Enhanced exoenzyme activities in sediments in the presence of deposit-feeding Chironomus riparius larvae

1. The combined effects of deposit‐feeding, bioturbation and bioirrigation by benthic macrofauna on the enzymatic hydrolysis of organic matter were studied in microcosms. Chironomus riparius larvae (Insecta, Diptera) served as model macrofauna and stinging nettle leaves (Urtica dioica) were used as a detrital food source. 2. In the upper 10 mm of the sediment (the habitat of C. riparius larvae), the activities of several exoenzymes, the contents of several fractions of particulate organic matter (POM), and the concentrations of dissolved oxidants (O₂, NO) were measured on a small scale. Fluorescent particles (luminophores) were used to quantify the vertical redistribution of particles within the same layer. 3. In control sediment, the addition of detrital food enhanced exoenzyme activities in the 0–2 mm layer only. In the presence of C. riparius larvae, exoenzyme activities increased to 10 mm depth. Further, the content of POM in the 0–2 mm layer was lower in the presence than in the absence of larvae, suggesting ingestion and subduction of the added detritus. After prolonged incubation without further food addition, exoenzyme activities returned close to background values in both treatments, whereas the vertical distribution of POM remained unchanged. 4. The overall penetration depth of O₂ and NO into the sediment was greater in the presence than the absence of C. riparius, the differences being more pronounced after prolonged incubation. Locally high O₂ and NO concentrations due to bioirrigation by C. riparius were measured deep in the sediment. Net downward transport of particles was observed only in the presence of C. riparius larvae and only at the beginning of the incubation. 5. I conclude that deposit‐feeding and bioturbation by macrofauna can quickly remove freshly deposited POM from the sediment surface and transfer it to less oxygenated sites (i.e. animal guts and deep sediment layers). Bioirrigation also increases the availability of oxidants deep in the sediment. The oscillation of oxidant supply to POM particles by ingestion–egestion, burial and re‐burial, and the intermittent bioirrigation of subsurface sediment, is probably the cause of the increased rate of organic matter hydrolysis, the rate‐limiting step in mineralization.     

The arrangement of resources in patchy landscapes: effects on distribution, survival, and resource acquisition of chironomids

The spatial arrangement of resources in patchy habitats influences the distribution of individuals and their ability to acquire resources. We used Chironomus riparius, a ubiquitous aquatic insect that uses leaf particles as an important resource, to ask how the dispersion of resource patches influences the distribution and resource acquisition of mobile individuals in patchy landscapes. Two experiments were conducted in replicated laboratory landscapes (38×38 cm) created by arranging sand and leaf patches in a 5×5 grid so that the leaf patches were either aggregated or uniformly dispersed in the grid. One-day-old C. riparius larvae were introduced into the landscapes in one of three densities (low, medium, high). In experiment 1, we sampled larvae and pupae by coring each patch in each landscape 3, 6, 12, or 24 days after adding larvae. In experiment 2, emerging adults were collected daily for 42 days from each patch in each landscape. In aggregated landscapes, individuals were aggregated in one patch type or the other during a particular developmental stage, but the ”preferred” type changed depending on developmental stage and initial density. Adult emergence was lower by about 30% in all aggregated landscapes. In dispersed landscapes, individuals used both types of patch throughout their life cycles at all initial densities. Thus, patch arrangement influences the distribution of mobile individuals in landscapes, and it influences resource acquisition even when average resource abundance is identical among landscapes. Regardless of patch arrangement, high initial density caused accumulation of early instars in edge patches, 75% mortality of early instars, a 25% increase in development time, and a 60% reduction in adult emergence. Because mortality was extremely high among early-instar larvae in high-density treatments, we do not have direct evidence that the mechanism by which patch arrangement operates is density dependent. However, the results of our experiments strongly suggest that dispersion of resource patches across a landscape reduces local densities by making non-resource patches available for use, thereby reducing intraspecific competition. Received: 20 July 1999 / Accepted: 28 January 2000     

Habitat selection by chironomid larvae: fast growth requires fast food

1. Sediments have been considered as a habitat, a cover from predators and a source of food, but also as a source of potential toxic compounds. Therefore, the choice of a suitable substrate is essential for the development of chironomids. 2. For the midge Chironomus riparius (Meigen 1804) the growth rate of larvae has often been related to the food quality in sediments rather than to the amount of toxicants in the sediment. Both food quality and sediment-bound toxicants have been reported to determine the field distribution of chironomid larvae. 3. We therefore studied the habitat selection by C. riparius larvae of floodplain lake sediments, differing in both food quality and concentrations of sediment-bound toxicants. We offered the different sediments pairwise to the chironomid larvae in a choice experiment and their settlement in the paired sediments was determined after 10 days. 4. It was observed that larvae showed a clear preference for sediments with higher food quality, which also provided better growth conditions, and that the food quality overruled avoidance of the sediments with higher toxicant concentrations. 5. Our observations correspond with the persistence of this fast growing opportunistic chironomid species in organically enriched aquatic ecosystems independent of the contamination level.     

