Effects of Acidity and Aluminum on the Physiology and Migratory Behavior of Atlantic Salmon Smolts in Maine, USA

Atlantic salmon, Salmo salar, smolts of hatchery origin were held for 5 to 16 days in ambient (pH 6.35, labile Al = 60 µg L^?1), limed (pH 6.72, labile Al = 58.4 µg L^?1), or acidified (pH 5.47, labile Al=96 µg L^?1) water from the Narraguagus River in Maine, USA. Wild smolts were captured in the same river in rotary traps and held for up to two days in ambient river water. Osmoregulatory ability was assessed by measuring Na⁺/K⁺ ATPase activity, hematocrit, and blood Cl concentration in freshwater, and after 24-hr exposure to seawater. Hatchery smolts exposed to acidic water and wild smolts displayed sub-lethal ionoregulatory stress both in fresh and seawater, with mortalities of wild smolts in seawater. Using ultrasonic telemetry, hatchery-reared ambient and acid-exposed, and wild smolts were tracked as they migrated through freshwater and estuarine sections of the river. The proportion of wild smolts migrating during daylight hours was higher than for hatchery-reared smolts. Wild smolts remained in the freshwater portions of the river longer than either group of hatchery smolts, although survival during migration to seawater was similar for all three treatments. Acid-exposed hatchery-origin and wild Narraguagus River smolts were both under ionoregulatory stress that may have affected their migratory behavior, but not their survival for the time and area in which we tracked them.     

Physiological response of brown trout (Salmo trutta) spawners and postspawners to acidic alminum-rich stream water

High immediate postspawning mortality due to inferior autumn water quality has been hypothesized to cause juvenilization in some brown trout populations in acidified areas. We exposed male and female spawners and female postspawners from a juvenile-dominated brown trout population to acidic streamwater (pH = 4.83, Al_i = 240 μg L^?1) and a limed control (pH = 5.70, Al_i = 55 μg L^?1) for 28 days in November and December, 1984. Water chemistry was monitored at least bi-daily, and physiological stress was assessed by analysis of plasma chloride, osmolality and haematocrit. Neither pronounced physiological stress nor mortality was observed at the control site. At the exposure site trout showed significant but moderate stress responses and 15 % died. The results are discussed in terms of potential population effects and physiological mechanisms, e.g., plasma volume reduction.     

Liming acid streams: Aluminium toxicity to fish in mixing zones

Liming to neutralize acidic surface waters involves a possible risk of toxicity to fish due to precipitation or changes in speciation of Al. We report the response of captive brown trout to the experimental liming of an acid stream rich in Al. Within 15 m of lime dosing 0.22 µm filterable Al fell from 580 to 230 µg L^?1, and to 120 jig L^?1, within 30 m, though total Al was unchanged. After 24 hr, fish mortality was 100% at untreated acidic sites, 80% up to 30 m downstream of liming, declining to zero within 100 m. Mortality was 70% at 15 m below the confluence of an acidic tributary with the limed stream, despite little change in pH or total Al concentration. Mortalities were significantly correlated with concentrations of Al and Fe in gill tissues, and with 0.22 µm filterable Al and Fe in the water, but not with particulate Al or Fe. AI(OH)₄ ^?, precipitating A1 or polymeric hydrolysis products are all possible causes of the observed toxicity. Iron may have also have contributed, but the stream concentrations of this metal were relatively low. The practical conclusion is that changes in Al chemistry, where waters of differing acidity mix, may be important in some circumstances where river systems are limed selectively.     

