Estimation of activities of some enzymes associated with energy-yielding metabolism in the polychaete, Glycera alba (Müller), and application of the methods to the study of the effect of organic pollution

Methods for coenzyme-linked spectrophotometric assays of activities of several enzymes associated with the energy-yielding metabolism of the predatory polychaete, Glycera alba (Müller), have been examined with respect to effects of methods of collection, preservation, and extraction of material, and the composition of assay reaction media on enzyme activities estimated on crude extracts of individual worms. Liquid nitrogen and Drikold (solid CO₂) were equally effective for the preservation of specimens prior to the enzyme assays, and phosphate buffer (0.1 M, pH 7.5) was a generally useful extractant. Effects of reaction conditions on phosphofructokinase activities are detailed. This enzyme had an optimum pH of 8.25 and was inhibited by ATP at pH 6.9 but not at the pH optimum.Activities of phosphofructokinase, pyruvate kinase, malate dehydrogenase, α-glycerophosphate dehydrogenase, lactate dehydrogenase, phosphoenolpyruvate carboxykinase, citrate synthase, and glutamate dehydrogenase have been assayed in crude extracts of G. alba from four sampling stations at various distances from the source of discharge of organic effluent from a seaweed factory into Loch Creran in the west of Scotland. Mean phosphofructokinase activity and to a lesser extent, mean pyruvate kinase activity, were lowest in the group of G. alba collected from the location most affected by the organic input.The results are discussed in relation to the reliability of the enzyme assays, the enzyme activity profile of G. alba with respect to its ecology, and the development of a biochemical index of effects of organic pollution on this representative of the “pollution-sensitive” macrobenthic invertebrate species found in unpolluted or “moderately” polluted areas of the marine environment.     

Surface nutrients,chlorophyll a and phaeopigment in some Scottish sea lochs

The distribution of several surface properties, both physical and chemical, of three Scottish sea lochs were studied during the spring phytoplankton bloom.Concentrations of nitrate, phosphate, and silicate have been determined and related to fluctuations in the freshwater input and uptake by the phytoplankton population. The ratios nitrate: phosphate and silicate: phosphate (on atom bases) in Loch Creran and Loch Linnhe were similar, but there were certain marked differences from those in Loch Etive.Nitrate, rather than phosphate, was a critical factor in the productivity of Loch Creran and Loch Linnhe. In Loch Etive the limitation of these nutrients was directly affected by the amount of fresh-water entering into the Loch.Patchiness of chlorophyll a and phaeopigments in Loch Creran and Loch Etive reflected those factors (e.g., freshwater/sea-water inflow, turbulence produced by the loch topography etc.) which affect the surface circulation in the two lochs.The delay in the development of the phytoplankton bloom in Loch Linnhe may be attributed to a poor stability of the surface layers due to low run-off.     

Biological availability and turnover rate of acetate in marine and estuarine sediments in relation to dissimilatory sulphate reduction

Abstract Acetate turnover rates were determined using ¹⁴C acetate in sediment slurries from two Scottish sea lochs and an estuary which had different rates of oxygen uptake and sulphate reduction. Turnover rates in Loch Etive and Loch Eil were 0.504 and 0.651 μMh⁻¹ respectively, but in the River Tay Estuary there was substantially higher acetate turnover (12.22 μMh⁻¹). The addition of 20 mM sodium molybdate, a specific metabolic inhibitor of sulphate-reducing bacteria (SRB), resulted in a complete inhibition of acetate turnover. These data suggest that SRB were solely responsible for acetate oxidation in these sediments. A comparison of acetate turnover rates in the absence of molybdate and accumulation rate in the presence of the inhibitor demonstrated that at least two pools of acetate with different biological availabilities existed. In Loch Etive only 19% of chemically measured acetate was available with corresponding values of 48% and 65% for Loch Eil and the Tay Estuary respectively.     

