The relative abundance of brown trout in acidic softwater lakes in relation to water quality in tributary streams

The influence of the water quality of tributary streams on the relative abundance in benthic gillnet catches (catch per unit effort, cpue) of allopatric brown Salmo trutta was assessed in associated acidic, softwater lakes. The study was carried out over 6 years (1989–1994) in 15 lakes located at altitudes between 230–715 m a.s.l. in two Norwegian catchments. The water quality of the main inlets and outlets varied little, as indicated by their of pH range (4·93–5·51) and calcium concentrations (0·19–0·44 mg 1⁻¹), but varied more with respect to concentrations of inorganic, monomeric aluminium (7·0–41·0 μg l⁻¹). Most of the lakes were also fed by secondary streams with better water quality: a maximum pH of 6·56, calcium levels of up to 0·74mg 1⁻¹, and inorganic aluminium levels as low as l·0 μg 1⁻¹. The cpue was inversely correlated with lake altitude ( r ²=0·50), and thus was adjusted to a mean altitude. The calcium concentration in the richest secondary stream to each lake, its richness judged on the basis of its acid-neutralizing capacity, had the highest predictive power of the variability in cpue ( r ² = 0·49).The calcium content in the other secondary streams or in the main inlets and outlets did not correlate with cpue. Alkalinity in the main outlets correlated to some extent with cpue ( r ² = 0·27). It is suggested that secondary streams with good water quality provide important refuges for the recruitment of brown trout in acidic softwater lakes.     

Effects of aluminium in hard, acid water on metabolic rate, blood gas tensions and ionic status in the rainbow trout

Rainbow trout, Salmo gairdneri , cannulated in the dorsal aorta, were exposed to 2 mgl⁻¹ aluminium (added as Al₂(SO₄)₃, 18H₂O) at pH = 5.0 in hard water of very high calcium concentrations (3.3 mmol 1⁻¹). No changes were observed in the concentration of the major plasma ions. The fish, however, became hypoxic, as seen from a fall in the dorsal aortic oxygen tension from around 100 mmHg to 30–40 mmHg, and a simultaneous increase in carbon dioxide tension. Moreover, exposure resulted in a 15% increase in standard oxygen uptake. The ventilation frequency doubled and swelling of the erythrocytes was observed. The results are discussed in the light of some earlier data on the effects of acid/aluminium exposure in water of low calcium concentration, and reveal an important role for calcium in the physiological responses to low pH and aluminium.     

Some effects of low pH and calcium on the growth and tissue mineral content of yearling brown trout, Salmo trutta

Yearling brown trout, Salmo trutta , were exposed to low mineral content water (nominal concentrations of 20μmol 1⁻¹ magnesium, 7.7 μmol 1⁻¹ potassium, 44 μmol 1⁻¹ sodium) over a pH range of 4.0–5.2 with ambient calcium concentrations of 2.5–60 μmol 1⁻¹. All fish died at pH 4.0 and 4.2 irrespective of ambient calcium concentration and also at pH 4.4 with only 2–3 μmol 1 ⁻¹ calcium (that is calcium-free water except for that leached from the diet or excreted by the fish). Good growth rates were obtained over the remaining treatments which extended down to pH 4.4 with as little as 7 μmol 1⁻¹ calcium. When starved, weight loss was inversely correlated with pH. Effects on plasma chloride, percentage dry weight and calcium, potassium sodium, and phosphorus contents of skin, muscle and bone tissue were also investigated. These demonstrated pH effects on mineral metabolism in starved fish, but no effects were detected in fed fish.     