Water quality requirement of Atlantic salmon (Salmo salar) in water undergoing acidification or liming in Norway

Atlantic salmon are severely affected by acidification in Norway. Water quality criteria for the salmon have to be based on the most sensitive stage, the smolt stage. The sensitivity to acidic water increases enormously during smolting, the seawater tolerance being especially vulnerable. Even moderately acidic water (pH about 6) with low inorganic monomeric aluminium (LA1) concentrations (<20μg. L^?1) and short-term episodes may be harmful. Mixing zones in limed or unlimed rivers may also represent a problem for seaward migrating smolts. In limed salmon rivers, the national liming goal has been increased to pH 6.5 during smolting (1 February to 1 July) and to 6.2 the rest of the year as a result of our experiments. In contrast to what has been found for brown trout, salmon strains originating from watercources undergoing acidification were not more tolerant than those from non-acidic watercourses. At the moment no such “tolerant” strains are available for restocking limed rivers in Norway.     

Base additions to flowing acidic water: Effects on smolts of Atlantic salmon (Salmo salar L.)

Acidic, Al-rich soft water (pH=5.1, Ca=1.0 mg L^?1 and labile Al=160 μg Al L^?1) was treated by addition of soda (Na ₂Co₃) and lime slurry (CaCO₃). Seven different water qualities of each type of treatment, covering the pH-range 5.1 to 8.2, were prepared in a flow-through system. Smolts of Atlantic salmon (Salmo salar L.) were used as test fish. In both types of treatment, mortality occurred at all pH-levels below 6.05. Above pH 6.05 no mortality occurred using lime slurry. Using soda, 10 % mortality occurred within 48 hr at pH above 7 due to the toxicity of aluminate at low levels of Ca. Plasma chloride levels indicated no physiological stress in the pH-range 6.45 to 7.0.     

Physiological responses of rainbow trout Salmo gairdneri exposed to plastic lake inlet and outlet stream waters

In Plastic Lake, Ontario, stocked rainbow trout (Salmo gairdneri) have failed to survive, one endemic fish species has become extinct and annual fish kills included up to five species, but especially pumpkinseeds (Lepomis gibbosus). The potential toxicity of Plastic Lake water was assessed by holding (hatchery) rainbow trout in the major inlet stream, the outlet, and in a portion of the outlet stream acidified to the pH of inlet No. 1. Stress on rainbow trout was assessed by measuring plasma and muscle concentrations of Na ⁺ Cl^?, and K⁺, plus gill A1 concentration. Trout held in Plastic Lake inlet No. 1 showed a rapid loss of plasma Na⁺ from 138 to 85 meq.L^?1and Cl^? from 120 to 75 meq.L^?1 within 24 hr. Gill A1 concentration increased from 20 to 105 μg.g^?1 dry weight. Trout held in the outlet steam showed only slight loss of plasma Na⁺ and Cl^? and no accumulation of Al on the gills. Trout held in the acidified outlet showed a significant loss of ions with plasma Na⁺ depressed from 140 to 115 meq.L^?1 and plasma Cl^? from 125 to 95 meq.L^?1over 24 hr. Gill Al concentration increased from 18 to 30 μg.g^?1 dry weight. The differences in stress response of rainbow trout held in the inlet and acidified outlet are likely due primarily to the difference in Al species concentrations in the two waters.     

Avoidance of low pH and elevated Al concentrations by Brook Charr (Salvelinus fontinalis) Alevins in laboratory tests

Laboratory studies were conducted to test the ability of brook charr (Salvelinus fontinalis) alevins, the earliest free-swimming life stage of the species, to detect and avoid toxic levels of H⁺ and inorganic Al. Alevins were tested in steep gradient choice tanks using a range of H⁺ (pH 4.0 to 5.5) and Al (0 to 500 μg L^?1) concentrations in low Ca (2.0 mg L^?1) water. The young brook charr actively avoided acidic water with a pH < 5.0. Aluminum additions of 500 μg L^?l increased the avoidance response. The observed behavioral response of alevins to low pH and elevated levels of Al, may be of significant adaptive advantage in systems undergoing acidification.     

