Comparing the responses of diatoms and macro- invertebrates to metals in upland streams of Wales and Cornwall

SUMMARY 1. Surprisingly few data compare the apparent responses of diatoms and macroinvertebrates to metals in streams. We examined variation in metals, diatoms and macroinvertebrates between 51 streams in metal‐mining areas of Wales and Cornwall, U.K., using a survey design with multiple reference and polluted sites. 2. To quantify variations in metals between sites, we calculated cumulative criterion unit (CCU) scores, a recently defined measure of total stream metal concentration and toxicity, to account for additive effects of each metal relative to putative toxic thresholds. We compared assemblage responses among epilithic diatoms and macroinvertebrates to CCU scores or individual metal concentrations using correlation and detrended correspondence analysis (DCA). 3. Macroinvertebrate diversity, richness and total abundance declined and evenness increased with increasing copper concentrations. Trends with CCU scores were significant but less pronounced. Some individual macroinvertebrate taxa varied significantly in abundance with CCU scores, copper or zinc, but overall assemblage composition correlated only with manganese, pH and nitrate. 4. Among diatoms, pH and conductivity explained the major variations in assemblage composition, and neither diversity, richness nor evenness varied with metal concentration. Nevertheless, the single strongest predictor of diatom assemblages on ordination axis 2 was the CCU score. The abundances of some macroinvertebrate taxa, particularly grazers, also explained significant variations in diatom assemblages that were linked to both metals and acid–base status. 5. Diatom species apparently tolerant of high metal concentrations included Psammothidium helveticum, Eunotia subarcuatoides, Pinnularia subcapitata and Sellaphora seminulum. Of these, P. helveticum, E. subarcuatoides and P. subcapitata were abundant at lower pH than S.seminulum and might indicate metal enrichment over different pH ranges. Sensitive species included Fragilaria capucina var. rumpens, Achnanthes oblongella and Tabellaria flocculosa. 6. We conclude that macroinvertebrates at these sites reflected metal pollution most strongly through variations in diversity while effects on diatoms were best reflected by changes in assemblage composition. We suggest that, with further refinement, CCU scores might be useful in evaluating the possible effects of metal pollution on benthic organisms in European rivers.     

Long-term effects of catchment liming on invertebrates in upland streams

1. Catchment liming to mitigate acidification causes major chemical change in freshwaters but longer‐term effects are poorly understood. Using a replicated basin‐scale experiment with a multiple BACI design (= before‐after‐control‐impact), we assessed chemical and biological effects for 10 years after the catchments of three acidified Welsh streams at Llyn Brianne were limed in 1987/88. 2. Stream chemistry was measured weekly to monthly, and macroinvertebrates monitored annually, between 1985 and 1998. Biological change through time was assessed from the abundance and taxon richness of invertebrates. We paid particular attention to 18 species known to be acid‐sensitive. The effects of liming were assessed by comparing chemical and biological trends among the three replicate limed streams, three acid reference streams and two naturally circumneutral streams. 3. Following single lime applications, acid‐base chemistry in treated streams changed significantly. High mean pH (> 6), increased calcium (> 2.5 mg L^?1) and low aluminium (< 0.1 mg L^?1) persisted throughout the 10 years following liming. 4. The effects of liming on invertebrates were modest. Acid sensitive taxa increased significantly in abundance in limed streams, but only during 2 years following treatment. Significant effects on richness were more sustained, but on average added only 2–3 acid‐sensitive species to the treated streams, roughly one‐third of their average richness in adjacent circumneutral streams. Only the mayfly Baetis rhodani and the stonefly Brachyptera risi occurred significantly more often in limed streams after treatment than before it. 5. Despite these modest long‐term effects on invertebrates, nearly 80% of the total pool of acid‐sensitive species has occurred at least once in the limed streams in the 10 years since treatment. This pattern of occurrence suggests that the colonization of limed streams by acid‐sensitive taxa reflects limited persistence rather than restricted dispersal. We present evidence to show that episodes of low pH continued to affect acid‐sensitive taxa even after liming. We highlight the importance of extending the time‐periods over which the effects of large‐scale ecological experiments are assessed.     