Soluble fractions of phosphorus compounds and alkaline phosphatase activity in loch creran and loch etive,Scotland

Reactive phosphorus, polyphosphate, monophosphate ester, dissolved organic phosphorus, and alkaline phosphatase activity were measured in Loch Creran and Loch Etive. Argyll, Scotland during the spring outburst of 1976. In addition, data on chlorophyll a, phacopigment, and salinity were obtained on each occasion.The mean levels of reactive phosphorus initially present in Loch Creran and Loch Etive were 0.50 and 0.22 μg-at. PO₄?P/I, respectively. As the diatom bloom progressed, it fell below the limit of detection. The mean values of polyphosphate were between 0.05–0.08 μg-at. PO₄?P/l in Loch Creran and 0.04 μg-at. PO₄?P/l in Loch Etive, and although present at the initiation and decline of the bloom, it was absent when chlorophyll a was high and the reactive phosphorus was low or undetectable. Monophosphate ester, was found in Loch Creran only, and then in the same concentration as that of the polyphosphate fraction. The mean levels of the dissolved organic phosphorus in Loch Creran and Loch Etive ranged between 0.08–0.18 μg-at. PO₄-P/l and 0.05–0.08 μg-at. PO₄?P/l, respectively. Alkaline phosphatase activity was detected when there was no reactive phosphorus in the water: the highest values in Loch Creran and Loch Etive were 0.003 and 0.005 I.U. of enzyme/4 ml of sample, respectively. The highest mean values of chlorophyll a were 8.0 μg/l in Loch Creran, and 6.0 μg/l in Loch Etive. The mean percentages of phaeopigment fluctuated between 20–55% in Loch Creran and 30–65% in Loch Etive. The highest values were found at the end of the bloom. The mean surface salinities ranged from 22.0 to 31.0‰ in Loch Creran and 3.7 to 17.0‰ in Loch Etive. The lowest values were recorded at the end of the bloom. The statistical treatment of the data, suggests a strong relationship between zooplankton grazing and levels of dissolved phosphorus compounds and phaeopigment.     

The Loch Eil project: Planktonic pigments in sediments from Loch Eil and the firth of Lorne

Photosynthetic pigments and their derivatives were measured in sediments in the fjordic Loch Eil and the Firth of Lome, Scotland, between November 1975 and November 1976. After acetone extraction from the top 10 mm of sediment cores, pigments were crudely separated, by fluorescence change on acidification, into (chlorophyll a + chlorophyllide a) and phaeopigments. The greatest pigment concentrations (mean 73 μg · g sediment dry wt^?1) were found in the most reducing sediments which also had a high average proportion (23%) of chlorophyll. The least mean pigment concentration (23 μg · g^?1) and proportion of chlorophyll (17%) were found in the most oxidizing sediments in the Firth of Lorne where there was a clear seasonal cycle, with a peak in sediment pigment concentration and chlorophyll proportion in May and June, just after the planktonic spring increase. The Loch Eil stations showed a less clear or no seasonal cycle; the station most affected by organic input was the most variable from month to month. It was concluded that redox status was the most obvious control of sediment pigment content, whereas the effect of sedimentation of phytoplankton was complex.     

Spectral intensity in some Scottish freshwater lochs

In the course of investigations on the ecology of submerged aquatic macrophytes a narrow-bandwidth spectroradiometer has been used to measure underwater spectral intensity (390–750 nm) in a series of lochs approaching the extremes in optical and chemical properties of Scottish fresh waters. Cosine response and immersion-effect properties of the collector were determined in the laboratory with a colUmated hght source. Diffuse attenuation coefficients, Ee, ranged from 0·55 in Loch Croispol, a calcareous loch, to 2·9 in Loch Leven, a lowland eutrophic loch. Underwater spectral intensity (1 m) relative to subsurface values show a proportional increase in short-wave radiation in the blue-green water of Loch Croispol, while the converse is true of the peaty brown water of Loch Uanagan. Attenuation coefficients were derived over 25 nm wavebands in Lochs Croispol, Leven and Uanagan. The water in the latter two lochs is optically similar although Leven is rich in phytopiankton and has an extinction peak at 675 nm. Uanagan represents the brown-water type of loch most common in Scotland. Loch Croispol has attenuation coefficients for shorter wave-lengths at least ten times lower than the other two lochs and only approaches their values at 750 nm. Croispol resembles Crater Lake, Oregon. The colour range at the maximum colonizable depths by rooted macrophytes of brown (Uanagan 4 m) to blue-green (Croispol 6 m) water is equivalent to 196·0 and 230·6 kJ/Einstein (48·2 and 55·6 kcal/Einstein). Red/far-red ratios even in plankton-rich water (Leven) were at least three times the value for sunlight (1·3) and within the total photic zone reached three-figure proportions. Possible implications for light-sensitive seeds of aquatic species and for morphogenesis and zonation are discussed.     

