Differences in the substrate spectrum of extracellular enzymes in shallow lakes of differing trophic status

Organic matter fluxes and food web interactions in lakes depend on the abilities of heterotrophic microbial communities to access and degrade organic matter, a process that begins with extracellular hydrolysis of high molecular weight substrates. In order to determine whether patterns of enzymatic hydrolysis vary among shallow lakes of different trophic status, we investigated the hydrolysis of six specific organic macromolecules (polysaccharides) in the spring and late summer in four adjacent shallow lakes of eutrophic, oligotrophic, and dystrophic status in coastal North Carolina, USA. The spectrum of enzyme activities detected differed strongly between lakes, with all six polysaccharides hydrolyzed in West Mattamuskeet in May, while only two substrates were hydrolyzed in Lake Phelps in August/September. Differences in the spectrum of enzyme activities, and therefore the capabilities of heterotrophic microbial communities, were likely driven by variations among lakes in primary productivity patterns, sediment–water interactions, and/or water chemistry. Our data represent a first step towards a better understanding of carbon substrate availability and differences in carbon cycling pathways in shallow lakes of different trophic status.     

Plankton photosynthesis, extracellular release and bacterial utilization of released dissolved organic carbon (RDOC) in lakes of different trophy

The major objective of these studies was to assess the extent of the flow of released dissolved organic carbon (RDOC) from phytoplankton to heterotrophic bacteria. The second aim was to test Nalewajko and Schindler's (1976) hypothesis that the percentage of extracellular release (PER) of RDOC by phytoplankton is higher in oligotrophic than eutrophic waters. The studies on the relationship between the bacterial assimilation of RDOC and productivity of phytoplankton in the lakes of different trophy have shown a direct correlation. It was observed that higher utilization rate of phytoplankton RDOC occurred in lakes with higher primary production. The inverse relationship between productivity of lakes, i.e. trophy of waters, and amount of RDOC or PER is closely dependent on the heterotrophic activity of bacteria in waters.     

Carbon flows in eutrophic Lake Rotsee: a <Superscript>13</Superscript>C-labelling experiment

The microbial segment of food webs plays a crucial role in lacustrine food-web functioning and carbon transfer, thereby influencing carbon storage and CO₂ emission and uptake in freshwater environments. Variability in microbial carbon processing (autotrophic and heterotrophic production and respiration based on glucose) with depth was investigated in eutrophic, methane-rich Lake Rotsee, Switzerland. In June 2011, ¹³C-labelling experiments were carried out at six depth intervals in the water column under ambient light as well as dark conditions to evaluate the relative importance of (chemo)autotrophic, mixotrophic and heterotrophic production. Label incorporation rates of phospholipid-derived fatty acid (PLFA) biomarkers allowed us to differentiate between microbial producers and calculate group-specific production. We conclude that at 6 m, net primary production (NPP) rates were highest, dominated by algal photoautotrophic production. At 10 m —the base of the oxycline— a distinct low-light community was able to fix inorganic carbon, while in the hypolimnion, heterotrophic production prevailed. At 2 m depth, high label incorporation into POC could only be traced to nonspecific PLFA, which prevented definite identification, but suggests cyanobacteria as dominating organisms. There was also depth zonation in extracellular carbon release and heterotrophic bacterial growth on recently fixed carbon. Large differences were observed between concentrations and label incorporation of POC and biomarkers, with large pools of inactive biomass settling in the hypolimnion, suggesting late-/post-bloom conditions. Net primary production (115 mmol C m^?2 d^?1) reached highest values in the epilimnion and was higher than glucose-based production (3.3 mmol C m^?2 d^?1, highest rates in the hypolimnion) and respiration (5.9 mmol C m^?2 d^?1, highest rates in the epilimnion). Hence, eutrophic Lake Rotsee was net autotrophic during our experiments, potentially storing large amounts of carbon.     