Exploitation of prey by a river bird, the dipper Cinclus cinclus (L.), along acidic and circumneutral streams in upland Wales

SUMMARY. 1. The diet of the Eurasian dipper Cinclus cinclus. a riverine bird from a globally widespread genus, was assessed through all stages of its annual cycle using published data and field studies from streams of contrasting chemistry. Time-activity budgets were also compiled throughout the year and used to estimate annual energy requirements from the stream ecosystem. 2. The annual energy requirements for a territorial pair ranged from 148,000 to 158,000 kJ yr^?1 depending on whether one or two broods were reared. After allowing for assimilation efficiency, these requirements were estimated to represent 10.5–11.0 kg dry mass of fish and invertebrates. Using representative values for territory size (4680–11,250 m²), annual exploitation of secondary production was estimated at 0.93–2.35 g dry mass m^?2. 3. Several features combined to focus the predatory load on certain organisms over different stages of the annual cycle. These included the availability and selection of alternative prey, the need to provision nestlings with large items such as trichopteran larvae, and the use by females of calcium-rich prey such as fish prior to egg formation. 4. Across their range of territory size, annual exploitation (dry mass) by dippers was estimated at 0.06–0.29 g m^?2 for Plecoptera. 0.02–0.22 g m^?2 for Ephemeroptera. 0.59–1.11 g m^?2 for Trichoptera and 0–0.78 g m^?2 for fish. Exploitation of Ephemeroptera. I richoptera and fish were all highest in circumneutral streams, but exploitation of Plecoptera was highest in acidic streams because other prey were scarce. 5. Cottids dominated the fish component of the diet. Influences on their density could be substantial according to available data on production in Welsh streams. Hydropsychids and limnephilids dominated the trichopteran component, with exploitation again representing potentially substantial amounts of production. Contemporaneous data are required on benthic production and exploitation by the birds. 6. The ecological role of birds in rivers particularly, and aquatic ecosystems generally, is currently neglected but worthy of considerable research effort.     

The effects of catchment liming on the chemistry and biology of upland Welsh streams: testing model predictions

1. The catchments of three acidified streams in mid Wales were limed in 1987/88. Here we assess their chemical and biological response in comparison with unmanipulated reference streams over a period of 5 years post-liming. 2. Stream chemistry was measured weekly/fortnightly between 1985 and 1992, while macroinvertebrates were sampled annually. Colonization by acid-sensitive taxa was assessed and trends in community structure were monitored by TWINSPAN. Real biological responses were compared with those predicted by an empirical model constructed using chemical data. 3. There were marked changes in stream chemistry following liming: calcium concentrations and pH values increased, while aluminium concentrations decreased to levels similar to those in naturally circumneutral streams. These conditions have persisted since liming. 4. Empirical models predicted that stream invertebrates would respond to the altered stream chemistry, with the establishment of communities typical of circumneutral conditions. 5. Following liming, the taxon richness and abundance of acid-sensitive taxa was significantly higher in limed compared with reference streams. Colonization by, and persistence of acid-sensitive taxa was patchy, however, and richness was still substantially lower than in naturally circumneutral streams. Moreover, contrary to the model predictions, there were no wholesale changes in the structure of macroinvertebrate communities. 6. We conclude that liming has created and maintained chemical conditions suitable for macroinvertebrate communities typical of circumneutral streams, but these chemical changes have not been matched by sustained responses among the biota. The views expressed in this article are those of the authors and do not necessarily reflect those of the NRA.     

Macroinvertebrate colonization of stones in two upland southern Australian streams

Experiments on the colonization by macroinvertebrates of stones were performed in the Acheron and Toorongo Rivers, two upland rivers in south-eastern Australia. The three experiments were each about one month in duration. In the first experiment — Acheron I — and the third experiment — Toorongo — river stones were used. In the Acheron II experiment both river stones and quarry stones were used.The number of colonizing species with time in the Acheron I and Toorongo experiments fitted a logarithmic model with the colonization and elimination (extinction) rates both declining with time. The population densities of the total animals and of most of the common species fitted the power model (linear model on logarithmically transformed axes). In the Acheron II experiment the major difference between the colonization of river stones and quarry stones was the low numbers of scrapers on the quarry stones. The colonization of the stones in Acheron I and Toorongo by the various functional feeding groups is described. The colonization of stones in streams is a rapid process suggesting that the stream stone assemblage has a high resilience — an attribute of deterministic communities.     

