Effects of Deposit-Feeding Macrofauna on Benthic Bacteria,Viruses, and Protozoa in a Silty Freshwater Sediment

In microcosm experiments, we simultaneously tested the effects of increased numbers of deposit-feeding macrofauna (chironomids, oligochaetes and cladocerans) on the standing stock, activities and interactions of heterotrophic bacteria, viruses, and bacterivorous protozoa (heterotrophic nanoflagellates and ciliates) in the aerobic layer of a silty littoral freshwater sediment. On average, bacterial secondary production was stimulated between 11 and 29% by all macrofaunal groups compared to control experiments without macrofauna addition. Bacterial standing stock increased significantly by 8 and 13% in case of chironomids and cladocerans, respectively. Oligochaetes and chironomids produced significant negative effects on viral abundance while the results with cladocerans were inconsistent. The addition of oligochaetes and chironomids resulted in a significant decrease by on average 68 and 32% of viral decay rates, respectively, used as a measure of viral production. The calculated contribution of virus-induced lysis to benthic bacterial mortality was low, with 2.8 to 11.8% of bacterial secondary production, and decreased by 39 to 81% after the addition of macrofauna compared to the control. The abundances of heterotrophic nanoflagellates were significantly reduced by 20% by all tested macrofauna groups, while ciliates showed inconsistent results. The importance of heterotrophic nanoflagellate grazing on benthic bacteria was very low (<1% of bacterial secondary production) and was further reduced by elevated numbers of macrofauna. Thus, the selected deposit feeding macrofauna groups seem to have several direct and indirect and partly antagonistic effects on the benthic bacterial compartment through the enhancement of bacterial production and the reduction of virus-induced cell lysis and protozoan grazing.     

Biological consequences of stream routing through a final-cut Strip mine pit: Benthic macroinvertebrates

Benthic macroinvertebrates were collected from the Plachek Pit, a final-cut coal strip mine pit at the Big Horn Mine near Sheridan, Wyoming, USA. Goose Creek was routed through the pit to allow mining under the original creek bed. The benthic community in the pit was dominated by the worm Limnodrilus hoffmeisteri and various midge genera, including Chironomus sp., Procladius sp. and Tanypus stellatus. Insects other than chironomids were minor components of the benthic fauna, and were collected only in the upstream portions of the pit; they were concluded to have originated from drift deposition. The pit functions as a sink for drifting macroinvertebrates. Based on community composition and supplemental water quality information it was concluded that the Plachek Pit best exemplified an organically-enriched, lake-like ecosystem. Overall, rerouting of Goose Creek through the Plachek Pit was not viewed as deleterious to Goose Creek, but rather benefited the creek by serving as a sink for inputs from upstream municipal and agricultural effluents.     

Production and Ecology of Benthic Chironomid Larvae (Diptera) in Lake Hayes,New Zealand,a Warm-monomictic Eutrophic Lake

The distribution and abundance of larval chironomids in Lake Hayes were studied from December 1973 to March 1975. The mean annual production of the two dominant species, Chironomus zealandicus and Chironomus sp. a, was 29.2 g m⁻² dry weight which is approximately 4.3 % of the average annual phytoplankton production in the lake. A high annual P/B ratio of 18.5 is consistent with the multivoltine life cycle of C. zealandicus. Larval chironomid production in the second summer when Anabaena blooms were absent was only one quarter of that in the first summer and is consistent with the hypothesis that the production of benthic chironomids in Lake Hayes is closely linked to that of the phytoplankton through the sedimentation of autochthonous organic matter.     

