Embryonic and larval development of brown trout, Salmo trutta L.: exposure to aluminium, copper, lead or zinc in soft, acid water

Freshly fertilized ova, eyed ova and yolk-sac fry of brown trout, Salmo trutta L., were exposed to each of four trace metals (aluminium: 6000 nmol l^?1; copper: 80 nmol l^?1; lead: 50 nmol l^?1; zinc: 300 nmol l^?1) while held in flowing artificial soft-water media maintained at pH 4.5 or 5.6 and [Ca] 20 or 200 μmol l^?1. In continuous exposure from fertilization, survival of ova was severely affected at pH 4.5 and [Ca] 20 μmol l^?1, regardless of the presence of Cu, Pb or Zn; Al reduced embryonic mortality and improved hatching success. High ambient [Ca] at pH 4.5 increased egg survival. At ‘swim-up’, surviving fry exposed to Al or Pb had lower whole body Ca, Na and K content, irrespective of pH or ambient [Ca]. Cu reduced whole body Ca and K content at pH 5.6 and [Ca] 200 μmol^?1, and whole body Ca, Na and K content in the other media. Zn reduced whole body mineral content at pH 5.6 and [Ca] 20 μmol l^?1. Whole body Mg content was reduced by all trace metals at pH 5.6 and [Ca] 20 μmol l^?1, and by Cu at pH 5.6 and [Ca] 200 μmol l^?1. Al and Cu impaired skeletal calcification at pH 5.6 at both ambient [Ca]; Pb only at [Ca] 20 μmol I^?1. Zn enhanced calcification at pH 4.5 and [Ca] 200 μmol l^?1. In the absence of trace metals, low pH reduced body Ca, Na, K content and skeletal calcification at [Ca] 200 μmol l^?1. The uptake of Ca, Na and K, measured at regular intervals from hatching was impaired to the same extent by all treatments at pH 4.5, irrespective of ambient [Ca] or trace metal presence. At pH 5.6, irrespective of ambient [Ca], Al, Cu and Pb impaired Ca and K uptake. The rate of Na uptake was reduced by Al and Cu. Al-treated yolk-sac fry, exposed to low ambient [Ca] from 200–300° days post-hatch, suffered high mortalities regardless of pH. Ca, Na and K uptake was impaired by all treatments at pH 4.5, and by Al and Cu at pH 5.6 in a similar exposure period. The development of the early stages of brown trout in the presence of trace metals is discussed in relation to recruitment failure in areas of soft, acid water.     

The effects of eight trace metals in acid soft water on survival, mineral uptake and skeletal calcium deposition in yolk-sac fry of brown trout, Salmo trutta L.

Recently hatched fry of brown trout were exposed to each of eight trace metals (Al, Cd, Cu, Fe, Mn, Ni, Pb, Zn) and to a mixture of all eight, at concentrations typical of soft acid waters, in flowing artificial soft water medium at pH 4·5 and 6·5, for 30 days. At pH 4·5, in the absence of trace metals, net uptake of Ca, Na and K and skeletal calcification were impaired but mortalities were low (10%). At pH 4·5, in the presence of Al, Cu, Pb and Zn individually and the mixture, mortalities were high (87–100%). Cd, Fe, Mn and Ni at pH 4·5 each caused some mortalities (17–43%) and impaired skeletal calcification. At pH 6·5, each of Al, Cd, Cu, Fe, Mn, Pb and the mixture of metals impaired net Ca uptake and (except Al, Fe and Pb) skeletal calcification. Net K uptake was impaired by Al and by the mixture, and less severely by Cu and by Fe. Net Na uptake was impaired by Al and by the mixture, and less severely by Fe and by Mn. The role of trace metals other than Al in fisheries'decline in soft acid waters is discussed.     