Aluminum toxicity to fish in acidic waters

An important consequence of acidification is the mobilization of Al from the edaphic to the aquatic environment. Elevated Al levels in acidic waters may be toxic to fish. Eggs, larvae, and postlarvae of white suckers (Catostomus commersoni) and brook trout (Salvelinus fontinalis) were exposed in laboratory bioassays to pH levels 4.2 to 5.6 and inorganic Al concentrations of 0 to 0.5 mg l^?1. Aluminum toxicity varied with both pH and life history stage. At low pH levels (4.2 to 4.8), the presence of Al (up 0.2 mg l^?1 for white suckers; 0.5 mg l^?1 for brook trout) was beneficial to egg survival through the eyed stage. In contrast, Al concentrations of 0.1 mg l^?1 (for white suckers) or 0.2 mg l^?1 (for brook trout) and greater resulted in measurable reductions in survival and growth of larvae and postlarvae at all pH levels (4.2 to 5.6). Aluminum was most toxic in over-saturated solutions at pH levels 5.2 to 5.4. The simultaneous increase in Al concentration with elevated acidity must be considered to accurately assess the potential effect of acidification of surface waters on survival of fish populations.     

The Return of the Salmon

The Atlantic salmon population in the River Otra, southern Norway was lost during the 1960's due to acid rain and industrial and municipal pollution. The industrial and municipal pollution sources were sanitized by 1995. A concurrent reduction in acid deposition has during the last 10 years raised pH from 5.2 to 5.7 and reduced inorganic monomeric Al from 71 to 28µg Al L^?1 above the industrial area. The water quality improvement resulted in salmon fry again being caught from 1995. Physiological measurements (blood parameters and seawater tolerance) performed on smolts of Atlantic salmon exposed within the river during the spring of 1999 suggests that the smolts were fully smoltified and seawater tolerant, despite having moderate gill morphological changes and having moderate high gill Al concentrations (70–80 µg Al g^?1 dw). The smolt quality measured suggests that the river again can support a native salmon population, provided no negative change in water quality. Winter episodes and acid tributaries within the watershed can, however, offset the recovery process.     

Localization of aluminum in tissues of fish

The concentrations of Al in fish gills has been used as a measure of fish exposure to this metal in acidified waters. This experiment was designed to determine if other fish tissues also accumulate Al and thus possibly contribute to the cause of death. Rainbow trout (Salmo gairdneri) were exposed to the following fours test conditions for 48 hr or until death: (1) pH 6.8, <0.001 mg.L^?1 Al (n=6); (2) pH 5.2, <0.001 mg₁L^?1 Al (n=2);(3) pH 5.2,1.0 mg.L^?1 Al (n=5); (4) pH 6.8, 1.0 mg.L^?1 Al (n=3). The trout were held in synthetic, low Ca water prior to, and during, experimentation. Esophagus-stomach, gonad, gall bladder, gill (left and right), heart, intestine, kidney, liver, muscle (epaxial), and spleen were digested in a 4:1 mixture of HNO₃:HClO₄ and analyzed by Inductively Coupled Plasma Emission Spectrophotometry. Elevated Al concentrations were found in gill and gastrointestinal tissues. Left and right gills of fish exposed to pH 5.2, 1.0 mg.L^?1 Al were the only tissues found to be significantly different (p<0.01) between the test conditions. The mean total Al concentrations of these test 3 fish, for the left and right gill were 3.61 and 4.33 mg.g^?1 Al dw. The Al concentration in thle gastrointestinal tissues of the fish exposed to pH 6.8 at 1.0 mg.L^?1 Al was greater than that of the control fish, but not statistically significant. These results suggest that the analysis of whole gill remains an effective indicator of Al exposure in fishes at low pH.     

The effects of low pH,low calcium concentrations and elevated aluminium concentrations on sodium fluxes in brown trout,Salmo Trutta L.

The effects of pH (c. 7.0, 5.4, 4.5 and 4.0), nominal Al levels (0 and 8 μmol L^?1) and Ca levels (10 and 50 μmol L^?1) on Na influx, efflux and netflux of brown trout have been investigated using artificial lake water of known composition. Low pH had little effect on influx, but tended to increase efflux, particularly in the low Ca treatments. A nominal addition of 8 μmol Al L^?1 at pH 4.5 and 4.0 reduced influx significantly. Efflux was unaffected. Aluminium addition at pH c. 7.0 and 5.4 had no such effect. The measured Al concentrations at the end of the static 8 hr flux measuring experiments were markedly lower than the nominal amount of A1 added to the start.     