Increased Benthic Algal Primary Production in Response to the Invasive Zebra Mussel (Dreissena polymorpha) in a Productive Ecosystem, Oneida Lake, New York

Increased water clarity associated with zebra mussel (Dreissena polymorpha) populations may favor benthic algal primary production in freshwater systems previously dominated by pelagic phytoplankton production. While zebra mussel-mediated water clarity effects on benthic primary production have been implicated in published reports, few production estimates are available. This study estimates benthic primary production in Oneida Lake, NY before and after zebra mussel invasion (1992), using measured photosynthetic parameters ( , α^B and β) from sampled benthic algal communities. In the summers of 2003 and 2004, primary production was measured as O₂ evolution from algal communities on hard (cobble) and soft (sediment) substrate from several depths. We also backcast estimates of benthic primary production from measurements of light penetration since 1975. Estimates of whole-lake epipelic and epilithic algal primary production showed a significant (4%) increase and exhibited significantly less interannual variability subsequent to the establishment of zebra mussels. We applied our model to two lakes of differing trophic status; the model significantly overestimated benthic primary production in a hypereutrophic lake, but there was no significant difference between the actual and predicted primary production values in the oligotrophic lake. The hypereutrophic lake had higher zebra mussel densities than Oneida (224 vs. 41 per sample respectively). Though total community respiration (measured in total darkness) was factored into our model predictions of production, our model may need modification when heterotrophic respiration is a large portion of total community metabolism.     

Microbial Food Webs in Marine Sediments. I. Trophic Interactions and Grazing Rates in Two Tidal Flat Communities

Abstract The role of grazing by marine sediment flagellates, ciliates, and meiobenthic animals in controlling production of their bacterial and diatom prey was investigated. Several novel or modified techniques were used to enumerate prey (bacteria and diatoms), measure bacterial production, quantify proto- and micrometazoan predators, and evaluate rates of bacterivory and herbivory. The results indicated that, in a temperate, marine intertidal flat composed of fine sand, colorless nanoflagellates, ciliates, and nematodes were the most important bacterivores. Together, these organisms were responsible for removing up to 53% of bacterial production, by grazing. The observed rates of bacterivory were high enough to hypothesize that periods of grazing control of bacterial production might occur regularly in similar habitats. Colorless microflagellates, ciliates, and nematodes had high rates of diatom consumption. The combined small diatom consumption rate was equivalent to 132% of diatom standing stock per day. Trophic interactions between diatoms and micro- and meiobenthos might be a factor limiting growth of small (around 10 μm) diatoms. In coarse sands of an open beach, all micrograzers except pigmented nanoflagellates were rare, whereas bacterial and diatom assemblages were rather abundant and active. In this type of sediment, the micrograzers were able to consume only a marginal percentage of bacterial production (<1%) and diatom standing stock (3.8%), thus playing a minor role in controlling the dynamics of their prey. Received: 11 June 1996; Accepted: 13 August 1996     

Modelling the importance of sediment bacterial carbon for profundal macroinvertebrates along a lake nutrient gradient

In dimictic, temperate lakes little is known about the quantitative importance of trophic coupling between pelagic and profundal communities. Although it is a generally accepted paradigm that profundal secondary production is dependent on autochthonous pelagic production (primarily diatoms), the importance of interactions between phytodetrital inputs, sediment bacteria, and macroinvertebrates are still not well understood. In this study, we used theoretical models to estimate macroinvertebrate carbon requirement (production + respiration) and bacterial production for lakes of different trophic categories. Comparisons of estimates show that the importance of bacterial production as a carbon source for benthic macroinvertebrates is inversely related to lake trophic state. Assuming that infauna assimilates 50% of ingested bacterial carbon, this food source could account for between 47% (oligotrophic lakes) and 2% (hypertrophic lakes) of their carbon demand. These calculations indicate that bacterial carbon should not be an important C-resource for profundal macroinvertebrates of eutrophic and hypertrophic lakes.     