The effects of shredding invertebrates on the transfer of organic carbon from littoral leaf litter to water-column bacteria

Leaf litter can be of great importance for the productivity of small oligotrophic lakes surrounded by deciduous forests. Feeding invertebrate shredders produce particulate organic leftovers, but their feeding also enhances the release of dissolved organic carbon (DOC). We tested whether invertebrate-mediated DOC release affects the production of heterotrophic water-column bacteria. Submersed leaves were incubated in microcosms with and without shredders; and DOC, absorbance, bacterial abundance and bacterial production in the water column were monitored. We also measured dry weight of the organic particles (FPOC, fine particulate organic carbon, leaf residues and shredders). Total leaf-litter carbon decreased by nearly 80% in the presence of shredders, and on average 56% of the initial leaf carbon ended up as FPOC after 126 days of incubation. Without shredders FPOC production was almost zero, and 72% of the added leaf carbon could be retrieved as leaves when the experiment ended. Both these figures include the rapid release of DOC during the first week of leaf incubation in the lake water (equivalent to 16–19% of total added leaf carbon). Although bacterial production in the water was several times higher in treatments with shredders, bacterial consumption of leaf-derived DOC from shredding was obviously of minor importance in the total carbon budget. This result suggests, although shredders have a strong impact on transformation of leaves to FPOC, they do not greatly enhance the initial rate of mineralization of the leaf-derived detritus.     

Net heterotrophy in Faroe Islands clear-water lakes: causes and consequences for bacterioplankton and phytoplankton

1. Five oligotrophic clear‐water lakes on the Faroe Islands were studied during August 2000. Algal and bacterial production rates, community respiration, and CO₂ saturation were determined. In addition, we examined the plankton community composition (phytoplankton and heterotrophic nanoflagellates) and measured the grazing pressure exerted by common mixotrophic species on bacteria. 2. High respiration to primary production (6.6–33.2) and supersaturation of CO₂ (830–2140 μatm) implied that the lakes were net heterotrophic and that the pelagic heterotrophic plankton were subsidised by allochthonous organic carbon. However, in spite of the apparent high level of net heterotrophy, primary production exceeded bacterial production and the food base for higher trophic levels appeared to be mainly autotrophic. 3. We suggest that the observed net heterotrophy in these lakes was a result of the oligotrophic conditions and hence low primary production in combination with an input of allochthonous C with a relatively high availability. 4. Mixotrophic phytoplankton (Cryptomonas spp., Dinobryon spp. and flagellates cf. Ochromonas spp.) constituted a large percentage of the plankton community (17–83%), possibly as a result of their capacity to exploit bacteria as a means of acquiring nutrients in these nutrient poor systems.     

Variable Production by Different Pelagic Energy Mobilizers in Boreal Lakes

We studied production by three key pelagic energy mobilizer communities, phytoplankton (PP), heterotrophic bacteria (HB), and methanotrophic bacteria (MOB), in five boreal lakes of varying size and concentration of dissolved organic carbon (DOC). Production by PP was responsible for most (>55%) of the total pelagic energy mobilization in all five lakes. Production by HB and PP estimated for the whole water column during the ice-free period were positively correlated, but with the exception of the clearest and most eutrophic lake PP apparently could not support the total carbon demand of bacteria. However, the DOC concentration did not explain the variability of heterotrophic bacterial production (HBP) within or between the lakes. Thus, our results provide circumstantial evidence for the “priming effect” whereby labile organic matter from autochthonous production enhances decomposition of allochthonous DOC. However, HBP was only 10–23% of the total pelagic energy mobilization in the lakes, suggesting that only a minor fraction of allochthonous DOC became available for higher trophic levels. High MOB activity was detected in the water columns of the stratified lakes when the molar ratio of CH₄:O₂ varied between 0.5 and 12. In the small stratified lakes (area < 0.01 km²), MOB production contributed 13–52% of the total pelagic energy mobilization, being greatest during the autumn mixing period. Our results indicate that in small stratified lakes (area < 0.01 km²) bacteria, especially MOB, are potentially quantitatively important supplementary food resources for zooplankton. However, in larger lakes primary producers are the most important (>70%) potential food source for zooplankton.     