Catecholamine release and interrenal response of brown trout, Salmo trutta, exposed to aluminium in acidic water

Cannulated brown trout, Salmo trutta , were exposed for 36 h to synthetic water with a low calcium content of pH 5 and similar synthetic water dosed with aluminium to raise the filterable A1 from 5 to 290 μg 1⁻¹ over the 36-h period. There were no significant disturbances of plasma concentrations of glucose, cortisol or catecholamine (adrenaline and noradrenaline) in fish held in water of pH 5. The addition of aluminium to this acidic synthetic water resulted in a generalized endocrine stress response with a four-fold increase in plasma glucose concentration after 18 h and a significant increase in plasma cortisol concentration from 24 h onwards when filterable A1 exceeded 200 μg 1⁻¹. Plasma catecholamine concentration indicated an adrenergic stress response in aluminium-exposed brown trout. A transient doubling in noradrenaline after 6 h in A1 was followed by a larger increase in both plasma adrenaline and noradrenaline concentrations in fish surviving the 36-h exposure to A1.     

A model relating the results of low pH bioassay experiments to the fishery status of Norwegian lakes

SUMMARY. 1. Data from the literature concerning several Norwegian brown trout (Salmo trutta) populations are used to construct Leslie matrices in order to estimate their capacity to withstand egg and fry mortalities. These estimates are compared with the results of bioassay studies conducted at pH 4.5 and at various calcium concentrations in order to predict the percentage of trout populations which would be able to survive these conditions.
2. These predictions are in good agreement with the observed fishery status of Norwegian lakes at calcium concentrations <30 /μeq I⁻¹ but above this concentration the recorded status is worse than predicted.
3. Likely causes for this difference are considered to be effects acting on fish older than those used in the bioassay experiments (either mortality or reduction in fecundity possibly due to impaired growth) or the effects of other water quality factors, for example aluminium.
4. These effects need only be fairly slow acting as compared with the egg and fry mortalities so far studied in detail. Thus the discrepancy between predicted and observed results at pH 4.5 and 50μeq 1-⁻¹ calcium could be caused by an additional 80% mortality acting over the 4–8 year period before maturation.
5. An additional factor which could result in the observed fishery status being worse than predicted is temporal variation in water chemistry which is not assessed in the survey of Norwegian lakes. In particular, the adverse conditions occurring at time of snowmelt could be critically important.     

The influence of calcium on aluminium-induced changes in the growth rate and mortality of brown trout, Salmo trutta L.

Yearling brown trout, Salmo trutta L., were maintained in a synthetic medium at pH 5.2 with various concentrations of calcium (4–400 μumol l⁻¹) and aluminium (0–3 μumol l⁻¹). In general, higher mortality and lower growth rates were found at higher aluminium concentrations, with these effects being reduced at high ambient calcium. Aluminium toxicity was greater in an experiment conducted in February–March than in an experiment conducted in October–December; this could be due to a seasonal variation in tolerance of the fish. Aluminium exposure resulted in an initial period of weight loss and high mortality followed by some recovery over the 6-week period of the experiments. In the first experiment, as in previously reported work, adaptation was incomplete in that aluminium-induced growth rate reduction was still evident to the end of the test. In the second experiment, however, no significant effect of aluminium treatment on growth was found during weeks 4 to 6 of the exposure period.     

Growth, mineral uptake and skeletal calcium deposition in brown trout, Salmo trutta L., yolk-sac fry exposed to aluminium and manganese in soft acid water

In 30-day exposures in artificial soft water medium, survival of brown trout alevins was not affected by low pH (4.5,4.8, 5.4), by low calcium concentration (10.25 μmol l⁻¹) or by manganese (≤20 μmol l⁻¹), but was impaired by aluminium (6–8 μmol l⁻¹) at low calcium concentration (10 μmol l⁻¹) irrespective of pH (4.5 or 5.4). Manganese (6.6, 20 μmol l⁻¹) impaired net calcium uptake and calcium deposition in the skeleton at low calcium concentration (25 μmol l⁻¹) irrespective of pH. Aluminium (2–8 μmol l⁻¹) impaired gross development, net uptake of calcium, potassium and sodium, and calcium deposition in the skeleton, and slightly increased the net loss of magnesium, some of these effects being more severe at calcium concentration 10 μmol l⁻¹ than 50 μmol l⁻¹, and some more severe at pH 5.4 than pH 4.5. Net uptake of calcium and sodium were impaired at low pH (4.5, 4.8), and skeletal calcium deposition was impaired at low calcium concentration (10 μmol l⁻¹), but these effects of low pH and low calcium concentration were slight compared with those of the trace metals. The possible role of trace metals in reports of the deleterious effects on fish of low pH levels is discussed.     