Benthic Bacterial Production and Protozoan Predation in a Silty Freshwater Environment

The interrelation of heterotrophic bacteria with bacterivorous protists has been widely studied in pelagic environments, but data on benthic habitats, especially in freshwater systems, are still scarce. We present a seasonal study focusing on bacterivory by heterotrophic nanoflagellates (HNF) and ciliates in the silty sediment of a temperate macrophyte-dominated oxbow lake. From January 2001 to February 2002 we monitored the standing stock of bacteria and protozoa, bacterial secondary production (BSP, ³H-thymidine, and ¹⁴C-leucine incorporation), and grazing rates of HNF and ciliates on bacteria (FLB uptake) in the oxic sediment of the investigated system. BSP ranged from 470 to 4050 µg C L^–1 wet sediment h^–1. The bacterial compartment turned out to be highly dynamic, indicated by population doubling times (0.6–10.0 d), which were comparable to those in the water column of the investigated system. Yet, the control mechanisms acting upon the bacterial population led to a relative constancy of bacterial standing stock during a year. Ingestion rates of protozoan grazers were 0–20.0 bacteria HNF^–1 h^–1 and 0–97.6 bacteria ciliate^–1 h^–1. HNF and ciliates together cropped 0–14 (mean 4)% of BSP, indicating that they did not significantly contribute to benthic bacterial mortality during any period of the year. The low impact of protozoan grazing was due to the low numbers of HNF and ciliates in relation to bacteria (1.8–3.5 × 10⁴ bacteria HNF^–1, 0.9–3.1 × 10⁶ bacteria ciliate^–1). Thus, grazing by HNF and ciliates could be ruled out as a parameter regulating bacterial standing stock or production in the sediment of the investigated system, but the factors responsible for the limitation of benthic protistan densities and the fate of benthic BSP remained unclear.     

Composition and Productivity of the Benthic Macroinvertebrate Community of a Subtropical Reservoir

Physical-chemical conditions, phytoplankton productivity, community structure and productivity of the macroinvertebrate benthic community were determined during 1976–77 in a subtropical reservoir. Physical-chemical results revealed high nitrate and phosphate concentrations with highest values in the riverine segment. Large phytoplankton populations were present during most of the year. Phytoplankton productivity was high, producing an annual mean of 87 mg C · m⁻³ · h⁻¹ (12 hours light day). High turbidity in the riverine segment limited phytoplankton productivity during winter and spring. Macrobenthos was dominated by chironomids (Chironomus, Procladius, Coelotanypus and Tanypus) and oligochaetes (Limnodrilus). The annual mean benthic population was estimated at 1,626 · m⁻² with a mean dry weight of 0.66 g · m⁻². Mean benthic species diversity was 1.80. A lacustrine-riverine community gradient was revealed. Benthic productivity was 6.8 g · m⁻² · yr⁻¹ (dry weight) with a P: B ratio of 10. A low correlation was observed between benthic and phytoplankton productivity, and between phytoplankton standing crop and benthic macroinvertebrate numbers throughout the reservoir. Algal food supplies had little impact on the benthic community which was composed predominately of species which fed mostly on organic detritus. Stressful conditions caused by low dissolved oxygen concentrations probably inhibited development of the benthic community throughout the reservoir during summer months, while high sedimentation rates limited development in the head waters.     

Benthic bacterial biomass supported by streamwater dissolved organic matter

Bacterial biomass in surface sediments of a headwater stream was measured as a function of dissolved organic carbon (DOC) flux and temperature. Bacterial biomass was estimated using epifluorescence microscopic counts (EMC) and ATP determinations during exposure to streamwater containing 1,788g DOC/liter and after transfer to groundwater containing 693g DOC/liter. Numbers of bacteria and ATP concentrations averaged 1.36×10⁹ cells and 1,064 ng per gram dry sediment, respectively, under initial DOC exposure. After transfer to low DOC water, biomass estimates dropped by 53 and 55% from EMC and ATP, respectively. The decline to a new steady state occurred within 4 days from ATP assays and within 11 days from EMC measures. A 4°C difference during these exposures had little effect on generation times. The experiment indicated that 27.59 mg/hour of natural DOC supported a steady state bacterial biomass of approximately 10g C/g dry weight of sediment (from EMC determinations). Steady state bacterial biomass estimates on sediments that were previously muffled to remove organic matter were approximately 20-fold lower. The ratio of GTPATP indicated differences in physiological condition or community composition between natural and muffled sediments.     