Effects of six trace metals on calcium fluxes in brown trout (Salmo trutta L.) in soft water

Summary Calcium fluxes were measured simultaneously in brown trout fry maintained in an artificial soft water medium of [Ca] 20 mol·l^-1 and pH 5.6, and exposed to each of six trace metals (Al, Cu, Fe, Ni, Pb, and Zn). The trace metal concentrations represented typical and maximum levels found in acid waters experiencing declining fishery status. In the absence of trace metals, evidence is presented which suggests that ca. 91% of Ca taken up from the external medium was by extraintestinal active transport. Calcium efflux was stimulated by both concentrations of Al, Cu, Fe, and Pb. Efflux was also stimulated by [Ni] 170 nmol·l^-1 and [Zn] 3000 nmol·l^-1. In some cases, response to increased efflux was stimulation of influx. Lack of stimulation of influx resulted in negative net Ca fluxes. Net Ca losses were recorded at both concentrations of Al, Pb, and Ni, lower concentrations only of Fe, and higher concentrations only of Cu and Zn.Abbreviations J _in influx - J _net net flux - J _out efflux Henceforward in this paper, chemical elements are referred to by their chemical symbols rather than by full names     

Initial feeding in migratory brown trout (Salmo trutta L.) alevins

Initial feeding of brown trout was investigated under laboratory conditions. Fifty per cent feeding occurred when yolk constituted approximately 31% of total alevin dry weight, and feeding rate was positively correlated to developmental stage. The possible ecological implications of initial feeding are discussed.     

Interpopulation variation in reproductive traits of anadromous female brown trout, Salmo trutta L.

We studied reproductive traits in nine anadromous brown trout, Salmo trutta L., populations in seven Norwegian rivers. Within populations we found a positive significant correlation between fish length and fecundity in all populations, and between fish length and egg diameter in five populations. There were significant differences in these relationships between populations from different rivers, and between populations from different locations within rivers. When adjusted for variation in fish length, mean fecundity and mean egg diameter showed a negative significant correlation among populations. The ratio of gonadal weight to somatic weight (gonadosomatic index) varied significantly among populations but was not associated with variation in fish length. Comparatively few large eggs were found in brown trout populations co-existing with several other fish species.     

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.     

Cortisol and thyroid hormone responses to acid stress in the brown trout, Salmo trutta L.

Individual cannulated brown trout monitored during exposure to acidic water showed increased plasma cortisol after 3 h at pH 4.0 with low (0.05 mm) or high (2.8 mm) calcium (Ca) content, and after 2 days in acidic water with a high Ca content. Most fish did not survive for 2 days in acidic water with a low Ca content. Non-cannulated fish showed a similar increase in mean plasma cortisol after 2 days in high-Ca acidic water (pH 4.0–4.6), but not in acidic water of a low Ca content. After 7 days of exposure to acidic water, plasma cortisol appeared to recover when there was a high Ca content but increased 20-fold when Ca content was low. In cannulated fish severe acid stress resulted in a marked and rapid thyroid response. Plasma thyroxine (T₄) was elevated after 3 h exposure to acidic water of both low and high Ca content and remained elevated for 2 days of acid exposure with high Ca. In non-cannulated fish an increase in mean T₄ was apparent only after 7 days in low-Ca acidic water. Plasma triiodothyronine (T₃) levels were not significantly altered by any of the acid regimes. Plasma glucose of cannulated fish was elevated within 3 h of acid-exposure and remained elevated after 2 days in high-Ca acidic water.     

The effect of calcium concentration on the toxicity of copper, lead and zinc to yolk-sac fry of brown trout, Salmo trutta L., in soft, acid water

Yolk-sac fry of brown trout were exposed to three levels of single trace metals (Cu, 20,40,80 nmol 1^-1; Pb, 12·5,25,50 nmol 1^-1; Zn, 75,150,300 nmol 1^-1) typical of concentrations reported for acid soft waters, in flowing, artificial, soft water media maintained at pH 4·5 and [Ca] of 20 or 200 μmol 1^-1for 30 days.
Mortalities were high in fry subjected to all levels of the three trace metals at [Ca] 20 μmol 1^-1, with 80% of the total deaths occurring between days 11 and 15 of the experiment. 25% mortality occurred at low [Ca] and pH 4·5 in the absence of trace metals, with only one death recorded at [Ca] 200 μmol1^-1'(Cu, 80 nmol 1^-1). At high [Ca] all three levels of Cu and Pb impaired net Na and K uptake; Cu was the only metal to reduce the uptake of Ca. The Zn treatments had no significant effect on mineral uptake. Calcification of centra was reduced by all three Cu treatments at [Ca] 200 μmol 1^-1. The lowest Zn concentration (75 nmol 1^-1) was the only other treatment to impair skeletal development. In the absence of trace metals, low [Ca] significantly reduced Ca, Na and K uptake, skeletal calcification and dry mass at pH 4·5.
The deleterious effects of low Cu, Pb and Zn concentrations at low pH and low [Ca], and the ameliorative effect of higher ambient [Ca], are discussed in relation to fishery status in soft, acid waters.     