Influence of Chironomus plumosus larvae on ammonium flux and denitrification (measured by the acetylene blockage- and the isotope pairing-technique) in eutrophic lake sediment

Bythotrephes invaded Harp Lake, Ontario, Canada, in the early 1990s. Here we describe seasonal changes in the size, abundance and life history of the invader. The weight of Bythotrephes could be accurately estimated (r² =0.90) from the length of its body and the state of development of its brood, but substantial (22%) corrections for shrinkage in sugarformalin were required. The seasonality of the size and abundance of Bythotrephes was apparently regulated by the interaction of fish predation, temperature-mediated growth, the availability of the invader's prey, and switches in its reproductive biology. The vernal increase in the growth of the Bythotrephes population lagged behind increases in water temperature and abundances of preferred prey, most probably because of predation by lake herring (Coregonus artedi). Bythotrephes abundance increased rapidly in early July; however, the population maximum was brief. After the abundance of their preferred prey crashed in mid-July, Bythotrephes body size declined, and adult females switched from parthenogenic to resting egg production. Population size declined in consequence. In the summer, female Bythotrephes committed most of their energy to growth between instar I and II, but to egg production for instars II and III. The biology of the Harp Lake population differed from that observed in other recently invaded lakes in several ways – the lag in population growth in the spring, the small change in body size between instars II and III, the decline in body size in August, and the co-incident switch by mature females to resting egg production. These inter-lake differences in biology indicate we have more to learn about Bythotrephes before we can predict its influence on the inland lakes of North America.     

The effect of tubificids and chironomids on particle redistribution of lake sediment

The effects of chironomid larvae,Chironomus plumosus, and tubificid worms,Limnodrilus spp., on particle redistribution in lake sediment were investigated experimentally using pots containing sediments obtained from Lake Suwa, Japan. The chironomids and tubificids increased the water content of surface sediment. The chironomid larvae had no effect on particle size distribution, while tubificids continuously accumulated small particles on the surface sediment through their selective feeding activity. Particles larger than 0.125 mm were buried at a sediment depth of 6 cm. In Lake Suwa, long diatom frustules, large plant debris and blue-green algal flocs were found to accumulate in the deeper layer of the lake sediment inhabited by tubificids at high density.     

大型底栖动物对河口沉积物的扰动作用

利用微宇宙示踪技术,定量研究了天津北塘河口地区的天津厚蟹、沙蚕、泥螺和青蛤4种底栖动物对沉积物的扰动作用.结果表明：生物扰动对沉积物的分布具有明显影响.不同底栖动物扰动能力不同,对表层沉积物的扰动强度（扩散系数,10^-3 cm²·d^-1）大小依次为：沙蚕（2.95）> 天津厚蟹（1.00）> 青蛤（0.78）> 泥螺（0.35）> 对照（0.05）;对深层沉积物则是天津厚蟹（3.10）>沙蚕（2.33）>青蛤（0.28）>泥螺（0.15）> 对照（0.05）.这种差异主要与不同底栖动物所属的功能群不同有关.底栖动物体积与表层沉积物扩散系数显著相关（P<0.05）,可采用底栖动物体积预测其对沉积物的扰动强度.     

Manipulation of habitat quality: effects on chironomid life history traits

1. To understand how habitat heterogeneity affects organisms, the present authors measured the response of chironomid life-histories to single patch types in a controlled laboratory experiment. The objective was to determine whether the size and type of leaf particles in the substratum affect development rate and survival of larvae, and the fecundity of the resulting adults.
2. Chironomus riparius larvae were raised in sand supplemented with dogwood, Cornus florida, (a rapid decomposer) or American sycamore, Platanus occidentalis, (a refractory decomposer) leaves fully crossed in a factorial design with four leaf particle sizes (0.2, 0.5, 1 and 5 cm²), or in sand only.
3. Development time was shorter in dogwood habitats than sycamore habitats, and shorter in sycamore habitats than in sand habitats. Survival was higher and larval head capsule lengths of instars III and IV were larger in dogwood than in sycamore or sand habitats. Development time was longer and head capsule lengths of instars III and IV were smaller in habitats with 5-cm² leaf particles than in 0.2-, 0.5- and 1-cm² particle size treatments. Female thorax length (a measure of fecundity) did not differ among treatments.
4. The present authors conclude that, for C. riparius confined to a single patch type, fitness is significantly enhanced when the patch contains small particles of dogwood (labile) leaves.     