Bioturbation enhances the aerobic respiration of lake sediments in warming lakes

While lakes occupy less than 2% of the total surface of the Earth, they play a substantial role in global biogeochemical cycles. For instance, shallow lakes are important sites of carbon metabolism. Aerobic respiration is one of the important drivers of the carbon metabolism in lakes. In this context, bioturbation impacts of benthic animals (biological reworking of sediment matrix and ventilation of the sediment) on sediment aerobic respiration have previously been underestimated. Biological activity is likely to change over the course of a year due to seasonal changes of water temperatures. This study uses microcosm experiments to investigate how the impact of bioturbation (by Diptera, Chironomidae larvae) on lake sediment respiration changes when temperatures increase. While at 5°C, respiration in sediments with and without chironomids did not differ, at 30°C sediment respiration in microcosms with 2000 chironomids per m² was 4.9 times higher than in uninhabited sediments. Our results indicate that lake water temperature increases could significantly enhance lake sediment respiration, which allows us to better understand seasonal changes in lake respiration and carbon metabolism as well as the potential impacts of global warming.     

Biogenic Stabilization of Intertidal Sediments: The Importance of Extracellular Polymeric Substances Produced by Benthic Diatoms

The sediment-stabilizing effect of benthic diatoms was investigated in a laboratory setting. Axenic cultures of the benthic diatoms Nitzschia cf. brevissima and Cylindrotheca closterium were inoculated in Petri dishes containing sand and incubated under axenic conditions. By ensuring aseptic routines throughout the experiments, interference from other organisms occurring with diatoms in natural photothrophic biofilms was avoided. This allowed the examination of the role of benthic diatoms in sediment stabilization. Increases in the critical erosion shear stress of the sediment were observed in the presence of both diatom taxa relative to sterile sediment. However, N. cf. brevissima was more effective than C. closterium. Values of critical shear stress in the experimental system were in the same range as those observed in natural biofilms, which indicates that diatoms are important agents for biogenic stabilization. Extracellular carbohydrate contents in the microcosms were similar for both diatom species. However, in the presence of N cf. brevissima, extracellular carbohydrate correlated significantly to critical shear stress, explaining up to 80% of the variation, whereas this was not the case for C. closterium. Therefore, it was concluded that the quantity of extracellular polymeric substances (EPS) alone did not explain the biogenic stabilization. Observed adsorption of EPS to sediment particles depended on the relative amount of uronic acids in the exopolymers. Using fluorescently labeled lectins, confocal laser scanning microscopy showed that EPS secretion by N. cf. brevissima resulted in ordered three-dimensional matrix structures. It is suggested that the structuring of EPS plays an prominent role in the process of biostabilization, and that diatoms such as N. cf. brevissima are actively involved in producing the structure of EPS, whereas others such as C. closterium do not do so to the same extent.     

Microphytobenthic Primary Production and Sedimentary Carbohydrates Along Salinity Gradients in the Lagoons of Grado and Marano (Northern Adriatic Sea)

Primary production of the microphytobenthic community and carbohydrates concentrations were studied in the lagoonal system of Grado and Marano, located in the Northern Adriatic coast. Sediment samples were collected along a salinity gradient. Abundance and species composition of the microphytobenthic communities were analysed and the benthic microalgal biomass was estimated as Chlorophyll a (Chl a). Primary production of benthic diatoms was estimated using ¹⁴C-tracer. Extracellular carbohydrates were extracted from the sediment and separated in two operationally defined fractions (colloidal and EDTA-extractable). Salinity was higher in the Grado lagoon, where the benthic microalgal community was mainly composed of marine diatoms. In the Marano lagoon, which has a lower salinity, freshwater species were also found. In both lagoons, photosynthetic efficiency showed an inverse relationship with salinity and a direct relationship with the main biological variables. Photosynthetic activity was directly related to Chl a and abundance of benthic microalgae, suggesting that in the benthic system microalgal community is responsible for primary production. Overall, salinity was also influent on the microphytobenthic primary production, which was greater in the more saline Grado lagoon.     

