to date little is known of how such transfer might vary across species and lakes. We gathered data from 87 lakes to determine how widespread this phenomenon might be and to define boundaries for its likely magnitude. Carbon stable isotope values of chironomid larvae varied greatly between taxa. Very marked 13C-depletion was evident only in certain taxa, especially Chironomus plumosus, C. anthracinus, and C. tenuistylus, all characteristic of eutrophic or dystrophic lakes and known to be tolerant of low oxygen conditions. Furthermore, marked 13C-depletion was only found in larvae from lakes in which late-summer hypolimnetic oxygen depletion near the sediment surface was below an apparent threshold concentration of 2-4 mg O2/L. Similarly, application of a two-source mixing model suggested that methanotrophic bacteria made the greatest contribution to profundal chironomid growth (0-70% of larval carbon) when the late-summer oxygen concentration dropped below approximately 2 mg O2/L. Our study demonstrates that methane-derived carbon is an important, but often neglected, contribution to the flux of carbon through the food webs of many productive or dystrophic lakes.

An Empirical Evaluation of the Utility of Convex Hull and Standard Ellipse Areas for Assessing Population Niche Widths from Stable Isotope Data

Stable isotope analyses are increasingly employed to characterise population niche widths. The convex hull area (TA) in a δ13C–δ15N biplot has been used as a measure of isotopic niche width, but concerns exist over its dependence on sample size and associated difficulties in among-population comparisons. Recently a more robust method was proposed for estimating and comparing isotopic niche widths using standard ellipse areas (SEA), but this approach has yet to be tested with empirical stable isotope data. The two methods measure different kind of isotopic niche areas, but both are now widely used to characterise isotopic niche widths of populations. We used simulated data and an extensive empirical dataset from two fish populations to test the influence of sample size on the observed isotopic niche widths (TA and SEA). We resampled the original datasets to generate 5000 new samples for different numbers of observations from 5 to 80 to examine the statistical distributions of niche area estimates for increasing sample size. Our results illustrate how increasing sample size increased the observed TA; even sample sizes much higher than nu200a=u200a30 did not improve the precision for the TA method. SEA was less sensitive to sample size, but the natural variation in our empirical fish δ13C and δ15N data still resulted in considerable uncertainty around the mean estimates of niche width, reducing the precision particularly with sample sizes n<30. These results confirm that the TA method is less appropriate for estimating population isotopic niche areas using small samples, especially when considerable population level isotope variation is expected. The results also indicate a need for caution when using SEA as a measure of trophic niche widths for consumers, particularly with low sample sizes and when the distribution and range for population isotope values are not known.

Distinct and diverse anaerobic bacterial communities in boreal lakes dominated by candidate division OD1

Lakes have a central role in the carbon cycle of the boreal landscape. These systems typically stratify in summer and their hypolimnetic microbial communities influence burial of biogenic organic matter in sediments. The composition of bacterial communities in these suboxic habitats was studied by pyrosequencing of 16S rRNA amplicons from five lakes with variable dissolved organic carbon (DOC) concentrations. Bacterioplankton communities in the hypolimnetic waters were clearly different from the surface layer with candidate division OD1, Chlorobi and Bacteroidetes as dominant community members. Several operational taxonomic units (OTUs) affiliated with candidate division OD1 were abundant and consistently present in the suboxic hypolimnion in these boreal lakes. The overall representation of this group was positively correlated with DOC and methane concentrations. Network analysis of time-series data revealed contrasting temporal patterns but suggested similar ecological roles among the abundant OTUs affiliated with candidate division OD1. Together, stable isotope data and taxonomic classification point to methane oxidation and autotrophic denitrification as important processes in the suboxic zone of boreal lakes. Our data revealed that while hypolimnetic bacterial communities are less dynamic, they appear to be more diverse than communities from the oxic surface layer. An appreciable proportion of the hypolimnetic bacteria belong to poorly described phyla.