The rates of sediment-water exchange of oxygen and sediment bioturbation in Lough Neagh,Northern Ireland

Oxygen is transported 30 mm into the sediment at an 8 m depth site in eutrophic Lough Neagh by the irrigational activities of the benthic fauna. Faunal activity also mixes the upper 20 mm of sediment. Sediment oxygen uptake rate, redox potential-depth profile and the chlorophylla concentration were measured in the upper sediment layers from February to November 1979. Chlorophylla input to the sediment, following the Spring phytoplankton maximum, remained in the 0–1 cm sediment layer but did cause the redox potential profile to change from one with potentials around 400 mV in the upper 50 mm to one with a strong gradient over the 0–30 mm region. The start of benthic faunal activity in May caused the chlorophylla to be mixed into the 1–2 cm layer and also caused oxygen to be transported into the sediment at a rate sufficient to change the redox potential back to its initial state. The biodiffusion coefficient for solids in the upper 20 mm was estimated to be 6 × 10⁻⁸ cm² s⁻¹. Oxygen transport in the pore, waters of the upper sediment layers was considered to be best described as advection, caused by the irrigational activities of the benthic fauna.     

水丝蚓对太湖沉积物有机磷组成及垂向分布的影响

以太湖常见底栖动物——水丝蚓为研究对象,借助室内流动培养装置和31P核磁共振技术(phosphorus-31 nuclearmagnetic resonance spectroscopy)(31P-NMR),研究生物扰动对太湖沉积物有机磷组成及垂向分布的影响。结果表明:短时间内水丝蚓扰动对沉积物有机磷化合物种类组成影响并不显著,但会引起上层沉积物中稳定性较差的有机磷化合物磷脂和DNA含量出现显著降低,同时沉积物中总磷和有机磷的垂向分布亦发生明显改变。此外,水丝蚓扰动下沉积物含水率、孔隙率和碱性磷酸酶活性显著增加,使活性较高的有机磷化合物分解加速,最终导致表层沉积物中的磷脂与DNA含量降低。     

The influence of lime and biological activity on sediment pH,redox and phosphorous dynamics

The addition of powdered limestone to intact sediment cores from oligotrophic, acid Lake Hovvatn caused pH to increase, redox potential (E₇) to drop, and permitted net precipitation of phosphorous (P) from the water column. Significant pH increase was found to a sediment depth of 6 cm and a maximum increase in pH from 4.9 to 6.5 was found at a depth of 0.5 cm when dosed with 36 g m^–2 of lime. Such pH increase creates important changes in sediment equilibrium chemistry and enhances habitat suitability. In the case of Hovvatn, however, sediments would consume only 5 kg of the 91 tons of applied limestone. Superficial sediments remained oxidized, but below 0.5 cm, E₇ in limed sediment declined significantly more than in unlimed sediments, with a maximum difference of 102 mV versus –66 mV at a depth of 6 cm in unlimed and limed cores, respectively. Abiotic reactions account for 82 ± 54% of this reduction and the remainder is due to the oxidation of organic matter by bacteria. Precipitation of CaSO₄, reduction of the sediments by organic compounds at elevated pH and inhibition of the downward diffusion of O₂ by the limestone powder are potential abiotic mechanisms which could drive E₇ down. Enhanced P release was not found at lowered E₇, and supernatent TP concentrations dropped from 11.7 to 4.4 µg P l^–1. More P was swept from solution in cores which recieved larger lime doses. The presence of chironomids caused sediment pH to increase by as much as 1.2 pH units, presumably due to NH₄ release, reduced sediment E₇ by as much as 171 mV and facilitated TP release during the first 17 d of core incubation. Field measurements of vertical distributions of sediment pH and E₇ before and after the liming of Hovvtn corroborated laboratory findings.     