Benthic invertebrate recolonization of small-scale disturbances in the littoral zone of a subtropical Florida lake

Construction of spawning beds by the exotic cichlid fish, Sarotherodon aurea, produces numerous small-scale disturbances in shallow Florida lakes. Two experimental methods were used to simulate these disturbances (containers filled with defaunated sediment and disturbed natural substratum) and to determine benthic invertebrate recolonization patterns and rates.Recolonization of experimental containers was rapid and densities of macroinvertebrates reached equilibrium with the control habitat by Day 6 following disturbance. However, Chironomidae and miscellaneous taxa usually were more abundant (p < 0.05) in the experimental containers, and Oligochaeta always were more abundant (p < 0.01) in the control. Species of the planktonically dispersed insects either attained equilibrium rapidly or showed significantly higher densities in the experimental containers (i.e. opportunistic species).In the second experimental approach, disturbance of natural substratum removed 91% of the benthic invertebrates. The common species of chironomids reached equilibrium by Day 6 but the two predominant oligochaetes, Limnodrilus and Pristina, reached equilibrium at 9 and 15 days respectively. For the entire community, similarity on Day 6 was 87.1% and over the remainder of the sample period averaged 86.8%. Chironomid instar analyses showed significantly higher proportions of early instars in the disturbed patch indicating that colonization is principally by younger organisms. Thus, small-scale disturbance in freshwater benthic communities may be more important in ordering the biomass or age-group structure than the numerical abundance.     

Benthic communities on a sandy Ligurian beach (NW Mediterranean)

The different components of the benthic community of a sandy microtidal beach (Arenzano) in Liguria (NW Mediterranean) were investigated during late spring (May) 2002 and 2003. Sampling was carried out in two transects, chosen in order to represent the characteristics of the entire beach and their eventual spatial variations. Each transect included two stations: one placed in the swash zone (SW) and one in the surf zone (SF). Although no significant differences were found in the sediment texture over the 2 years (t-tests p > 0.1 for all the granulometric fractions), notwithstanding an increase in the mean grain size (from 0.8 to 1.1 mm) between the sampling periods, 2002 was characterised by a higher quantity of organic matter (on average 14.4 vs. 3.6 gC m⁻² for the sum of proteins, carbohydrates and lipids) and higher bacterial biomass (on average 1.9 vs. 0.9 gC m⁻²). The metazoan assemblages (meiofauna and macrofauna) were also richer (density = 2.9 × 10⁵ vs. 1.0 × 10⁵ ind. m⁻², biomass = 0.09 vs. 0.03 gC m⁻² for meiofauna; density = 1988 vs. 739 ind. m⁻², biomass = 0.14 vs. 0.03 gC m⁻² for macrofauna) in 2002. A significant quantitative reduction (t-test for proteins, carbohydrates and lipids, at least p = 0.004) in the food supply in 2003 affected the abundance of the metazoans, as confirmed by a multivariate analysis that clearly differentiated the 2 years, and seemed to inhibit their relationships within the benthic food web. The bacterial biomass was always dominant, even under the least favourable trophic conditions, due to the ability of bacteria to adapt to a very harsh environment. Our results suggest that the food supply played an important role in the benthic community structures of the beach during late spring, bacteria being the key organisms within the benthic system. The communities seemed to be bottom-up controlled, while predation seemed to be irrelevant.     

Instar distribution and biomass of Chironomidae larvae in Lago El Junco, Isla San Cristobal, the Galápagos

The distribution, abundance and standing crop biomass of chironomid larvae were determined at one-meter depth intervals along three radial transects. Samples were collected by coring soft sediments while diving. Three genera were found in the lake: Chironomus sp. (collector-filtering larvae), Ablabesmyia sp. (predatory larvae) and Goeldichironomus sp. (collector-filtering larvae). Standing crop densities of chironomids, averaged over the entire lake, varied from 30,594 larvae/m² to 11,428 larvae/m² at different depths. No statistically significant zonation in density was found for the two most common taxa, Chironomus sp. (87.8% of specimens) and Ablabesmyia sp. (9.0%), however the deepest zones (>4 m) had the lowest estimated densities. Significant differences in standing crop biomass were detected, with the 6 m depth having greatest biomass. The increase in standing crop biomass was a function of (1) lower frequency of first instars of Chironomus sp. and Ablabesmyia sp. at 6 m (2) higher average larval biomass of both species at 6 m and (3) very significant increase in average biomass of fourth instars of Chironomus sp. at 6 m compared to fourth instars at shallower depths. These results indicate that the lentic chironomids of this isolated oceanic habitat consist of a small number of species that are ecological generalists and tolerant of low oxygen concentrations.     