Response of lake trout (Salvelinus namaycush) and brook trout (S. fontinalis) to surface water acidification in Ontario

Netting surveys of lakes varying in pH (4.4–7.1) showed that lake trout (Salvelinus namaycush) populations fail to recruit at pH <5.5 and are lost from lakes with pH<5.2. Brook trout (S. fontinalis) were extirpated in lakes with pH <5.0. In regional chemical surveys of Ontario lakes, it was found that 2% of sampled brook trout lakes and 2.5% of lake trout lakes were acidified (alkalinity <0 uEq L^?1). Threshold pH levels determined from fisheries assessments were used to estimate that 1% of lake trout and brook trout populations have been lost due to acidification.     

Increased mortality of fish due to changing Al-chemistry of mixing zones between limed streams and acidic tributaries

The present study is mainly focusing on mortality variations of fish due to changing Alchemistry of mixing zones. An artificial mixing zone was made by pumping water from a limed stream and an acidic tributary into a mixing channel. Atlantic salmon (Salmo salar L.) parr were exposed to the mixed water, limed stream water, and acidic tributary water. Mortality, blood haematocrit and plasma Cl^?-concentration were recorded. Neither mortality, nor changes in haematocrit and plasma Cl^? were observed when fish were exposed to limed water, while in both acidic and mixed water, mortalities and loss of plasma Cl^? were observed. The highest mortality rates were found within the initial part (0 to 20 s) of the mixing zone. Blood haematocrit increased only in fish exposed to acidic tributary water. Our results shows that changes in Al-chemistry and subsequent Al-polymerization occur when acidic tributary water is mixed with limed stream water. We have also demonstrated that the toxicity which can arise in mixing zones are greater than in the original acidic water before mixing. The variations in mortality observed are associated with the quality and quantity of Al-polymerization as well as ageing of the polymers.     

Field and laboratory studies of exposures of brown trout to acid waters

Some recent work on the effects of acid waters on brown trout are presented. Laboratory bioassay experiments have demonstrated that yearling trout are relatively insensitive to pH >4.3. Aluminium is demonstrated to be extremely toxic with suppression of growth occurring at concentrations above 20 μg L^?1 at pH 4.4 to 5.2. Aluminium toxicity is reduced at high pH (5.9 and 6.3). Field studies carried out on 61 acidic and circumneutral streams in upland areas of England and Wales showed a strong relationship between water quality and standing crop of 1+ brown trout. Measured pH levels per se were too high to be directly toxic. On the other hand, heavy metal and Al concentrations could account for low or zero brown trout biomass in the more acidic streams. A mobile bioassay laboratory has been developed to allow controlled bioassay experiments to be carried out in the field. Natural and synthesised waters can be tested concurrently in multi-factorial experiments with in situ determinations of pH, Ca, Al (total and monomeric) and other water quality characteristics.     

Ecological and physiological responses of Atlantic salmon in acidic organic rivers of Nova Scotia,Canada