Bacterial consumption of total and dissolved organic carbon in the Great Barrier Reef

Heterotrophic bacteria typically take up directly dissolved organic matter due to the small molecular size, although both particulate and dissolved organic matter have labile (easily consumed) compounds. Tropical coastal waters are important ecosystems because of their high productivity. However, few studies have determined bacterial cycling (i.e. carbon uptake by bacteria and allocation for bacterial biomass and respiration) of dissolved organic carbon in coastal tropical waters, and none has determined bacterial cycling of total and dissolved organic carbon simultaneously. In this study we followed bacterial biomass and production, and organic carbon changes over short-term (12 days) dark incubations with (total organic carbon, TOC) and without particulate organic carbon additions (dissolved organic carbon, DOC). The study was performed at three sites along the middle stretch of the Great Barrier Reef (GBR) during the dry and wet seasons. Our results show that the bacterial growth efficiency is low (0.1–11.5%) compared to other coastal tropical systems, and there were no differences in the carbon cycling between organic matter sources, seasons or locations. Nonetheless, more carbon was consumed in the TOC compared to the DOC incubations, although the proportion allocated to biomass and respiration was similar. This suggests that having more bioavailable substrate in the particulate form did not benefit bacteria. Overall, our study indicates that when comparing the obtained respiration rates with previously measured primary production rates, the GBR is a heterotrophic system. More detailed studies are required to fully explore the mechanisms used by bacteria to cycle TOC and DOC in tropical coastal waters.

     

Links between Terrestrial Primary Production and Bacterial Production and Respiration in Lakes in a Climate Gradient in Subarctic Sweden

We compared terrestrial net primary production (NPP) and terrestrial export of dissolved organic carbon (DOC) with lake water heterotrophic bacterial activity in 12 headwater lake catchments along an altitude gradient in subarctic Sweden. Modelled NPP declined strongly with altitude and annual air temperature decreases along the altitude gradient (6°C between the warmest and the coldest catchment). Estimated terrestrial DOC export to the lakes was closely correlated to NPP. Heterotrophic bacterial production (BP) and respiration (BR) were mainly based on terrestrial organic carbon and strongly correlated with the terrestrial DOC export. Excess respiration over PP of the pelagic system was similar to net emission of CO₂ in the lakes. BR and CO₂ emission made up considerably higher shares of the terrestrial DOC input in warm lakes than in cold lakes, implying that respiration and the degree of net heterotrophy in the lakes were dependant not only on terrestrial export of DOC, but also on characteristics in the lakes which changed along the gradient and affected the bacterial metabolization of allochthonous DOC. The study showed close links between terrestrial primary production, terrestrial DOC export and bacterial activity in lakes and how these relationships were dependant on air temperature. Increases in air temperature in high latitude unproductive systems might have considerable consequences for lake water productivity and release of CO₂ to the atmosphere, which are ultimately determined by terrestrial primary production.     

CO2 Saturation and Trophic Shift Induced by Microbial Metabolic Processes in a River-Dominated Ocean Margin (Tropical Shallow Lagoon,Chilika, India)

An effort has been made for the first time in Asia's largest brackish water lagoon, Chilika, to investigate the spatio-temporal variability in primary productivity (PP), bacterial productivity (BP), bacterial abundance (BA), bacterial respiration (BR) and bacterial growth efficiency (BGE) in relation to partial pressure of CO₂ (pCO₂) and CO₂ air–water flux and the resultant trophic switchover. Annually, PP ranged between 24 and 376 µg C L^?1 d^?1 with significantly low values throughout the monsoon (MN), caused by light limitation due to inputs of riverine suspended matter. On the contrary, BP and BR ranged from 11.5 to 186.3 µg C L^?1 d^?1 and from 14.1 to 389.4 µg C L^?1 d^?1, respectively, with exceptionally higher values during MN. A wide spatial and temporal variation in the lagoon trophic status was apparent from BP/PP (0.05–6.4) and PP/BR (0.10–18.2) ratios. The seasonal shift in net pelagic production from autotrophy to heterotrophy due to terrestrial organic matter inputs via rivers, enhanced the bacterial metabolism during the MN, as evident from the high pCO₂ (10,134 µatm) and CO₂ air–water flux (714 mm m^?2 d^?1). Large variability in BGE and BP/PP ratios especially during MN led to high bacteria-mediated carbon fluxes which was evident from significantly high bacterial carbon demand (BCD >100% of PP) during this season. This suggested that the net amount of organic carbon (either dissolved or particulate form) synthesized by primary producers in the lagoon was not sufficient to satisfy the bacterial carbon requirements. Lagoon sustained low to moderate autotrophic–heterotrophic coupling with annual mean BCD of 231% relative to the primary production, which depicted that bacterioplankton are the mainstay of the lagoon biogeochemical cycles and principal players that bring changes in trophic status. Study disclosed that the high CO₂ supersaturation and oxygen undersaturation during MN was attributed to the increased heterotrophic respiration (in excess of PP) fuelled by allochthonous organic matter. On a spatial scale, lagoon sectors such as south sector, central sector and outer channel recorded “net autotrophic,” while the northern sector showed “net heterotrophic” throughout the study period.     