The role of chironomids (Diptera) in the profundal macrozoobenthos in Finnish reservoirs

The proundal macrozoobenthos of 12 Finnish inland polyhumic reservoirs was studied by quantitative sampling in 1984 and 1986. Additionally, the succession of the macrozoobenthos in one young reservoir (Lake Kyrkösjärvi) was studied during its first six years. The zoobenthos material was analyzed using the DCA (detrended correspondence analysis) method, and the ordination of sites (i.e. reservoirs), was related to associated environmental data. The role of chironomids was, in general, important in the profundal of the reservoirs. Because of the coarse bottom quality at many sampling sites, many littoral taxa were found in the profundal zone. The genusChironomus dominated (>50% of the total zoobenthic biomass) especially in the acidic, upper-reach reservoirs.Chironomus larvae also dominated in the zoobenthos of Lake Kyrkösjärvi during its first three years. Some of the reservoirs showed similarities with the dystrophic lake type of the lake typology system (Brundin, 1956). Because of the initially eutrophic character of the bottom, many reservoirs belonged to eutrophic lake types. Finnish reservoirs appear to contain a zoobenthic fauna intermediate between natural lakes and bog pools. This theme was discussed from the ecological and evolutionary point of view ofChironomus species.     

The production of five genera of Chironomidae (Diptera) in Lake Norman,a North Carolina reservoir

The production of five genera of chironomids collected at 4- and 8-m depths at four stations in Lake Norman, North Carolina, was estimated using the size-frequency method. To correct for multivoltinism, the estimates were adjusted by multiplying by 365/Cohort Production Interval. Ranges of production estimates (P) in milligrams dry mass m^-2 yr^-1 and production/mean annual biomass ratios (P/B ratios) for each genus over both depths were: Tanytarsus, P = 744 to 8788, P/BB = 66 to 176; Cladotanytarsus, P = 74 to 2387, P/B = 69 to 100; Stempellina, P = 19 to 412, P/ B = 74 to 132; Chironomus, P = 166 to 7293, P/ B = 50 to 70; Cryptochironomus, P = 92 to 398, P/ B = 71 to 221. These estimates are among the highest reported for chironomids, due primarily to the rapid larval development times estimated for these genera.     

The protective role of endogenous bacterial communities in chironomid egg masses and larvae

Insects of the family Chironomidae, also known as chironomids, are distributed worldwide in a variety of water habitats. These insects display a wide range of tolerance toward metals and organic pollutions. Bacterial species known for their ability to degrade toxicants were identified from chironomid egg masses, leading to the hypothesis that bacteria may contribute to the survival of chironomids in polluted environments. To gain a better understanding of the bacterial communities that inhabit chironomids, the endogenous bacteria of egg masses and larvae were studied by 454-pyrosequencing. The microbial community of the egg masses was distinct from that of the larval stage, most likely due to the presence of one dominant bacterial Firmicutes taxon, which consisted of 28% of the total sequence reads from the larvae. This taxon may be an insect symbiont. The bacterial communities of both the egg masses and the larvae were found to include operational taxonomic units, which were closely related to species known as toxicant degraders. Furthermore, various bacterial species with the ability to detoxify metals were isolated from egg masses and larvae. Koch-like postulates were applied to demonstrate that chironomid endogenous bacterial species protect the insect from toxic heavy metals. We conclude that chironomids, which are considered pollution tolerant, are inhabited by stable endogenous bacterial communities that have a role in protecting their hosts from toxicants. This phenomenon, in which bacteria enable the continued existence of their host in hostile environments, may not be restricted only to chironomids.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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