Ecological and toxicological data from field studies on acidic rivers of Nova Scotia were examined to review the effects of low pH on Atlantic salmon (Salmo salar) populations in waters rich in organic acids where noexchangeable forms of Al dominate at all times. There were no survival of salmon past the dry stage at pH <4.7, and survival rates for salmon from egg to smolt only increased at pH >4.9. Annual production of juvenile salmon and potential yield of smolts were lower at pH 4.7 to 5.4 than at pH 5.6 to 6.3 because of reduced densities attributable to the high mortality of fry at pH ≤5.0. However, acidity episodes to pH <4.7 also resulted in mortality of parr, reducing densities and often completely eliminating year-classes. The physiological responses of juvenile salmon to chronic acid conditions and to acute acidity typical of episodic events were also reviewed in relation to toxicity. Decreased in plasma Na and Cl were well correlated with ambient pH, but not with exchangeable Al concentrations in rivers. These plasma electrolytes provided reliable indicators of the thresholds for sublethal effects on ionoregulatory mechanisms. There was no morphological evidence of damage or lesions in gill epithelia, indicating that accumulation of Al in the gills of parr was not a significant factor in the lethal effects observed in acidic rivers. High organic matter content in the water apparently protected gills from adverse Al effects. Toxicity was considered to result from the effect of low ambient pH on branchial ionoregulatory mechanisms.     

Acute and sub-chronic toxicity of lead to the early life stages of smallmouth bass (Micropterus Dolomieui)

The early life stages of smallmouth bass (Micropterus dolomieui) were exposed to Pb in acute (96 hr) and sub-chronic (90 day) bioassays (water hardness = 152 mg L^?1 as CaCO₃). After 96-hr static exposures at nominal Pb concentrations up to 15.9 mg L^?1, eggs and sac fry showed no increased mortality over that in controls. Swim-up fry (96-hr LC₅₀ = 2.8 mg Pb L^?1) were more sensitive to Pb exposure than were fingerlings (96-hr LC₅₀ of 29.0 mg Pb L^?1 ). The relation between dissolved Pb and mortality was non-significant for either swim-up fry or fingerlings. Fingerlings were exposed to Pb concentrations as high as 405 μg L^?1 for 90 day to evaluate effects on substrate selection, locomotor activity, hematology, and weight. Dark or light substrate selection (cover-seeking) and locomotor activity, weight and hemoglobin concentration in the blood were not significantly altered by any treatment. Hematocrit and leucocrit varied significantly but not in relation to Pb levels. Sub-chronic Pb exposure did not appear to represent a threat to smallmouth bass in waters of medium hardness and above-neutral pH (7.1 to 7.9).     

The effects of liming on juvenile stocks of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in a Norwegian river

The effects of liming on juvenile stocks of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in the river Vikedalselva in southwestern Norway were assessed. From 1987 to 1989, the river was limed only during the spring snow melt, and pH varied in the range between 5.5 and 7.0. In 1990 to 1993, the river was limed to pH 6.2 from 15 February to 1 June and to pH 5.7 during the rest of the year. Since 1994, the pH during late winter and spring was maintained above 6.5. Prior to liming fish kills were evident during spring snow melt, but these have not occurred since liming. Electrofishing in the autumn between 1981 and 1994 showed no significant change in densities of juvenile salmon and brown trout after liming, mean densities ranged between 19–50 and 9–32 individuals 100 m^–2 respectively. A significant linear correlation between production and biomass of both species was found, indicating that factors directly controlling density affect juvenile production and cause production to remain below carrying capacity. In spite of a clear increase in pH and a reduction in the concentration of labile aluminium after liming, the conditions still do not seem to be optimal for juvenile salmonids. We suggest that a complexity of different factors impose limitations on fish production in the river: inadequate egg deposition, environmental factors such as water temperature and flow, osmoregulatory failure in mixing zones between limed and acidic water and gill damage through deposition of aluminium and iron. However, there are several indications of a reduction in toxic effects after the pH was raised to 6.5 during spring snow melt.     