The Loch eil project: Simulation of the hydrology and sediment inputs to Loch Eil

The two objectives of the study were the assessment of the water and sediment response of the ungauged catchments draining to Loch Eil, and the analysis of the sensitivity of erosion processes to the variation of input precipitation and sediment characteristics. The technique employed was the application of the Strathclyde Sediment Model I, which had already been tested on the Scottish river basins using data from several sources. The method showed some potential but provided only a rough comparison with the actual sedimentation in the Loch.     

Impacts of acidification on brown trout Salmo trutta populations and the contribution of stocking to population recovery and genetic diversity

Anthropogenic acidification in SW-Scotland, from the early 19th Century onwards, led to the extinction of several loch (lake) brown trout (Salmo trutta) populations and substantial reductions in numbers in many others. Higher altitude populations with no stocking influence, which are isolated above natural and artificial barriers and subjected to the greatest effect of acidification, exhibited the least intrapopulation genetic diversity (34% of the allelic richness of the populations accessible to anadromous S. trutta). These, however, were characterised by the greatest interpopulation divergence (highest pairwise D_EST 0.61 and F_ST 0.53 in contemporary samples) based on 16 microsatellite loci and are among the most differentiated S. trutta populations in NW-Europe. Five lochs above impassable waterfalls, where S. trutta were thought to be extinct, are documented as having been stocked in the late 1980s or 1990s. All five lochs now support self-sustaining S. trutta populations; three as a direct result of restoration stocking and two adjoining lochs largely arising from a small remnant wild population in one, but with some stocking input. The genetically unique Loch Grannoch S. trutta, which has been shown to have a heritable increased tolerance to acid conditions, was successfully used as a donor stock to restore populations in two acidic lochs. Loch Fleet S. trutta, which were re-established from four separate donor sources in the late 1980s, showed differential contribution from these ancestors and a higher genetic diversity than all 17 natural loch populations examined in the area. Genetically distinct inlet and outlet spawning S. trutta populations were found in this loch. Three genetically distinct sympatric populations of S. trutta were identified in Loch Grannoch, most likely representing recruitment from the three main spawning rivers. A distinct genetic signature of Loch Leven S. trutta, the progenitor of many Scottish farm strains, facilitated detection of stocking with these strains. One artificially created loch was shown to have a population genetically very similar to Loch Leven S. trutta. In spite of recorded historical supplemental stocking with Loch Leven derived farm strains, much of the indigenous S. trutta genetic diversity in the area remains intact, aside from the effects of acidification induced bottlenecks. Overall genetic diversity and extant populations have been increased by allochthonous stocking.     

The effects of artificial enrichment upon the planktonic and benthic communities in a mesotrophic to hypertrophic loch series in lowland Scotland

Phytoplankton, submerged macrophytes, zoobenthos and zooplankton communities were studied concurrently in three lochs which progressively increased in trophic state. The phytoplankton showed a change in dominance from Bacillariophyceae (Pennales) and Desmidaceae in the mesotrophic Loch of the Lowes to Bacillariophyceae (Pennales) and Cyanobacteria in eutrophic Balgavies Loch and to Bacillariophyceae (Centrales) and Chlorophyceae in hypertrophic Forfar Loch. The Nygaard Compound phytoplankton quotient rose from 1.2 in the Lowes to 12 in Forfar. There was a decrease in number of species of submerged macrophytes, and in depth of colonisation from 5 m in the Lowes to 2 m in Forfar which was proportional to the decrease in mean transparency. Macrophyte biomass, however, increased in proportion to the concentration of total phosphorus in the lochs. There was a decrease in species of littoral zoobenthos, but an increase in density of profundal benthos from the Lowes to Forfar. Zooplankton also showed an increase in density with enrichment, accompanied by changes in dominance. Calanoida were dominant in the Lowes; Cladocera in Balgavies and Cyclopoda in Forfar. The importance of these differences in understanding the effects of progressive enrichment and to the management of the lochs is discussed.     