Seasonal and ontogenetic shifts in the diet of Arctic charr Salvelinus alpinus in a subarctic lake

Seasonal and ontogenetic shifts in the diet of Arctic charr Salvelinus alpinus were studied in a deep, ultra-oligotrophic lake in subarctic Finland from both stomach contents and the stable carbon (delta(13)C) and nitrogen (delta(15)N) isotope compositions of muscle and liver tissues. Both diet and isotope results indicated that the S. alpinus population relied mainly on littoral benthic energy sources. The strong littoral reliance appeared largely independent of season or fish size, although the data lacked small (total length, L(T), <130 mm) and young (<3 years) S. alpinus. Liver isotope values of intermediate-sized S. alpinus (200-350 mm), however, suggested exploitation of the increase in the abundance of pelagic zooplankton in the late open-water season. The results suggest that, in general, a strong littoral reliance of fishes can be a feature in subarctic lakes throughout the year. Due to its faster isotopic turnover rate and thus higher resolution for temporal diet changes, liver could be more commonly used in stable-isotope studies of fish trophic niche shifts instead of using only the less responsive muscle tissue.

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food-web stability. In lakes, littoral and pelagic food-web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high-latitude lakes. We analyzed food-web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km2) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food-chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate-dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high-latitude lakes.

The importance of sedimenting organic matter, relative to oxygen and temperature, in structuring lake profundal macroinvertebrate assemblages

We quantified the role of a main food resource, sedimenting organic matter (SOM), relative to oxygen (DO) and temperature (TEMP) in structuring profundal macroinvertebrate assemblages in boreal lakes. SOM from 26 basins of 11 Finnish lakes was analysed for quantity (sedimentation rates), quality (C:N:P stoichiometry) and origin (carbon stable isotopes, δ13C). Hypolimnetic oxygen and temperature were measured from each site during summer stratification. Partial canonical correspondence analysis (CCA) and partial regression analyses were used to quantify contributions of SOM, DO and TEMP to community composition and three macroinvertebrate metrics. The results suggested a major contribution of SOM in regulating the community composition and total biomass. Oxygen best explained the Shannon diversity, whereas TEMP had largest contribution to the variation of Benthic Quality Index. Community composition was most strongly related to δ13C of SOM. Based on additional δ13C and stoichiometric analyses of chironomid taxa, marked differences were apparent in their utilization of SOM and body stoichiometry; taxa characteristic of oligotrophic conditions exhibited higher C:N ratios and lower C:P and N:P ratios compared to the species typical of eutrophic lakes. The results highlight the role of SOM in regulating benthic communities and the distributions of individual species, particularly in oligotrophic systems.

Inferring Phytoplankton, Terrestrial Plant and Bacteria Bulk δ¹³C Values from Compound Specific Analyses of Lipids and Fatty Acids.

Stable isotope mixing models in aquatic ecology require δ13C values for food web end members such as phytoplankton and bacteria, however it is rarely possible to measure these directly. Hence there is a critical need for improved methods for estimating the δ13C ratios of phytoplankton, bacteria and terrestrial detritus from within mixed seston. We determined the δ13C values of lipids, phospholipids and biomarker fatty acids and used these to calculate isotopic differences compared to the whole-cell δ13C values for eight phytoplankton classes, five bacterial taxa, and three types of terrestrial organic matter (two trees and one grass). The lipid content was higher amongst the phytoplankton (9.5±4.0%) than bacteria (7.3±0.8%) or terrestrial matter (3.9±1.7%). Our measurements revealed that the δ13C values of lipids followed phylogenetic classification among phytoplankton (78.2% of variance was explained by class), bacteria and terrestrial matter, and there was a strong correlation between the δ13C values of total lipids, phospholipids and individual fatty acids. Amongst the phytoplankton, the isotopic difference between biomarker fatty acids and bulk biomass averaged -10.7±1.1‰ for Chlorophyceae and Cyanophyceae, and -6.1±1.7‰ for Cryptophyceae, Chrysophyceae and Diatomophyceae. For heterotrophic bacteria and for type I and type II methane-oxidizing bacteria our results showed a -1.3±1.3‰, -8.0±4.4‰, and -3.4±1.4‰ δ13C difference, respectively, between biomarker fatty acids and bulk biomass. For terrestrial matter the isotopic difference averaged -6.6±1.2‰. Based on these results, the δ13C values of total lipids and biomarker fatty acids can be used to determine the δ13C values of bulk phytoplankton, bacteria or terrestrial matter with ± 1.4‰ uncertainty (i.e., the pooled SD of the isotopic difference for all samples). We conclude that when compound-specific stable isotope analyses become more widely available, the determination of δ13C values for selected biomarker fatty acids coupled with established isotopic differences, offers a promising way to determine taxa-specific bulk δ13C values for the phytoplankton, bacteria, and terrestrial detritus embedded within mixed seston.