Juvenile salmonid production in a Lake Erie nursery stream

Normandale Creek (2531m²) provides spawning and nursery grounds for lake-run rainbow trout, Salmo gairdneri Richardson, brown trout, Salmo frutta L. , and coho salmon, Oncorhynchus kisutch (Walbaum). Upstream movements are significantly correlated with peak stream discharge (P < 0.05). In 1973–1974, 59 adult salmonids constructed 86 nests of which 60% were disturbed by re-use or sand deposition. From a calculated deposition of 90 400 ova, 483 juveniles, 6 cm in fork length ( f.l. ), were resident in the stream when estimates by electrofishing first became valid. Estimated within-stream (i.e. all sections combined) monthly densities of wild juveniles >6 cm f.l. ranged from six to 22 fish (100 m²)⁻¹, and biomass from 1.3 to 3.1 gm⁻². The highest within-section biomass was 8.3 gm⁻² in November. Additions of hatchery-reared rainbow trout temporarily increased monthly within-stream juvenile density up to 23 fish (100 m²)⁻¹, and biomass to 8.2 g m⁻². Density and biomass were positively correlated (P < 0.05) with both instream and bank cover, and biomass negatively correlated (P < 0.05) with gradient. The importance of substratum and flow characteristics on juvenile density and biomass was strongly indicated although not statistically significant. Annual within-section production of juveniles > 6 cm f.l. ranged from 4.79 to 5.93 g m⁻² year⁻¹ in Sections III and IV, respectively. Inclusion of calculated probable numbers of fish < 6 cm f.l. increased within-section production from 5.93 to 18.65 g m⁻² year^−l in upstream Section IV.     

Effect of rearing density on thyroid and interrenal gland activity and plasma and hepatic metabolite levels in rainbow trout, Salmo gairdneri Richardson

There were significant inverse correlations between rearing density of rainbow trout, Salmo gairdneri Richardson, and final body weight, plasma L-thyroxine (T₄), trüodo-L-tryronine (T₃), cortisol and protein concentrations, plasma T₄/T₃ ratios and thyroid epithelial cell height. In addition, hepatosomatic indices and plasma free fatty acid concentrations were higher in fish reared at low (134 g 1⁻¹) density compared with groups reared at medium (210g1⁻¹) or high density (299g 1⁻¹), and the post-feeding (3.5-4h) elevation in plasma glucose and triglyceride levels evident in trout maintained at low rearing density was not found in those fish reared at higher densities. There were no significant effects of rearing density on hematocrit, carcass composition, hepatic glycogen and lipid levels and interregnal nucleus size.     

Some effects on the growth of brown trout from exposure to aluminium at different pH levels

Yearling brown trout, Salmo trutta L., were exposed to various concentrations of inorganic aluminium (0–3.7 μM1⁻¹) over a pH range of 4.3–6.5 in a flow-through bioassay apparatus using synthetic test media. Low pH, in the absence of aluminium, produced little effect on growth or survival except at the lowest pH tested (4.3). At pH less than 5.5, concentrations of total aluminium in excess of 1 μM 1⁻¹ (27μg 1⁻¹) were found to retard growth. The effects of a given aluminium concentration were markedly reduced at pH above 5.5.
The change in aluminium toxicity with pH must be related to changes in aluminium chemistry. When growth rates are correlated with the different aluminium species, calculated using thermodynamic equilibrium constants given in the literature, it appears that the Al(OH)^{2 +} species is the most toxic, with a small contribution also coming from polymeric complexes.     