Soybean root growth in relation to ionic composition in magnesium-amended acid subsoils: Implications on root citrate ameliorating aluminum constraints

Abstract

Hydroponic studies with soybean (Glycine max [L.] Merr.) have shown that µmol L^?1 additions of Mg²⁺ were as effective in ameliorating Al rhizotoxicity as additions of Ca²⁺in the mmol L^?1 concentration range. The objectives of this study were to assess the ameliorative effects of Mg on soybean root growth in acidic subsoils and to relate the soil solution ionic compositions to soybean root growth. Roots of soybean cultivar Plant Introduction 416937 extending from a limed surface soil compartment grew for 28 days into a subsurface compartment containing acid subsoils from the Cecil (oxidic and kaolinitic), Creedmoor (montmorillonitic) and Norfolk (kaolinitic) series. The three Mg treatments consisted of native equilibrium soil solution concentrations in each soil (50 or 100 µmol L^?1) and MgCl₂ additions to achieve 150 and 300 µmol L^?1 Mg (Mg150 and Mg300, respectively) in the soil solutions. Root elongations into Mg-treated subsoils were compared with a CaCO₃ treatment limed to achieve a soil pH value of 6. Subsoil root growth responses to the Mg treatments were less than for the lime treatments. Root length relative to the limed treatments for all subsoils (RRL) was poorly related to the activity of the soil solution Al species (Al³⁺ and Al-hydroxyl species) and Mg²⁺. However, the RRL values were more closely related to the parameters associated with soil solution Ca activity, including (Ca²⁺), (Al³⁺)/(Ca²⁺) and (Al³⁺)/([Ca²⁺] + [Mg²⁺]), suggesting that Ca could be a primary factor ameliorating Al and H⁺ rhizotoxicity in these subsoils. Increased tolerance to Al rhizotoxicity of soybean by micromolar Mg additions to hydroponic solutions, inducing citrate secretion from roots to externally complex toxic Al, may be less important in acid subsoils with low native Ca levels.     

Population dynamics of brook trout (Salvelinus fontinalis) during maintenance liming of an acidic lake

Maintenance liming of an acidic lake in the Adirondack Mountains of New York state (Woods Lake) was conducted three times over a 5 yr period in an attempt to establish a self maintaining brook trout population. Various strains and age classes of marked brook trout were stocked annually and the population was inventoried semi annually to evaluate survival, growth, and reproductive success. The Woods Lake brook trout population was dominated by young, stocked fish throughout the maintenance liming period of 1985-89. Based on spring emergent fry trap catches and fall trap net catches of unmarked fish, only one naturally produced year class (1986) was successfully recruited to the Woods Lake brook trout population. Low annual survival rates (G 20%) of juvenile trout were observed throughout the study period. Although initial growth rates and condition of young trout were satisfactory, increased intraspecific competition for food resulted in declining growth rates and condition of older age classes. Fall standing crops of brook trout remained at relatively low levels of 6 to 10 kg ha^?1 and both production per unit biomass and growth efficiency decreased over the 5-yr. Repeated whole lake liming and limited spawning habitat improvement were not sufficient to sustain brook trout natural reproduction in Woods Lake. Low productivity, marginal spawning habitat quality, and low survival rates of stocked trout in Woods Lake resulted in the failure to establish a self maintaining, productive brook trout population in Woods Lake.     

On the prediction of acid precipitation events and their effects on fishes

It is possible to predict acid rain events and melts of acid snow some 12 to 24 hr in advance, including estimation of the magnitude and duration of such events. This is sufficient notice to permit monitoring of stream chemistry and fish plasma and muscle ions before acid stress, and to continue this monitoring throughout and after specific events. Such a program has been in place for 2 yr in waters tributary to the Milford Bay Trout Hatchery, Ontario. During one snow melt in February 1984 surface waters showed a decline to pH 4 and associated negative ANC. Rainbow trout held in such water lost plasma Na and Cl rapidly and died within 28 hr. The hatchery water supply, consisting of a mixture of spring and surface water, showed a decline in alkalinity from 300 to 30 μeq.L^?1, and a pH change from 6.6 to 5.4, during snow melt. Total A1 concentration increased from 42 to 222 μg.L^?1 during snow melt with the “reactive” component increasing from 17 to 112 μg.L^?1. Rainbow trout held in this water did not show physiological stress. More rapid run-off of melt water could be expected to exhaust all of the alkalinity in the hatchery water supply permitting the pH to decline and A1 concentration to rise to levels lethal to the hatchery stock of rainbow trout.