Top consumer abundance influences lake methane efflux

Lakes are important habitats for biogeochemical cycling of carbon. The organization and structure of aquatic communities influences the biogeochemical interactions between lakes and the atmosphere. Understanding how trophic structure regulates ecosystem functions and influences greenhouse gas efflux from lakes is critical to understanding global carbon cycling and climate change. With a whole-lake experiment in which a previously fishless lake was divided into two treatment basins where fish abundance was manipulated, we show how a trophic cascade from fish to microbes affects methane efflux to the atmosphere. Here, fish exert high grazing pressure and remove nearly all zooplankton. This reduction in zooplankton density increases the abundance of methanotrophic bacteria, which in turn reduce CH4 efflux rates by roughly 10 times. Given that globally there are millions of lakes emitting methane, an important greenhouse gas, our findings that aquatic trophic interactions significantly influence the biogeochemical cycle of methane has important implications.

Bacterial and phytoplankton responses to nutrient amendments in a boreal lake differ according to season and to taxonomic resolution.

Nutrient limitation and resource competition in bacterial and phytoplankton communities may appear different when considering different levels of taxonomic resolution. Nutrient amendment experiments conducted in a boreal lake on three occasions during one open water season revealed complex responses in overall bacterioplankton and phytoplankton abundance and biovolume. In general, bacteria were dominant in spring, while phytoplankton was clearly the predominant group in autumn. Seasonal differences in the community composition of bacteria and phytoplankton were mainly related to changes in observed taxa, while the differences across nutrient treatments within an experiment were due to changes in relative contributions of certain higher- and lower-level phylogenetic groups. Of the main bacterioplankton phyla, only Actinobacteria had a treatment response that was visible even at the phylum level throughout the season. With increasing resolution (from 75 to 99% sequence similarity) major responses to nutrient amendments appeared using 454 pyrosequencing data of 16S rRNA amplicons. This further revealed that OTUs (defined by 97% sequence similarity) annotated to the same highly resolved freshwater groups appeared to occur during different seasons and were showing treatment-dependent differentiation, indicating that OTUs within these groups were not ecologically coherent. Similarly, phytoplankton species from the same genera responded differently to nutrient amendments even though biovolumes of the majority of taxa increased when both nitrogen and phosphorus were added simultaneously. The bacterioplankton and phytoplankton community compositions showed concurrent trajectories that could be seen in synchronous succession patterns over the season. Overall, our data revealed that the response of both communities to nutrient changes was highly dependent on season and that contradictory results may be obtained when using different taxonomic resolutions.

Enhanced greenhouse gas emissions and changes in plankton communities following an experimental increase in organic carbon loading to a humic lake

Organic carbon concentrations in the surface waters of the boreal region have increased during the past two decades. We investigated the impact of elevated dissolved organic carbon (DOC) loading to a humic lake by a whole-lake experiment in which DOC in the form of cane sugar was added monthly during the ice-free period over two consecutive years. The sugar addition represented an increased concentration of 2xa0mgxa0l−1 of DOC in the epilimnion and led to an increase in CO2 emission and also an apparent increase in CH4 emission to the atmosphere from the lake surface. The composition of the bacterial, phytoplankton and zooplankton communities altered during the study period and the bacterial abundance in the metalimnion and hypolimnion of the lake decreased. No changes were detected in epilimnetic primary production or respiration, but there was an increase in bacterial production in the epilimnion. The nutrient and particulate organic carbon concentrations also suggested possible changes in the activity of heterotrophic bacteria in the metalimnion. Carbon stable isotope analyses indicated transfer of some added sugar carbon through the food web to zooplankton consumers. Overall the results suggest that future increases in organic carbon loading to boreal lakes will increase greenhouse gas emissions, although the magnitude of any change is likely to depend on the availability of nutrients like phosphorus and nitrogen which influence organic matter processing and the development of plankton communities.