Predicting local biological characteristics in streams: a comparison of landscape classifications

1. Landscape classifications group tracts of land based on similar physico‐chemical attributes that may affect the biological characteristics of streams at local scales. We tested the ability of five landscape models to account for variation in algal and macroinvertebrate biomass, brook trout (Salvelinus fontinalis) growth and macroinvertebrate community composition from 132 riffles in 15 catchments in the Big Horn Mountains, Wyoming. 2. A model created by the U.S. Forest Service (FS) combined catchment and ecoregion approaches to classify the landscape. Our model used digital elevations to create a landscape classification for streams (DEM). The last three models were based on: (1) standard ecoregions (Ecoregion), (2) the type of underlying bedrock (Geology) and (3) the geographical distance between sites (Site Proximity model). 3. Overall, the Ecoregion and Geology models performed better than the two catchment models (FS and DEM) in predicting local biological characteristics. The Geology model was best at predicting differences in algal and macroinvertebrate biomass, the Site Proximity and Ecoregion models were best at predicting patterns of similarity in macroinvertebrate community composition, and the Site Proximity, Ecoregion, Geology, and FS models, in order from best to worst, accounted for significant variation in brook trout growth. The Site Proximity model performed well because of the effects of spatial autocorrelation. The DEM was consistently one of the worst models at predicting local biological characteristics because it failed to include important attributes (e.g. dominant geology). Calcareous geology was positively associated with greater macroinvertebrate biomass and faster brook trout growth, but it was inversely related to algal biomass. 4. None of the models accounted for a large amount of variation in local biological characteristics. Single‐scale, landscape classifications may never accurately predict variation in local biological characteristics because: (1) landscapes show a high degree of spatial heterogeneity, (2) local effects are stronger than landscape attributes and (3) there are too many intervening levels between landscape and local scales in the nested hierarchy of streams. However, landscape classifications did account for significant variation in biological characteristics. Thus, they would be a valuable management tool as part of a multi‐scale, hierarchical technique for classifying stream ecosystems.     

Thermal adaptation in biological membranes: interacting effects of temperature and pH

Summary The interacting effects of pH and temperature on membrane fluidity were studied in plasma membranes isolated from liver of rainbow trout (Oncorhynchus mykiss) acclimated to 5 and 20°C. Fluidity was determined as a function of temperature under conditions of both constant (in potassium phosphate buffer) and variable pH (in imidazole buffer, consistent with imidazole alphastat regulation) from the fluorescence anisotropy of two probes: 1,6-diphenyl-1,3,5-hexatriene, which intercalates into the bilayer interior, and 1-(4-trimethylammoniumphenyl)-6-phenyl-1,3,5-hexatriene which is anchored at the membrane/water interface. The temperature dependence of the anisotropy parameter for 1,6-diphenyl-1,3,5-hexatriene in plasma membranes of 20°C-acclimated trout was greater when determined in phosphate (AP per °C=-0.047) than in imidazole buffer (AP per °C=-0.022); similar, but less significant, trends were noted with 1-(4-trimethylammoniumphenyl)-6-phenyl-1,3,5-hexatriene. In contrast, the temperature dependence of fluidity (AP/°C in the range-0.0222 to-0.027) did not vary with buffer composition in membranes of 5°C-acclimated trout. In phosphate buffer, anisotropy parameter values for 1,6-diphenyl-1,3,5-hexatriene were significantly lower in 5°C-than 20°C-acclimated trout, indicating a less restricted probe environment following cold acclimation and nearly perfect compensation (91%) of fluidity. Temperature-dependent patterns of acid-base regulation were estimated to account for 11–40% of the fluidization evident in membranes of 5°C-trout, but a period of cold acclimation was required for complete fluidity compensation. In contrast, no homeoviscous adaptation was evident in imidazole buffer, indicating that membrane fluidity is sensitive to buffer composition. Accordingly, vesicles of bovine brain phosphatidylcholine, suspensions of triolein, and plasma membranes of 5°C-acclimated trout were consistently more fluid in imidazole than phosphate buffer. Membranes of 5°C-acclimated trout were enriched in molecular species of phosphatidylcholine containing 22:6n3 (at the expense of species containing 18:1n9 and 18:2n6) compared to membranes of 20°C-trout; consequently, the unsaturation index was significantly higher (3.29 versus 2.73) in trout maintained at 5 as opposed to 20°C. It is concluded that: 1) the chemical composition of the internal milieu can significantly influence the physical properties of membrane lipids; 2) temperature-dependent patterns of intracellular pH regulation may partially offset the ordering effect of low temperature on membrane fluidity in 20°C-acclimated trout transferred to 5°C, but not in 5°C-acclimated trout transferred to warmer temperatures; 3) the majority of the thermal compensation of plasma membrane fluidity resulting from a period of temperature acclimation most likely reflects differences in membrane composition between acclimation groups; 4) imidazole apparently interacts with trout hepatocyte plasma membranes in a unique way.Abbreviations _im netcharge stateofproteins - AP anisotropyparameter - bw body weight - DPH 1,6-diphenyl-1,3,5-hexatriene - HEPES N-2-hydroxyethylpiperazine-N-2-ethanesulphonicacid - PC phosphatidylcholine - pH_e pHofarterial blood - pH_i intracellular pH - TMA-DPH 1-(4-trimethylammoniumphenyl)-6-phenyl-1,3,5-hexatriene - TRIS tris(hydroxymethyl)aminomethane     