The importance of herbivory by protists in lakes of a tropical floodplain system

Inland aquatic ecosystems play a critical role in the global carbon cycle, processing a great fraction of the organic matter coming from terrestrial ecosystems, and the microbial food web is crucial in this process. Thus, we aimed to evaluate whether the food resource of planktonic protozoa comes mainly from small primary producers or heterotrophic bacteria in tropical shallows lakes, assuming the hypothesis that, in general, picocyanobacteria would be the main food resource for protists. We also expected that the autotrophic fraction would be mainly related to protists at the surface of the environments, while the heterotrophic fraction would be more important at the lower strata of the water column. We performed size-fractionation experiments to evaluate the effects of predation of protists on heterotrophic bacteria and picocyanobacteria. We also sampled planktonic organisms at the subsurface and bottom of 20 lakes in a Neotropical floodplain. We found an herbivory preference of heterotrophic flagellates, while ciliates seem to exert a stronger impact on heterotrophic bacteria. We also found no relationship between heterotrophic bacteria and protists in the field data, whereas positive relationships between picocyanobacteria and protists were observed in environments where there was sunlight. Thus, both heterotrophic bacteria and picocyanobacteria were important components in the food webs of tropical shallow lakes. Moreover, the trophic cascade caused by zooplankton predation suggests that protists are efficient in transferring the energy from the base of microbial food webs to higher trophic levels.     

Carbon dioxide supersaturation promotes primary production in lakes

A majority of the world's lakes are supersaturated with respect to carbon dioxide (CO(2) ). By experimental manipulation of the CO(2) concentration in supersaturated boreal lakes, we demonstrate that phytoplankton primary production was up to 10 times higher in supersaturated lake water in comparison with water with CO(2) at equilibrium concentrations and that CO(2) , together with nutrients, explained most of the variation in pelagic primary production and phytoplankton biomass over a wide variety of unproductive lakes. These results suggest that phytoplankton can be co-limited by CO(2) and nutrients in unproductive lakes. As import of terrestrial organic carbon and its subsequent microbial mineralisation in lakes is a driving force of CO(2) -supersaturation our results suggest that lake productivity and carbon cycling may respond to variations in terrestrial organic carbon export, (e.g. caused by land use or climate change) in ways not described before.     

Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l⁻¹) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes. This paper is dedicated to the memory of Prof. Peter Blomqvist (deceased 2004).     

Net pelagic heterotrophy in mesotrophic and oligotrophic basins of a large, temperate lake

Understanding the effects of trophic status and dissolved organic carbon concentration (DOC) on lake carbon cycling is essential for accurate ecosystem carbon models. Using isotopically labelled substrates we assessed spatial and temporal variability in bacterial respiration (BR) and algal primary production (PP) in two trophically, morphometrically and hydrologically different basins in Loch Lomond, a large temperate lake in Scotland. GIS modelling was used to construct a whole lake balance for bacterial production/respiration and PP, and from this the proportion of heterotrophy fuelled by allochthonous carbon was estimated. We tested the hypotheses that trophic status and DOC concentration affect the balance between PP and BR and examined which is the more significant driving factor. Additionally we estimated the percentage of BR that is fuelled by terrestrial carbon. PP varied seasonally and showed inter-basin homogeneity. BR was greatest in the mesotrophic south basin in autumn, which corresponded to measured peak DOC input, though over an annual cycle no relationship was observed between BR and DOC concentration. The PP:BR ratio was 0.37 ± 0.30 and 0.3 ± 0.45 in the north and south basins, respectively, assuming a bacterial growth efficiency of 0.1. We have found that allochthonous carbon potentially supports a substantial quantity of pelagic production, even during periods of high photosynthesis. Less productive systems are thought to be dominated by heterotrophic processes. However, we have found that the mesotrophic basin of a large lake to be as heterotrophic as its neighbouring oligotrophic basin, an observation that has implications for our understanding of modelling of the role of lakes in linking the terrestrial-atmospheric carbon cycle.     