Body shape and robustness response to water flow during development of brown trout Salmo trutta parr

Domesticated brown trout Salmo trutta parr were subjected to increased, variable flow under controlled experimental conditions. Using geometric morphometric analyses, (a mass–length index) and caudal fin area–body length ratio, this study assessed morphological responses in lateral body depth, growth and robustness and propulsive potential, respectively, of parr over the course of 32 weeks. Geometric morphometric analyses did not reveal an effect of exercise on either lateral body depth or caudal fin area. However, improved overall robustness and growth trajectories in exercised parr showed a positive adaptive response to the enriched habitat. Exercise and habitat heterogeneity thus have the potential to improve survivability of domesticated salmonids in the wild.     

Wild adult hybrids of Salmo salar L. and Salmo trutta L.

Ovarian development was impaired in three adult Salmo salar L. × S. trutta L. hybrids identified among adult salmonids in Scottish fisheries. Species-specific variation at enzyme loci indicated that the fish were F1 hybrids and mitochondrial DNA analysis showed them to be the progeny of S. salar females.     

Seasonal and spatial microhabitat selection and segregation in young Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Norwegian river

Seasonal microhabitat selection by sympatric young Atlantic salmon and brown trout was studied by diving. Both species, especially Atlantic salmon, showed seasonal variation with respect to surface and mean water velocities and depth. This variation is partly attributed to varying water flows and water temperatures. In winter the fish sought shelter in the substratum. A spatial variation in habitat use along the river due to different habitat availabilities was observed. Both species occupied habitats within the ranges of the microhabitat variables, rather than selecting narrow optima. It is hypothesized that the genetic basis allows a certain range to the behavioural response. Microhabitat segregation between the two species was pronounced, with brown trout inhabiting the more slow-flowing and partly more shallow stream areas. Atlantic salmon tolerated a wider range of water velocities and depths. Habitat suitability curves were produced from both species. It is suggested that habitat suitability curves that are based on observations of fish occupancy of habitat at median or base flow may not be suitable in habitat simulation models, where available habitat is projected at substantially greater water flows.     

Morphological variability among seven populations of brown trout, Salmo trutta L., in Greece

Morphological variation in five meristic and 21 morphometric characters was examined in the brown trout, Salmo trutta L., from seven distinct streams and rivers from N and NW Greece. Variation within each sample, estimated as the multivariate generalization of the coefficient of variation, did not vary substantially among the populations. The phenetic relationships of these populations were examined using the stepwise discriminant analysis. The taxonomic implications of these results are discussed.     

The post‐spawning movements of migratory brown trout Salmo trutta L.

The post spawning behaviour of sea trout Salmo trutta was studied over a 2 year period in the river and estuary of the River Fowey, south‐west England. Forty‐five sea trout kelts were trapped immediately after spawning in December and intraperitoneally tagged with miniature acoustic transmitters. The subsequent emigration into coastal waters was monitored using acoustic receivers deployed throughout the river catchment. The levels of gill Na⁺K⁺ATPase activity in sea trout kelts sampled at the same time as the tagged fish were within the range of 2·5 to 4·5 μmol Pi per mg protein per h indicating that the post‐spawning fish were not physiologically adapted to salt water. The tagged kelts were resident in fresh water between 4 and 70 days before entering the estuary. Sixty two per cent of the tagged kelts subsequently migrated successfully into coastal waters, with a higher success rate for male fish (75%) than females (58%). There was a significant size related difference in the run‐timing of the kelts with the larger fish moving more quickly into coastal waters after spawning than smaller fish. Seaward migration within fresh water was predominantly nocturnal and generally occurred in conjunction with increasing river discharge and rising water temperature. Migration through the estuary continued to be predominantly nocturnal and occurred during an ebbing tide. Residency within the estuary varied amongst individuals although it was invariably short, with most fish moving out into coastal waters within one to two tidal cycles. Five tagged kelts returned from the coastal zone and re‐entered fresh water during April and June. Marine residence time varied between 89 and 145 days (mean 118 days) and the minimum estimated marine survival was c. 18%. One of these sea trout was subsequently recaptured after successfully spawning in the vicinity where it had been previously tagged demonstrating a degree of spawning site fidelity.     