Sea trout in North Argyll sea lochs: II. diet

The diet of sea trout in some of the sea lochs of the west coast of Scotland was investigated. The contents of 986 sea trout stomachs from the Loch Etive area (1970–1973), and 291 stomachs from the Loch Eil area (1964–1973), were examined and the composition of the diet, seasonal changes and the effect of trout size were analysed using frequency of occurrence, dry weight and number of organisms methods. Benthic feeding (crustacea and annelids) was more important in winter while midwater and surface organisms (young fish and insects) were preferred in summer. Young fish (mainly clupeids and sand eels) featured more in the diet of larger trout (≤21 cm) than in the smaller size range (≥21 cm). Surveys of 24-h were completed in June and September, 1972, and indicated that availability of food was the main factor influencing the presence or absence of trout. Bottom feeding was greatest during the day while the amount of midwater and surface feeding tended to increase between sunset and sunrise.     

Verification of a mathematical growth model ofPhragmites australis using field data from two scottish lochs

A growth model ofPhragmites australis was verified using two independent sets of published field data. The model simulates the growth pattern of a well-established, monospecific stand ofP. australis in the absence of genetic diversity and environmental stresses of mainly nutrient and water deficiency. The model formulated using first order differential equations was combined with plant phenology and comprises five subroutines in which photosynthetically active radiation, shoot, root, rhizome and new rhizome biomass are calculated. Using the model, experimental results were reproduced within reasonable limits having concordance correlation coefficients of more than 0.75 for 70% of the output parameters, which was the main objective of the study. The modelled efficiencies of PAR were 7.15% and 3.09%, as opposed to 7.7% and 2.53% in experimental estimations, for Loch of Forfar and Loch of Balgavies, respectively. Production and seasonal fluxes of dry matter ofP. australis in Scottish lochs were estimated using the modelled quantities for the 1975 growing season in g m⁻². They showed that 31% and 37% of total net photosynthate translocated to rhizomes before shoot senescence began in Loch of Forfar and Loch of Balgavies, respectively. Also in both lochs approximately 45% of total downward translocation came from accumulated shoot dry matter during senescence, while the rest came from photosynthesis before the shoots started to senesce.     

Zooplankton in Loch Lomond: perspectives,predation and powan

Published and unpublished accounts of the zooplankton community of Loch Lomond are reviewed. Loch Lomond's zooplankton community is unique amongst those of Scotland's other large lochs. Holopedium gibberum appeared in Loch Lomond in the years between the studies of Slack & Hamilton in the 1950s and zooplankton surveys in the late 1970s.The annual cycles of Eudiaptomus gracilis, Cyclops strenuus abyssorum and Mesocyclops leuckarti are described for the mid basin of Loch Lomond. E. gracilis overwinter mainly as adult with variable egg production during their lifespan of 5–8 months. The cyclopoid copepods have variable life history characteristics. C. str. abyssorum produce a single generation per year, overwinter by means of resting eggs and a small population of planktonic adults, while M. leuckarti is present in the plankton throughout the year, but in low numbers during winter.Zooplankton populations in Loch Lomond are predated by the diverse fish community which includes a facultative planktivore in the powan, Coregonus lavaretus. Powan feed heavily on zooplankton from late spring until late autumn. Powan display selectivity in the prey species taken, in the size distribution and morph types taken within prey species. Zooplankton species, morphs and forms in Loch Lomond tend to reflect this predation.Zoology Department, University of St. AndrewsZoology Department, University of St. Andrews     