Bacterial Growth on Allochthonous Carbon in Humic and Nutrient-enriched Lakes: Results from Whole-Lake 13C Addition Experiments

Organic carbon (C) in lakes originates from two distinct sources—primary production from within the lake itself (autochthonous supply) and importation of organic matter from the terrestrial watershed (allochthonous supply). By manipulating the ¹³C of dissolved inorganic C, thereby labeling within-lake primary production, we examined the relative importance of autochthonous and allochthonous C in supporting bacterial production. For 35 days, NaH¹³CO₃ was added daily to two small, forested lakes. One of the lakes (Peter) was fertilized so that primary production exceeded total respiration in the epilimnion. The other lake (Tuesday), in contrast, was low in productivity and had high levels of colored dissolved organic C (DOC). To obtain bacterial C isotopes, bacteria were regrown in situ in particle-free lake water in dialysis tubes. The contribution of allochthonous C to bacterial biomass was calculated by applying a two-member mixing model. In the absence of a direct measurement, the isotopic signature of the autochthonous end-member was estimated indirectly by three different approaches. Although there was excess primary production in Peter Lake, bacterial biomass consisted of 43–46% allochthonous C. In Tuesday Lake more than 75% of bacterial growth was supported by allochthonous C. Although bacteria used autochthonous C preferentially over allochthonous C, DOC from the watershed contributed significantly to bacterial production. In combination with results from similar experiments in different lakes, our findings suggest that the contribution of allochthonous C to bacterial production can be predicted from ratios of chromophoric dissolved organic matter (a surrogate for allochthonous supply) and chlorophyll a (a surrogate for autochthonous supply).     

Tidal impact on planktonic primary and bacterial production in the German Wadden Sea

 Tidal variation of biological parameters was studied at three anchor stations in selected inlet channels of the northern German Wadden Sea in May and July 1994. Concentrations of bacteria, chlorophyll a and suspended matter as well as primary and bacterial production were assessed over a period of 25 h in the surface and in the bottom water. Diurnal variation in primary production was found both under in situ light conditions and under constant illumination. Tidal turbulence caused the introduction of detritus, bacteria and pigments from the sediment into the water column. The impact of sediment resuspension was most evident in the bottom water, leading to tidally oscillating bacterial production rates which were high during high stream velocity and low during the slack times. Estimations of the areal daily phytoplankton production and corresponding bacterial carbon demands were unbalanced. Primary production accounted for only 25–45% of the total bacterial carbon requirement. This discrepancy is due to the shallow euphotic depth in the Wadden Sea, allowing net primary production only in the upper 2–3 m of the water column, while the relatively high levels of bacterial activity do not show a vertical decline. Assuming that the specific biological activities in the water columns over the tidal flats are similar to those found in the inlet channels, it was found that production processes dominate in shallow areas whereas decomposition processes dominate in the deep channels. Moreover, the predominance of heterotrophic processes in the inlet channels means that additional organic carbon sources must contribute to the heterotrophic metabolism in the deep parts of the Wadden Sea, and that the horizontal flux of material is important in this turbid mesotidal ecosystem. Received: 18 April 1998 / Accepted: 12 November 1998     

Mass flux calculations show strong allochthonous support of freshwater zooplankton production is unlikely