Preliminary empirical models of the historical and future impact of acidification on the ecology of Welsh streams

SUMMARY 1. We describe a preliminary approacb to modelling the impact of acidification on the ecology of two Welsb streams. Output from the hydrochemical Model of Acidification of Groundwaters in Catchments (MAGIC) was used to drive empirical models which predicted brown trout Salmo trulta (L.) survival, trout density and invertebrate assemblage type. The models were used for hindcasts between 1844 and 1984 under conifer forest and moorland conditions. Forecasts involved each of these land uses with sulphate deposition either continued at 1984 levels or reduced by 50%.
2. Trout survival times and trout densities in the models declined markedly between 1844 and 1984. The most severe decline occurred under simulated forest, where high aluminium concentration led to the virtual elimination of trout in both streams.
3. In forecasts, only in simulated moorland with sulphate deposition reduced by 50% of 1984 levels, was further decline in trout population retarded. There was no marked recovery in trout density under any of the conditions examined.
4. Invertebrate assemblages in streams during the nineteenth century may have differed from those now existing in nearby moorland streams which are presently circumneutral. Past chemical conditions were unusual (<3mg 1⁻¹ total hardness, but pH >5.7 and low aluminium) by present-day standards, and were outside the range of the invertebrate model until -1940.
5. Between the 1940s and 1984 there was no change in invertebrate fauna under the moorland scenario despite some acidification. However, simulated forest advanced the appearance of the most impoverished assemblage type, which did not recover in spite of reduced deposition.
6. We discuss several uncertainties with the models in their present form, but suggest some methods for their testing and validation.     

The effect of sodium and calcium concentrations on the hatching of eggs and the survival of the yolk sac fry of brown trout, Salmo trutta L. at low pH

The effect of various external concentrations of sodium and calcium on the survival and hatching of brown trout, Salmo trutta , eggs at pH 4.5 was tested. A calcium concentration of approximately 10 ppm (500 μE l⁻¹) enables freshly fertilized eggs to survive whereas eyed ova are tolerant of deionized water acidified by sulphuric acid with no other ions added. Concentrations of sodium and calcium of 1 ppm (44 and 50 μE l⁻¹, respectively) are sufficient to ensure the successful hatching of eyed ova and subsequent survival of the alevins. At pH 4.5 hatching is prolonged by the alevins passing through a temporary encapsulated stage.     

Development of lamellar epithelial hyperplasia in gills of pantothenic acid-deficient rainbow trout, Salmo gairdneri Richardson

Two semisynthetic diets differing in the levels of pantothenic acid (PA) supplementation (0 or 40 mg kg⁻¹ diet) were fed to rainbow trout, Salmo gairdneri Richardson, initially weighing 0.7 g fish⁻¹. Each diet was fed to two tanks of fish and, at the start of the feeding trial, each tank contained 200 animals. The experiment was conducted for 28 days during which time, every 2 days, gill tissue was sampled and food consumption was determined. Lamellar hyperplasia was first detected in the gills of deficient fish on the 12th day sample, while reduced feed intake was first manifest at 16 days. Hyperplasia first appeared in the distal regions of the gill filaments, but the lesion rapidly progressed in a proximal direction and 35 of 40 fish examined on days 22–28 exhibited hyperplasia on more than 75% of the filament surface. Gill lamellar hyperplasia is a sensitive indicator of PA deficiency in the rainbow trout. Moreover, the lesion is specific to PA deficiency since its developmental pattern differs histologically from the lamellar hyperplasia of the non-nutritional gill diseases.     