The distribution of salmonids in upland streams in relation to depth and gradient

The distribution of various age classes of salmon and trout was assessed in upland streams by electrofishing. Water depths and site gradients were measured and correlated to fish densities. The fry of both species were significantly more abundant in shallow water; up to 75·3% of salmon fry and 72·2% of trout fry were captured in sites of mean depth < 20 cm. Older trout were found mainly in the deeper areas, with a maximum of 7·4% captured in sites < 20 cm mean depth. Yearling fish were found in all the depth-ranges sampled, but with a tendency for higher numbers in mid-range depths. There were similar correlations in the abundance of each age class with the actual areas of shallow, mid-range and deep water habitat available within sites. Correlations of fish density with gradient indicated that trout were limited in their distribution to areas of lower flow, whereas salmon were not. Since depth and gradient were significantly negatively correlated, there was an apparent preference of trout for slightly deeper habitats than the equivalent year classes of salmon. The observed habitat segregation is discussed in terms of competition and selection.     

The independent and interactive effects of fine sediment and flow on benthic invertebrate communities characteristic of small upland streams

SUMMARY 1. Floods are an important mechanism of disturbance operating in streams that can markedly influence the abundance and diversity of benthic fauna. In upland streams many studies cite the scouring effects of fine transported sediments as a potential source of disturbance to the biota during spates, but few studies have sought to test this hypothesis critically.
2. Here we used a series of eight artificial streams to test whether high suspended-sediment concentrations influenced the short-term response of benthic invertebrate fauna to increases in flow. In an experiment designed to simulate a small spate, flow and sediment loads were each manipulated to examine their independent and interactive effects. Benthic invertebrates were sampled before and after the manipulation, and drift samples were taken at regular intervals during the experiment. The experiment was repeated twice, once at the end of winter (June, 1998), and once in summer (February, 1999).
3. Flow increases caused large increases in the number and diversity of drifting animals, and significant declines in the numbers and diversity of organisms found in benthic samples, but these declines were apparently not affected by the addition of fine sediment. The addition of sediment alone had little effect on the fauna. These results were consistent across both experiments.
4. The results suggest that flow increases alone can disturb benthic fauna, and that neither substrate movement nor suspended sediment increases are necessary for floods and spates to disturb the benthic assemblage. However, as argued elsewhere, the effects of flow increases are likely to be contingent upon the presence or absence of local flow refugia, which can allow animals to escape the shear forces that would otherwise remove them from the surface or interstitial areas of the streambed.     