Many studies have concluded terrestrial carbon inputs contribute 20-70% of the carbon supporting zooplankton and fish production in lakes. Conversely, it is also known that terrestrial carbon inputs are of very low nutritional quality and phytoplankton are strongly preferentially utilized by zooplankton. Because of its low quality, substantial terrestrial support of zooplankton production in lakes is only conceivable when terrigenous organic matter inputs are much larger than algal production. We conducted a quantitative analysis of terrestrial carbon mass influx and algal primary production estimates for oligo/mesotrophic lakes (i.e., TP ≤ 20 μg L(-1)). In keeping with the principle of mass conservation, only the flux of terrestrial carbon retained within lakes can be utilized by zooplankton. Our field data compilation showed the median (inter-quartile range) terrestrial particulate organic carbon (t-POC), available dissolved organic carbon (t-DOC) inputs, and in-lake bacterial and algal production were 11 (8-17), 34 (11-78), 74 (37-165), and 253 (115-546) mg C m(-2) d(-1), respectively. Despite the widespread view that terrestrial inputs dominate the carbon flux of many lakes, our analysis indicates algal production is a factor 4-7 greater than the available flux of allochthonous basal resources in low productivity lakes. Lakes with high loading of t-DOC also have high hydraulic flushing rates. Because t-DOC is processed, i.e., mineralized or lost to the sediments, in lakes at ≈ 0.1% d(-1), in systems with the highest t-DOC inputs (i.e., 1000 mg m(-2) d(-1)) a median of 98% of the t-DOC flux is advected and therefore is not available to support zooplankton production. Further, advection is the primary fate of t-DOC in lakes with hydraulic retention times <3 years. When taking into account the availability and quality of terrestrial and autochthonous fluxes, this analysis indicates ≈ 95-99% of aquatic herbivore production is supported by in-lake primary production.     

Comparison of bacterial production in sediments, epiphyton and the pelagic zone of a lowland river

1. The microbial metabolism of organic matter in rivers has received little study compared with that of small streams. Therefore, we investigated the rate and location of bacterial production in a sixth‐order lowland river (Spree, Germany). To estimate the contribution of various habitats (sediments, epiphyton, and the pelagic zone) to total bacterial production, we quantified the contribution of these habitats to areal production by bacteria. 2. Large areas of the river bottom were characterized by loose and shifting sands of relatively homogenous particle size distribution. Aquatic macrophytes grew on 40% of the river bottom. Leaf areas of 2.8 m² m^?2 river bottom were found in a 6.6 km river stretch. 3. The epiphyton supported a bacterial production of 5–58 ng C cm^?2 h^?1. Bacterial production in the pelagic zone was 0.9–3.9 μg C L^?1 h^?1, and abundance was 4.0–7.8 × 10⁹ cells L^?1. Bacterial production in the uppermost 2 cm of sediments ranged from 1 to 8 μg C cm^?3 h^?1, and abundance from 0.84 to 6.7 × 10⁹ cells cm^?3. Bacteria were larger and more active in sediments than in the pelagic zone. 4. In spite of relatively low macrophyte abundance, areal production by bacteria in the pelagic zone was only slightly higher than in the epiphyton. Bacterial biomass in the uppermost 2 cm of sediments exceeded pelagic biomass by factors of 6–22, and sedimentary bacterial production was 17–35 times higher than in the overlying water column. 5. On a square meter basis, total bacterial production in the Spree was clearly higher than primary productivity. Thus, the lowland river Spree is a heterotrophic system with benthic processes dominating. Therefore, sedimentary and epiphytic bacterial productivity form important components of ecosystem carbon metabolism in rivers and shallow lakes. 6. The sediments are focal sites of microbial degradation of organic carbon in a sand‐bottomed lowland river. The presence of a lowland river section within a river continuum probably greatly changes the geochemical fluxes within the river network. This implies that current concepts of longitudinal biogeochemical relationships within river systems have to be revised.     

Spatial and Temporal Variability in the Ecosystem Metabolism of a High-elevation Lake: Integrating Benthic and Pelagic Habitats