Growth, production and bioenergetics of brown trout in upland streams with contrasting riparian vegetation

1. A series of laboratory-based equations on trout growth and bioenergetics developed by J.M. Elliott were applied to data collected for brown trout ( Salmo trutta L.) under field conditions in Co. Mayo, Western Ireland. Fish were collected by electrofishing eight upland streams with contrasting riparian vegetation; grassland, open canopy and closed canopy deciduous.
2. Stream temperatures, one of the main influencing factors on fish growth and energetics, did not differ significantly between riparian types.
3. Observed growth rates were lower than the predicted maximum growth rates and were not influenced by riparian vegetation type. Growth ranged between 0.66% day⁻¹ for 0 + trout to 0.08% day⁻¹ for 2 + trout.
4. Production estimates showed no clear difference between riparian vegetation types over the growing season.
5. Fish densities and biomass tended to be greater in closed canopy streams particularly in summer.
6. Actual ration sizes calculated for trout were similar to the ration required for maintenance metabolism and were only 45–63% of the maximum potential rations. Although there was an ontogenetic increase in ration size with increasing fish age, the proportion of ration available for growth (i.e. the difference between actual and maintenance rations) did not differ between age classes but was greatest in summer. 1+ and 2+ trout show greatest ration available for growth in grassland streams.
7. Trout growth did not differ between riparian vegetation types but did vary seasonally with greatest attainment in summer. Growth was limited in the present study possibly due to combined effects of reduced prey available to fish and low stream temperatures reducing metabolic requirements. In such food limited systems, terrestrial invertebrate energy subsidies could have significant benefits to brown trout growth, production and bioenergetics.     

The production of brown trout, Salmo trutta in tributaries of the Upper Wye, Wales

The population density, age structure, biomass, growth and production of brown trout were investigated in four tributaries of the upper River Wye. The populations at each site were largely maintained by immigration from nursery areas. Abundance of separate year classes at sites on the three largest tributaries reached a peak at age 2+. On the smallest stream numbers reached a peak at 1+. Recruitment occurred throughout the year but decreased with age of year classes. Maximum O+ densities ranged from 0.04 to 0.89 m⁻², and >0+ densities from 0.13 to 0.59 m⁻². Average total biomass in 1975 ranged from 2.6 to 14.2 g m⁻². Within the study sites annual trout production in 1975 ranged from 2.9–19.7 g m⁻². Production values were dependent on age structure and population mobility at the study sites. In the three largest streams 2+ and 3+ fish contributed 66.3–88.3% of total production whilst 1+ and 2+ fish contributed 74.5–84.5 % of the total in the smallest stream. The mobile (non-resident) component of the population accounted for up to 60–70% of production at certain times of the year, but over the year (1976) accounted for =30 % of total production. The resident component of the highest annual production value (19.7 g m⁻²) was estimated to be between 15.0 and 18.2 g m⁻².     

The interactions of water hardness and pH with the acute toxicity of zinc to the brown trout, Salmo trutta L.

Exposure of brown trout, Salmo trutta , to zinc under continuous flow conditions over 96 h showed that both water hardness and pH exert major influences on the toxicity of the metal. 96-h LC50 values for total zinc ranged from <0.14mg 1⁻¹ in alkaline soft water (pH 8; lOmg 1⁻¹ as CaCO₃) to 3.20 mg 1⁻¹ in acidic hard water (pH 5; 204 mg 1⁻¹ as CaCO₃). A variable reduction in zinc toxicity in hard water compared with soft water over the pH range 4–9 was attributed to high external calcium. Zinc toxicity was positively correlated with decreasing acidity over the pH range 5–7, the metal being most toxic at pH 8–9 where metal complexes predominate. Below pH 5 metal toxicity also increased, irrespective of hardness. Water hardness and pH interacted with zinc toxicity in a complex manner, apparently dependent on physical and chemical transformations of the metal, and as changes in uptake. detoxification and excretion by the fish.     

Observations on siting, dimensions and structure of salmonid redds

Redds of trout, Salmo trutta L., Atlantic salmon, S. salar L., and rainbow trout, S. gairdneri Richardson, were studied at sites in north-east England, south-west Wales and southern England (Dorset). Spawning females preferred gravels of 20–30 mm diameter and water velocities at 0·6 depth of 15 cm s^-1 and < c. 2·0 body lengths s^-1. After logarithmic transformation the major horizontal dimensions of redds could be related to one another by linear regressions, and redd tail length could, similarly, be related to female fish length. Egg burial depth in upland stony streams in north-east England and south-west Wales could be predicted from a linear regression of mean depth upon female length. Information is also given on gravel composition and redd structure.     