Contrasting effects of anthropogenic and natural acidity in streams: a meta-analysis

Large-scale human activities including the extensive combustion of fossil fuels have caused acidification of freshwater systems on a continental scale, resulting in reduced species diversity and, in some instances, impaired ecological functioning. In regions where acidity is natural, however, species diversity and functioning seem to be less affected. This contrasting response is likely to have more than one explanation including the possibility of adaptation in organisms exposed to natural acidity over evolutionary time scales and differential toxicity due to dissimilarities in water chemistry other than pH. However, empirical evidence supporting these hypotheses is equivocal. Partly, this is because previous research has mainly been conducted at relatively small geographical scales, and information on ecological functioning in this context is generally scarce. Our goal was to test whether anthropogenic acidity has stronger negative effects on species diversity and ecological functioning than natural acidity. Using a meta-analytic approach based on 60 datasets, we show that macroinvertebrate species richness and the decomposition of leaf litter -- an important process in small streams -- tend to decrease with increasing acidity across regions and across both the acidity categories. Macroinvertebrate species richness, however, declines three times more rapidly with increasing acidity where it is anthropogenic than where it is natural, in agreement with the adaptation hypothesis and the hypothesis of differences in water chemistry. By contrast, the loss in ecological functioning differs little between the categories, probably because increases in the biomass of taxa remaining at low pH compensate for losses in functionality that would otherwise accompany losses of taxa from acidic systems. This example from freshwater acidification illustrates how natural and anthropogenic stressors can differ markedly in their effects on species diversity and one aspect of ecological functioning.     

Nanomaterials in biological environment: a review of computer modelling studies

Nanotechnology is set to impact a vast range of fields, including computer science, materials technology, engineering/manufacturing and medicine. As nanotechnology grows so does exposure to nanostructured materials, thus investigation of the effects of nanomaterials on biological systems is paramount. Computational techniques can allow investigation of these systems at the nanoscale, providing insight into otherwise unexaminable properties, related to both the intentional and unintentional effects of nanomaterials. Herein, we review the current literature involving computational modelling of nanoparticles and biological systems. This literature has highlighted the common modes in which nanostructured materials interact with biological molecules such as membranes, peptides/proteins and DNA. Hydrophobic interactions are the most favoured, with π-stacking of the aromatic side-chains common when binding to a carbonaceous nanoparticle or surface. van der Waals forces are found to dominate in the insertion process of DNA molecules into carbon nanotubes. Generally, nanoparticles have been observed to disrupt the tertiary structure of proteins due to the curvature and atomic arrangement of the particle surface. Many hydrophobic nanoparticles are found to be able to transverse a lipid membrane, with some nanoparticles even causing mechanical damage to the membrane, thus potentially leading to cytotoxic effects. Current computational techniques have revealed how some nanoparticles interact with biological systems. However, further research is required to determine both useful applications and possible cytotoxic effects that nanoparticles may have on DNA, protein and membrane structure and function within biosystems.     

Classification and ordination of macroinvertebrate assemblages to predict stream acidity in upland Wales

Macroinvertebrates were collected from riffles at 104 sites in upland Wales during April and July 1984. Species assemblages were ordinated by DECORANA, classified by TWINSPAN and related to stream chemistry and other environmental factors using correlation and multiple discriminant analysis. DECORANA axis 1 was most strongly correlated with pH and aluminium concentration whilst axis 2 correlated with stream gradient and flow. Four TWINSPAN site groups established in each season were also principally related to pH and aluminium concentration, and reflected overall taxon-richness; differences between groups were most apparent during spring, when catchment forest cover and taxon-richness were also related. A dichotomous key based on indicator species was established for each season with the coleopteran Hydraena gracilis Germar and the Ephemeroptera, including Baetis rhodani Pictet, important indicators at Level 1. We propose that these indicator systems may be used for the rapid detection and assessment of acid waters throughout Wales, and that the methodology is applicable generally.     