Invertebrate bioturbation can reduce the clogging of sediment: an experimental study using infiltration sediment columns

1. Invertebrate bioturbation can strongly affect water‐sediment exchanges in aquatic ecosystems. The objective of this study was to quantify the influence of invertebrates on the physical characteristics of an infiltration system clogged with fine sediment. 2. Two taxa (chironomids and tubificids) with different bioturbation activities were studied in experimental slow infiltration columns filled with sand and gravel and clogged with a 2 cm layer of fine sediment at the surface. We measured the effects of each taxon separately and combined on hydraulic head, water mobility and sediment reworking. 3. The results showed that invertebrates could reduce sediment clogging and this effect was linked to the functional mode of bioturbation of each group. Tubificid worms dug networks of galleries in the fine sediment, creating pathways for water flow, which reduced the clogging of sediment. In contrast, the U‐shaped tubes of chironomids were restricted to the superficial layer of fine sediments and did not modify the hydraulic conductivity of experimental columns. The combination of invertebrates did not show any interactive effects between tubificids and chironomids. The occurrence of 80 tubificids in the combination was enough to maintain the same hydraulic conductivity that 160 worms did in monospecific treatment. 4. The invertebrates like tubificid worms can have a great benefit on functioning of clogged interfaces by maintaining high hydraulic conductivity, which contributes to increased water‐sediment exchanges and stimulates biogeochemical and microbial processes occurring in river sediments.     

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 influence of chironomus plumosus larvae on the exchange of dissolved substances between sediment and water

4th instar Chironomus plumosus larvae (about 1000·m^–2) were added to tubes containing sediment and overlying water. At a temperature of 20°C the larvae greatly increased the trasnport of silica, phosphorus and iron from the sediment to the water. Oxygen concentrations did not influence the exchange of silica. For two non-calcareous sediments the exchange of phosphorus and iron was much higher under anaerobic than under aerobic conditions while the difference was small for sediment from a hardwater lake. Exchange of inorganic nitrogen was little influenced by added chironomid larvae.     

Effect of cadmium exposure on the globin protein expression in 4th instar larvae of Chironomus riparius Mg. (Diptera: Chironomidae): An ecotoxicoproteomics approach

In order to identify Chironomus hemoglobin (Hb) as a biomarker of ecotoxicity monitoring; herein, the effects of cadmium chloride (Cd) on Hb parameters were investigated in the 4th instar larvae of Chironomus riparius. The expressions of globin mRNA and hemolymph protein, using ecotoxicoproteomic approach, were investigated. Conventional ecotoxicity tests were also conducted to validate the ecotoxicological relevance of the response of Chironomus Hb as a biomarker. The proteomic analysis indicated that exposure to Cd lead alteration in the expression of hemolymph protein, with the total expressions of 12 hemolymph protein spots decreasing in response to treatment, with that of two increasing in response to Cd exposure. In addition, all of the spots differentially expressed in response to Cd treatment were identified as globin proteins. The decreased total Hb content observed in the hemolymph of larvae exposed to Cd suggested that the decreased expression of selected globin proteins in response to Cd exposure impacted on Hb synthesis. The overall results suggested that Hb could be a target molecule for exposure to Cd in C. riparius, with a proteomic approach appearing to be an ideal tool for the discovery of biomarkers in ecotoxicological research.     

The impact of Chironomus plumosus larvae on organic matter decay and nutrient (N, P) exchange in a shallow eutrophic lake sediment following a phytoplankton sedimentation

The importance of Chironomus plumosus larvae onbenthic metabolism and nutrient exchange across thesediment–water interface was evaluated in a shalloweutrophic lake (Lake Arreskov, Denmark) following aphytoplankton sedimentation. Chironomus plumosuslarvae were added to laboratory sediment microcosms,corresponding to a density of 2825 larvae m⁻².Non-inhabited microcosms served as controls. Asedimentation pulse of organic matter was simulated byadding fresh algal material (Chlamydomonasreinhardii) to sediment cores (36 g dryweight m⁻²). The mineralization was followed bymeasuring fluxes of O₂, CO₂, dissolvedinorganic nitrogen and phosphate. A rapid clearance ofalgae from the water column in faunated microcosmssuggested that chironomids may be of major importancein controlling phytoplankton concentrations in shalloweutrophic lakes. Chironomids increased the sedimentO₂ uptake ≈ 3 times more than what wouldbe expected from their own respiration, indicating astimulation of microbial activity and decomposition oforganic matter in the sediment. Addition of algaeenhanced the release of CO₂, NH⁺ ₄ ando-P. The excess inorganic C, N and P released inamended non-inhabited sediment after 36 dayscorresponded to 65, 31 and 58% of the C, N and P inthe added algae. In sediment inhabited by Chironomus plumosus the corresponding numbers were147, 45 and 73%, indicating that mineralization oforganic matter also from the indigenous sediment poolwas stimulated by chironomids. This revised version was published online in July 2006 with corrections to the Cover Date.     