Ammonia assimilation and metabolism byBeggiatoa alba

Beggiatoa alba B18LD was investigated for its pathways of ammonia assimilation. The increase in growth yields ofB. alba with excess acetate was linear from 0.1 to 2.0 mM ammonia.B. alba had strong glutamine synthetase (GS) and glutamate synthase (GOGAT) activities, irrespective of the ammonia concentration in the medium. Glutamate dehydrogenase activity was not found, and alanine dehydrogenase (aminating) was observed only whenB. alba was grown at high (2.0 mM) ammonia. Methionine sulfoximine, an inhibitor of GS, inhibited growth ofB. alba irrespective of the ammonia concentration in the medium. Thus it appears the primary pathway for ammonia assimilation inB. alba is via the GS-GOGAT pathway at both low and high ammonia concentrations. Preliminary experiments were unable to discern if theB. alba GS is modified by covalent modification.Non-standard abbreviations GS Glutamine synthetase - GOGAT glutamate-oxoglutarate aminotransferase - GDH glutamate dehydrogenase - ADH alanine dehydrogenase - MSX methionine sulfoximine - GOT glutamate-oxaloacetate aminotransferase - GPT glutamate-pyruvate aminotransferase     

The macrophytic vegetation of Loch Urigill and other lochs of the Ullapool area

Summary

From the 15 lochs examined at altitudes between 9 and 267 m in western Sutherland and Wester Ross, a few areas of tall fen (with Hippuris, Sparganium erectum, Osmunda etc.) survive to indicate its potential distribution. Predominant grazed fen comprises patches of short Juncus articulatus and Carex nigra. Reedswamp is scarce, being dominated by Eleocharis palustris, Carex lasiocarpa and Schoenoplectus lacustris and, rarely, by Phragmites and Sparganium erectum. Floating-leaved vegetation was only encountered twice. Submerged vegetation to water depths of at least 1 m consists in the main of sparse Lobelia-Littorella with Juncus bulbosus, typical of the prevailing stony shores in every loch. Extensive Isoetes lacustris meadows may occur beyond that, with Potamogeton praelongus or P. perfoliatus stands in waters of alkalinities over 0·45 m-equiv 1^-1, reaching depths of 3 m in Loch Urigill. Loch Urigill is unusual in having dominant Isoetes at a water alkalinity of 0·76 m-equiv 1^-1 along with abundant Anabaena species and some Potamogeton filiformis.     

Shoot development and production studies of phragmites Australis (Cav.) Trin. Ex Steudel in Scottish Lochs

Studies were made on the development and production of Phragmites australis (Cav.) Trin. ex Steudel (= Phragmites communis Trin.) stands from Forfar Loch-polytrophic, Balgavies Locheutrophic and Loch of the Lowes-mesotrophic. Differences were detected in the shoot height, shoot dry weight, standing crop biomass, leaf area index, crop growth rate and net efficiency of solar energy conversion among the three lochs. Such differences were partly due to the different nutrient levels of the three locations. The percentage ratio of leaf to shoot weight, leaf area ratio, relative growth rate and net assimilation rate for the reed stands in the three lochs were also calculated and were compared with values from other study areas.     

Light limitation of phytoplankton development in an oligotrophic lake - Loch Ness, Scotland