Scales and frequencies of disturbances: rock size, bed packing and variation among upland streams

1. Disturbance is an important source of variability in species composition and diversity, but application of disturbance models is contingent upon a very good understanding of the spatial scales and frequencies of disturbance. Such information is particularly lacking from streams. In this study, we measured the disturbance levels of rocks (defined here as the proportion of the original sample disturbed after 6 months) of differing sizes and positions within the stream bed and looked at the variation between and within three upland streams. 2. Rocks were blazed with distinctive marks in situ and mapped using simple trigonometry and permanently marked points on the banks. Forty rocks were selected and marked completely at random, and a further sixty marked from random selections within three size classes (small, medium and large) crossed with two bed-packing classes (on top of the bed or packed into the bed). This sampling design was used at each of two sites (an upper, order 3 location and a lower, order 4 location separated by ≈ 8–16 km) on each of three rivers and in two periods during the year (a dry summer period and a wetter winter period) for a sample size of 1200 rocks in all. During summer, on-top rocks were removed from below the study sites and placed at random locations through the riffle after marking, to test whether human-placed rocks can provide estimates of natural disturbance levels. 3. Rocks were relocated and classified as disturbed (moved or buried) or not disturbed (found at the same location) after ≈ 6 months. Log-linear modelling revealed that human-placed rocks moved half as often as on-top rocks marked in situ. Overall, small rocks disappeared more frequently than medium ones, which disappeared more often than large ones, and rocks lying loosely on top of the bed were disturbed more often than those packed into the bed. There was no interaction between rock size and bed packing in their effects on disturbance and each of these factors affected disturbance levels in the same way at all six sites and in both seasons. During the summer, there were no differences between upper and lower sites, but disturbance was still relatively frequent and patchy in occurrence, with five of six sites showing significant spatial clumping of disturbed rocks. Disturbance levels were higher in the wetter, winter season than during the drier, summer season, but this was caused by a doubling of disturbance rates at all three lower sites, which also showed reduced (but, in two cases, still significant) levels of spatial aggregation. Disturbance levels at upper sites in winter were similar to summer rates, and the level of aggregation of disturbed rocks differed between sites. 4. The results obtained by this study suggest that disturbance levels should not be assessed using methods where rocks are placed in riffles. Disturbance models applied to rocky upland streams may need to heed differences seen at small scales (i.e. between individual rocks), as differences seen at these scales were a crucial source of variability and potentially as significant as variation between sites. However, small-scale differences in disturbance were expressed similarly in different locations. Potentially, the same disturbance model could be applied to all sites, with each of them sitting in different locations along common disturbance continua.     

Catastrophe modelling in the biological sciences

Catastrophe Theory was developed in an attempt to provide a form of Mathematics particularly apt for applications in the biological sciences. It was claimed that while it could be applied in the more conventional physical way, it could also be applied in a new metaphysical way, derived from the Structuralism of Saussure in Linguistics and Lévi-Strauss in Anthropology.Since those early beginnings there have been many attempts to apply Catastrophe Theory to Biology, but these hopes cannot be said to have been fully realised.This paper will document and classify the work that has been done. It will be argued that, like other applied Mathematics, applied Catastrophe Theory works best where the underlying laws are securely known and precisely quantified, requiring those same guarantees as does any other branch of Mathematics when it confronts a real-life situation.     

The influence of chemistry and habitat features on the microcrustacea of some upland Welsh streams

SUMMARY. 1. Microcrustaceans were sampled during September 1990 from three microhabitats (margins, stony riffles and gravel) at thirteen stream sites in mid-Wales.
2. Patterns in the distribution, abundance and community structure of microcrustacean taxa were related to stream chemistry, physiography, substratum composition and marginal habitat structure. Patterns of distribution between microhabitats were also assessed.
3. Chemical variables were the strongest correlates with microcrustacean parameters. Total numbers of microcrustaceans and harpacticoid copepods were reduced at sites with high aluminium concentrations, and ostracods were scarce or absent at the most acidic sites. In contrast, cyclopoid copepods were more abundant and species rich at low pH.
4. Microcrustacean communities were most species rich at sites with a high percentage cover of macrophytes on the substratum. Cladoceran species richness and abundance, and the abundance of several microcrustacean species, were also positively correlated with percentage macrophyte cover.
5. Three site groupings produced by TWINSPAN classification were primarily related to percentage macrophyte cover and marginal habitat structure, with chemistry and land use also of importance.
6. Total microcrustacean abundance was highest in margins, but of all the species, only the harpacticoid Bryocamptus cuspidatus showed a strong microhabitat preference.