Mitochondrial DNA sequence variation and phylogeography among Salmo trutta L. (Greek brown trout) populations

To investigate the phylogenetic relationships and geographical structure among brown trout S. trutta L. populations from the South Adriatic-Ionian and Aegean sea basins, mitochondrial DNA nucleotide sequence comparisons were used. A 310-base-pair (bp) segment of the control region (D-loop), and an additional 280-bp segment of the cytochrome b gene were sequenced from representatives of 13 brown trout populations. Phylogenetic analyses, conducted after combining the data presented with published data from other Eurasian brown trout, revealed four major phylogenetic groups, three of which were found widely distributed within the southern Balkan region. The phylogeographical patterns revealed by mtDNA represent one of the few cases where phylogenetic discontinuity in a gene tree exists without obvious geographical localization within a species' range and has most likely resulted from the differentiation of the major mtDNA clades during Messinian or early Pleistocene times. Finally, the genetic relationships among the populations suggested by mtDNA were generally not in accordance with either allozyme or morphological data.     

Episodic exposure to acid and aluminium in soft water: survival and recovery of brown trout,Salmo trutta L.

Yolk-sac fry, swim-up fry and 1–2 yr juveniles of brown trout, Sulmo trutta L., were exposed to episodes of aluminium and low pH, maximum aluminium concentration 12 μmol l⁻¹ (323 μg l⁻¹), minimum pH 4.5, total duration up to 54 h (yolk-sac fry) or up to 78 h (swim-up fry and juveniles), in an artificial soft water medium, [Ca] 20 μmol l⁻¹ (0–8 mg l⁻¹) (nominal baseline: pH 5.6, zero aluminium concentration). Yolk-sac fry mortality was nil or very low. A marked increase in susceptibility, with high mortalities, occurred when the yolk was fully absorbed. Mortality of juveniles exposed to two successive episodes was lower than would have been expected on the basis of comparisons with mortalities in single episodes, and mortality declined as the interval between the two episodes was increased. Disturbance of sodium, potassium or calcium balance or gill damage in surviving yolk-sac fry or juveniles was still evident 5 to 6 days after the end of a single episode.     

Endocrine responses of brown trout, Salmo trutta L., to acid, aluminium and lime dosing in a Welsh hill stream

Brown trout caged in an acidic, softwater, Welsh hill stream running through close-canopy conifer forest and in an adjacent, circumneutral, moorland stream were studied after a natural rainfall event and after experimental acid and aluminium dosing of the forest stream. Endocrine (cortisol and thyroxine) and metabolic (glucose) stress responses occurred in fish held in the forest stream. Liming downstream of the acid zones mitigated these responses.     

Microhabitat use by brown trout, Salmo trutta L. and Atlantic salmon, S. salar L., in a stream: a comparative study of underwater and river bank observations

Two methods, visual observation from the river bank and visual observation underwater by diving, were compared for microhabitat studies in young brown trout and Atlantic salmon in a stream. A wide range of habitat conditions were surveyed. Each method yielded different results with respect to microhabitat use. River bank observations missed small fish under surface turbulence and in deeper waters. Underwater observations missed small fish in shallow areas.     

Effects of low pH on the chorion of rainbow trout, Oncorhynchus mykiss, and brown trout, Salmo trutta f. fario

Morphological studies on the effect of low pH on egg shells (chorion) of rainbow trout, Oncorhynrhus mykiss Walbaum (formerly Salmo gairdneri ), and brown trout, Salmo trutta f. fario L., showed that the plugs of the outer layer of the chorion, which block the channels of the underlying layer, developed distinctive vacuolization at low pH exposure. This may lead to loss of chorional semi-permeability. Chorions were incompletely degraded at low pH inducing hatching problems. Partially hatched alevins showed only a small local zone of dissolution of the chorion around the head of the embryo preventing the larvae from being freed.