• 1 The underwater light climate in Loch Ness is described in terms of mixing depth (Z_m) and depth of the euphoric zone (Z_eu). During periods of complete mixing, Z_m equates with the mean depth of the loch (132 m), but even during summer stratification the morphometry of the loch and the strong prevailing winds produce a deep thermocline and an epilimnetic mixed layer of about 30 m or greater. Hence, throughout the year the quotient Z_m/Z_eu is exceptionally high and the underwater light climate particularly unfavourable for phytoplankton production and growth.
• 2 Phytoplankton biomass expressed as chlorophyll a is very low in Loch Ness, with a late summer maximum of less than 1.5 mg chlorophyll a m^-3 in the upper 30 m of the water column. This low biomass and the resulting very low photosynthetic carbon fixation within the water column are evidence that a severe restraint is imposed on the rate at which phytoplankton can grow in the loch.
• 3 The chlorophyll a content per unit of phytoplankton biovolume and the maximum, light-saturated specific rate of photosynthesis are both parameters which might be influenced by the light climate under which the phytoplankton have grown. However, values obtained from Loch Ness for both chlorophyll a content (mean 0.0045 mg mm^-3) and maximum photosynthetic rate (1–4 mg C mg Chla^-1 h^-1) are within the range reported from other lakes.
• 4 Laboratory bioassays with the natural phytoplankton community from Loch Ness on two occasions in late summer when the light climate in the loch is at its most favourable, suggest that even then limitation of phytoplankton growth is finely balanced between light and phosphorus limitation. Hence, for most of the year, when the light climate is less favourable, phytoplankton growth will be light limited.
• 5 Quotients relating mean annual algal biomass as chlorophyll a (c. 0.5 mg Chla m^-3) and the probable annual specific areal loading of total phosphorus (0.4–1.7 g TP m^-2 yr^-1) suggest that the efficiency with which phytoplankton is produced in Loch Ness per unit of TP loading is extremely low when compared with values from other Scottish lochs for which such an index has been calculated. This apparent inefficiency can be attributed to suppression of photosynthetic productivity in the water column due to the unfavourable underwater light climate.
• 6 These several independent sources of evidence lead to the conclusion that phytoplankton development in Loch Ness is constrained by light rather than by nutrients. Loch Ness thus appears to provide an exception to the generally accepted paradigm that phytoplankton development in lakes of an oligotrophic character is constrained by nutrient availability.

     

Genetic Variation at Structural Loci in the Glossina morsitans Species Group

Gene diversity was investigated in four taxa of tsetse flies (Diptera: Glossinidae) including Glossina morsitans morsitans, G. m. centralis, G. swynnertoni, and G. pallidipes. Histochemical tests were performed for 35–46 isozymes. Polymorphic loci were 20% in G. morsitans morsitans, 32% in G. m. centralis, 17.6% in G. swynnertoni, and 26% in G. pallidipes. Mean heterozygosities among all loci were 6.6% in G. morsitans morsitans, 6.0% in G. m. centralis, 7.1% in G. swynnertoni, and 6.8% in G. pallidipes. Allozyme gene diversities were considerably less than those reported for many Diptera. The low gene diversities are probably related to small effective population sizes.     

Chemical composition studies on some aquatic macrophytes in three Scottish lochs. I. Chlorophyll,ash, carbon,nitrogen and phosphorus.

The chlorophyll, ash, carbon, nitrogen and phosphorus levels in seven species of freshwater macrophytes (Juncus effusus L., Iris pseudacorus L., Carex rostrata Stokes, Glyceria maxima (Hartm.) Holmberg, Nuphar lutea (L) Sm., Polygonum amphibium L. and Schoenoplectus lacustris (L) Pallas) in three Scottish lochs of different trophic levels were studied during 1975. Mean chlorophyll levels varied from a minimum of 1.73 mg g^–1 dry weight in Balgavies Loch Juncus to 10.22 mg g^–1 dry weight for Forfar Loch Iris. Carbon contents ranged from 450 to 520 mg g^–1 ash-free dry weight. For ash, nitrogen and phosphorus, significant differences in mean concentrations were detected among plant species as well as within one plant species growing in different lochs. Positive correlations were apparent between the degree of eutrophication in the study areas and the amount of ash, phosphorus and nitrogen present in the plants growing in them.     

Aquatic angiosperm communities from lochs on Rhum

Summary

The plant communities from twenty lochs and lochans on Rhum are surveyed. Species lists are given for each site. Aquatic vegetational cover is sparse and only a limited number of plant communities are present, predominantly those involving Lobelia dortmanna and Littorella uniflora as dominants or co-dominants, and a more restricted representation of communities with Equisetum fluviatile, Phragmites australis or Nymphaea alba subsp. alba. Exposure and poor substratum are suggested as factors causing limited plant growth. This work is discussed in the light of other studies carried out in Scotland.