We characterized spatial and temporal variability in net ecosystem production (NEP), community respiration (CR), and gross primary production (GPP) over an ice-free season in an oligotrophic high-elevation lake using high-frequency measurements of dissolved oxygen. We combined the use of free-water and incubation chamber measurements to compare pelagic and benthic habitats and estimate their relative contributions to whole-lake metabolism. Despite a brief period of predominant heterotrophy after snowmelt, both free-water and incubation chamber measurements confirmed autotrophy of the epilimnion in all habitats throughout the ice-free season. In contrast, benthic incubation chambers showed the benthos to be consistently heterotrophic. Although temperature was the strongest seasonal driver of benthic metabolism, bacterioplankton density and indexes of organic matter quality explained the most variability in pelagic metabolism. Driven largely by benthic metabolism, free-water measurements of GPP and CR were twice as high in littoral than pelagic habitats. However, rates of water column primary production overlying the littoral benthos were high enough to overcome net benthic heterotrophy, and seasonal mean NEP in littoral habitats remained positive and not significantly different from pelagic habitats. Benthic rates averaged about 25% of whole-lake metabolism. Pelagic metabolism measurements were affected by littoral rates about half the time, with the degree of isolation between the two a function of advection and water column stability. These results emphasize the importance of characterizing spatial and temporal variability in metabolism within the context of physical dynamics and challenge the notion that benthic metabolism will necessarily be larger than pelagic metabolism in oligotrophic lakes.     

Relationship between δ13C of chironomid remains and methane flux in Swedish lakes

1. Methanogenic carbon can be incorporated by methane‐oxidising bacteria, leading to a ¹³C‐depleted stable carbon isotopic composition (δ¹³C) of chironomids that feed on these microorganisms. This has been shown for the chironomid tribe Chironomini, but very little information is available about the δ¹³C of other abundant chironomid groups and the relationship between chironomid δ¹³C and methane production in lakes. 2. Methane flux was measured at the water surface of seven lakes in Sweden. Furthermore, fluxes from the sediments to the water column were measured in transects in two of the lakes. Methane fluxes were then compared with δ¹³C of chitinous chironomid remains isolated from the lake surface sediments. Several different chironomid groups were examined (Chironomini, Orthocladiinae, Tanypodinae and Tanytarsini). 3. Remains of Orthocladiinae in the seven study lakes had the highest δ¹³C values (?31.3 to ?27.0‰), most likely reflecting δ¹³C of algae and other plant‐derived organic matter. Remains of Chironomini and Tanypodinae had lower δ¹³C values (?33.2 to ?27.6‰ and ?33.6 to ?28.0‰, respectively). A significant negative correlation was observed between methane fluxes at the lake surface and δ¹³C of Chironomini (r = ?0.90, P = 0.006). Methane release from the sediments was also negatively correlated with δ¹³C of Chironomini (r = ?0.67, P = 0.025) in the transect samples obtained from two of the lakes. The remains of other chironomid taxa were only weakly or not correlated with methane fluxes measured in our study lakes (P > 0.05). 4. Selective incorporation of methane‐derived carbon can explain the observed correlations between methane fluxes and δ¹³C values of Chironomini. Remains of this group might therefore have the potential to provide information about past changes in methane availability in lakes using sediment records. However, differences in productivity, algal δ¹³C composition and the importance of allochthonous organic matter input between the studied lakes may also have influenced Chironomini δ¹³C. More detailed studies with a higher number of analysed samples and detailed measurement of δ¹³C of different ecosystem components (e.g. methane, dissolved inorganic carbon) will be necessary to further resolve the relative contribution of different carbon sources to δ¹³C of chironomid remains.     

Extracellular dissolved organic carbon released from phytoplankton as a source of carbon for heterotrophic bacteria in lakes of different humic content

The quantitative importance of photosynthetically produced dissolved organic carbon (PDOC) released from phytoplankton as a source of carbon for pelagic, heterotrophic bacteria was investigated in four temperate Swedish lakes, of which two had low (≈20 mg Pt 1⁻¹), and two moderately high (60–80 mg Pt 1⁻¹) humic content. The bacterial assimilation of PDOC was estimated with the ¹⁴C method, and the total production of the heterotrophic bacteria was estimated with the [3H]thymidine incorporation method. The release of PDOC from natural communities of phytoplankton was not restricted to periods of photosynthesis, but often continued during periods of darkness. Heterotrophic bacteria often assimilated the labile components of the PDOC at high rates (up to 73% of the released PDOC was assimilated during the incubation in our experiments). The contribution of PDOC to bacterial production exhibited large within-lake seasonal variations, but PDOC was at certain times, both in humic and non-humic lakes, a quantitatively very important carbon source for the heterotrophic bacteria. Under periods of comparatively low primary production, heterotrophic bacteria in humic lakes appear to utilize allochthonous, humic substances as a substrate.