In vitro evidence for the ionoregulatory role of rainbow trout mucus in acid, acid/aluminium and zinc toxicity

Rainbow trout body mucus dialysed with acidified distilled water at pH 7,5 and 3 experienced ion depletion which was greatest at pH 3 and minimal between pH 7 and 5. Mucus Na⁺ loss is exacerbated in the presence of 1 mg I⁻¹ aluminium as A1₂(SO₄), at pH 5 and 7. Al₂(SO₄), causes greater depletion of Na⁺ from mucus than A1C1₃. A lethal level of zinc (2 mg 1⁻¹) does not deplete mucus Na⁻ or K⁺, unlike a lethal level of aluminium (1 mg 1⁻¹) at pH 7. The results are discussed in terms of the ionoregulatory role of mucus in heavy metal and acid toxicity.     

Flexible modes of anadromy in Baltic sea trout: making the most of marginal spawning streams

From examination of the ratios of strontium to calcium laid down as a lifetime record in the otoliths of sea trout Salmo trutta from Gotland, Baltic Sea, it was found that: (1) the shortest stream was used mostly by precociously emigrant or coastally hatched spawners; (2) longer streams had more fish that underwent normal smoltification; (3) sea-caught fish were predominantly coastally hatched (presumably near stream mouths). Furthermore, some otoliths showed no evidence of a freshwater history at all, raising the possibility of a contingent of the coastal population that does not depend on riverine spawning. The results emphasize the importance of the coastal zone as natal and early life habitat for sea trout in the Baltic, particularly with respect to a potential change to a warmer climate which may exacerbate conditions within small, ephemeral trout streams.     

Factors limiting fish populations in the Loch Fleet system, an acidic drainage system in south-west Scotland

Loch Fleet is an oligotrophic upland lake in Galloway, south-west Scotland. It once supported a brown trout, Salmo trutta L., sport fishery with low annual catches (< 150 fish year ^?1) but catches declined markedly after 1950 and no fish were caught after 1975. Diatom records for the lake sediments indicate acute acidification since 1975, pH changing from c. 5–8 to 4–6. In 1984 a project was set up at Loch Fleet to investigate techniques of acidity mitigation, with a view to restoring fisheries in this and similarly affected waters. An underlying assumption of the project was that fish had been lost as a direct result of acidity and associated factors. Studies were therefore undertaken during a 2-year baseline period (1984–1986) to validate this assumption. Fish surveys using a variety of techniques (gill-netting, trapping, electrofishing) confirmed that trout were absent from the Loch and its afferent streams, and also from its main outlet stream, the Little Water of Fleet, for a distance of 7 km downstream. Trout were present below this point but are prevented from passing upstream by a 5-m waterfall. Eels, Anguilla anguilla L., were present throughout the Little Water of Fleet, though not in the Loch itself. Population densities of both species were low, with less than 7 eels and 5 trout per 100 m². Survival studies using brown trout ova and yolk-sac fry indicated that conditions in the Loch and its afferent streams were acutely toxic to these stages as a result of the low pH (pH 4.5), low calcium (I mg l^?1) and high aluminium concentrations (200 μg 1 ^?1 total Al, 60 μg 1 ^?1 inorganic monomeric Al). Trout fingerlings could survive these conditions in short-term tests (9 days) but, in chronic exposure tests lasting up to 180 days carried out in situ in streams adjoining the loch, no fish survived this period. This toxicity was eliminated in experiments where pH was raised to 5.4 by KOH addition. It is concluded that the loss of the brown trout fishery at Loch Fleet occurred as a direct result of acidity and related factors, probably acting in the first instance on the sensitive intra-gravel ova and yolk-sac fry stages, leading to recruitment failure.