Land use effects on sediment yield and quality

Attempts to establish land use effects on sediment yield and water quality require consideration of (1) background biogeochemical cycling, (2) quantitative land use mapping, (3) selection of appropriate transport and attenuation models and (4) assessment of critical or threshold amounts, and types of land use change. Progress in each of these problem areas is considered.     

The relationship between sediment mobilisation and the entry of Baetis mayflies into the water column in a laboratory flume

Mass bedload movement is thought to play a key role in initiating stream invertebrate drift during extreme flood events. However, little is known of the importance of the shear of invertebrates from stone surfaces relative to their entrainment along with bed material at different discharges. In particular, it is unclear whether so-called catastrophic drift only occurs once mass bedload movement, and hence entrainment of invertebrates, occurs. We investigated the relationship between the mobilisation and transport of bed sediments and the entry of Baetis mayflies into the water column in a laboratory flume. Experiments quantified the percentage of Baetis drifting at a range of discharges that mobilised between 0 and 95% of the flume-bed sediments. Control experiments quantified drift losses from sediment fixed to the bed of the flume, such that sediments were immobile even at the highest discharges. Drift losses increased with increasing discharge and velocity in the flume. Sediment mobility contributed significantly to drift (ANCOVA, p < 0.001), with consistently greater drift losses in mobile sediment experiments than in those with fixed sediment. The discharge which resulted in a loss of 100% of Baetis from the mobile sediment bed (discharge 30 l s^–1) resulted in a loss of approximately 50% of individuals from the fixed bed. Results indicate that once bed sediments are mobilised, entry of Baetis into the drift is greater than expected from the shear of animals from stone surfaces alone. Thus, entrainment of animals along with sediment contributes significantly to drift at high flows. This implies that differences in bed stability between sites or streams, or temporal changes in sediment characteristics within a site, could influence patterns of drift.     

A New Approach for the Modelling of Sediment Reworking Induced by a Macrobenthic Community

A new model of bioturbation has been developed to describe short term sediment reworking induced by macrobenthic communities. The design of the model had to consider the mixing processes, firstly, at the organism level, and secondly, at community level. This paper describes the mixing mode of the four types of bioturbators defined by the authors: the biodiffusors, the upward-conveyors, the downward-conveyors and the regenerators. The mathematical formulation of these sub-models consists of ordinary differential equations. They take into account the size of the bioturbated zone, the output fluxes to the water column, tracer decay, physical mixing due to local currents and the type and intensity of the bioturbation processes. These sub-models make it possible to describe correctly the mixing events that have occurred in cores with each type of bioturbator. They also provide the basis for general bioturbation model, that will take into account the respective degrees of involvement of (i) the different bioturbation processes and their characteristics, (ii) the interference between the different processes, and (iii) make possible to predict the particle reworking in order to include it in studies of organic matter in early diagenesis. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of bioturbation on denitrification in a marine sediment from the West Mediterranean littoral

Thein vitro effect ofNereis diversicolor on denitrification has been studied in PVC tubes filled with a coastal marine sediment defaunated by sieving. The first aim of the experiment was to determine the effect of sediment defaunation on denitrification (denitrifying population and Denitrifying Enzyme Assays). Sieving induced a loss of 70% of the initial DEA. The number of denitrifying bacteria was 10 times lower than inin situ sediment. In the top two centimetres, the DEA rose by 75% of its initial value, after 82 days. Polychaetes were only added after a return to near pre-disturbance levels to ensure that our data on the effects of their addition would not be disturbed by changes in the sediment. Introduction of Polychaetes increased the denitrifying population and DEA in the first layer (0–2 cm) of the sediment after 15 days. After 45 days, the surface of the polychaete burrows in sediment was 1.3 to 1.5 times higher than after 15 days, resulting in an increase in solute exchange between seawater and the top layer of sediment. An inhibitory effect of oxygen on denitrification was detected in the uppermost layer only.     

生物扰动对沉积物中污染物环境行为的影响研究进展

生物扰动由于显著改变沉积物结构和性质,进而影响沉积物中污染物的环境行为。综述生物扰动对沉积物中氮、磷、重金属和疏水性有机污染物环境行为的影响。生物扰动促进这些污染物从沉积物向水体释放。生物扰动还对不同的污染物产生其它不同的影响。对于氮,生物扰动还影响其硝化与反硝化作用;对于磷,生物扰动不仅改变其化学形态,还提高有机磷降解。对于重金属,生物扰动还能改变其在沉积物中的分布及化学形态。对于疏水性有机污染物,生物扰动主要增强生物富集和代谢,以及提高生物降解。