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.     

Atlantic salmon resources in the Northeastern United States and the potential effects of acidification from atmospheric deposition

The Atlantic salmon (Salmo salar) was formerly abundant in northeast coastal rivers in the United States from the Canadian border to the Connecticut River, and possibly as far south as the Delaware River. It was eliminated from most of its former range by a combination of overfishing, construction of dams impassable to migrating fish, and municipal and industrial pollution. Reproducing populations are now limited to a few rivers in Maine, but attempts are under way to reintroduce the species to some rivers where populations formerly existed. Most of the native Atlantic salmon rivers are low in acid neutralizing capacity and receive acidic precipitation. The third order streams are not now acidic; however, in some first and second order streams in Maine, pH episodically declines to 4.7 and Al increases to 350 μg g^?1. These conditions could be toxic to sensitive early life history stages of Atlantic salmon. Comparison of chemical conditions in two Maine rivers in 1980–1982 with those in 1969–1970 indicated that the streams have not become more acidic during this interval. Data on the sport catch of Atlantic salmon indicated that populations have generally remained stable or recently increased in the Maine rivers having naturally reproducing populations. The recent increase probably resulted from introductions of hatchery smolts to supplement natural reproduction, and the occurrence of strong year classes in 1978 and 1980. The population in one stream has declined significantly in recent years, but the cause of the decline is probably not related to acidic precipitation. Atlantic salmon resources in the U.S. have apparently not been adversely affected by atmospheric deposition at the present time.     

Water Quality Dependent Recovery from Aluminum Stress in Atlantic Salmon Smolt

In Norway, a variable pH target (pH 6.2–6.4 during most of the year, but 6.4 during the smoltification period) is used to reduce the cost of liming salmon rivers. Here we test the adequacy of this liming strategy. Atlantic salmon presmolts exposed to sublethal acidic water (pH 5.9, <25 µg Ali·L^?1) for more than 3 months showed impaired sewater tolerance, elevated gill-Al concentrations, severe gill tissue changes, elevated blood plasma glucose concentrations, but no effect on blood plasma chloride. It is usually assumed that smolt will recover from prior aluminum (Al) exposure if water quality is restored. Recovery rate is here used as an indirect measure of water quality improvements achieved after treating acid water (pH 5.8, 85 µg Ali·L^?1) with lime to reach pH-target levels of 6.0 – 6.3. Fish were exposed in a channel-tank set-up for >210 hrs in water aged from 1 minute up to 2 hours after treatment (in a flow through system). More Al was eliminated from the gills when the fish were exposed to pH 6.3 than to pH 5.8 or 6.0, and when water was aged after pH increase. Recovery, defined as return of normal gill morphology, blood homeostasis and establishment of seawater tolerance was achieved within 210 hrs in channels treated with lime to pH 6.3, while a similar recovery was not obvious at lower pH values. Liming to pH 6.3 detoxified Al better than pH 6.1.     

Acid production,acid tolerance and growth rate of Lotus rhizobia in laboratory media

Twenty-seven strains of Lotus rhizobia were tested for acid tolerance in yeast-extract mannitol broth (pH 4.6) by multiplication from low initial cell densities. Acid production on unbuffered yeast-extract mannitol agar slopes incorporating bromothymol blue indicator was also determined. All slow-growing, alkali-producing strains were acid sensitive and four of the six fast-growing, acid-producing strains were acid tolerant as was one fast-growing, alkali-producing strain. The method of testing for acid tolerance proved suitable for fast-growing strains and results are discussed in relation to the ecological implications of acid and alkali production by rhizobia.     

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.     

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.     

Physiological responses to acid stress of an acid-soil tolerant and an acid-soil sensitive strain of Rhizobium leguminosarum biovar trifolii

Physiological responses to acid stress in two strains of Rhizobium leguminosarum bv trifolii of differing acid-soil tolerance were compared. Acidity affected the size and morphology of the acid-tolerant strain, WSM409, but not of the acid-sensitive strain, TA1. Acid grown cells of WSM409 and TA1 had less cell-associated Ca and Mg and more P than cells grown at pH 7.0. Potassium content was lower in acid grown cells; WSM409 was less affected by pH than that in TA1. WSM409 was more tolerant of pH shock at pH 3.5 when grown at pH 4.8 than when grown at pH 7.0. TA1 was more sensitive to pH shock when grown at pH 4.8 than when grown at pH 7.0. WSM409 shows a characteristic adaptive acid tolerance response, whereas TA1 shows an acid sensitive response.     

Ecological effects of acidification on tertiary consumers. Fish population responses

Acidification effects fish populations in various ways, dependant on several factors of biotic and abiotic nature. The most important abiotic factor is water quality and its variability through season and during episodes. Among important biotic factors are fish species, development stages and spawning strategy. Also the specimens residence at time of toxic water and their ability to seek areas of better quality (refuges), are of main importance. For inland populations, reproduction failure seems to be the most important factor for population losses. Which lifestage being most affected might, however, differ from one population to another, even among the same species. Eggs and. alevins have been considered to be the most sensitive stages, although high mortality of postspawners have been reported from several lakes. Anadromous species, like Atlantic salmon, have stages of development that are particularly sensitive to changes in water quality. These stages are connected to physiological changes when preparing for change in salinity (smoltification and return from sea). Unfortunately these physiological changes coincides with the most critical environmental periodes; spring melt and autumn rain.     

Monitoring fish stocks in relation to acidification in Norwegian watersheds

The Norwegian Monitoring Programme for Long-Range Transported Air Pollutants started in 1980. The biological part of this programme includes besides invertebrate studies in streams, (i) fish community status in lakes by means of interviews, test-fishing in lakes by using standard gill-net series, recruitment studies of brown trout in inland streams, and juvenile stock assess and monitoring of fish kills in salmon rivers. Damaged fish stocks are recognized within a land area of 51,500 km² in southern Norway and 30 km² in northern Norway. At least 6,000 lake-dwelling fish stocks have either been lost or are at various stages of reduction. Brown trout (Salmo trutta) is the most widespread and abundant species of fish in Norwegian watersheds, and is also most severe affected by acidification. More recently, there are some indications of an increase in the abundance of brown trout in some areas. However, analysis of age structure in lakes, and fry densities in streams in such areas revealed large annual variations in recruitment rate, which indicates unstable water chemical conditions. Atlantic salmon (Salmo salar) is virtually extinct in 25 rivers in southernmost, southwestern and western Norway.     

不同春小麦品种耐低磷性评价及种质筛选

筛选磷高效作物是充分利用土壤磷素和减少磷肥施用量的重要手段。本研究以162份春小麦种质资源为材料,对其苗期的株高、总根长、根表面积等8个指标的耐低磷系数进行分析,采用隶属函数法综合评价春小麦苗期的耐低磷特性,初步筛选耐低磷材料,并进一步进行成株期的耐低磷特性鉴定,筛选出耐低磷材料和磷敏感材料,分析其在低磷下酸性磷酸酶的活性变化。结果表明,低磷胁迫下春小麦材料苗期和成株期的各性状均受到不同程度的影响,并随着胁迫程度的增加,小麦生长受抑制程度增强。通过主成分分析将苗期8个指标转化成4个综合指标(累计贡献率为82.60%),将成株期的10个指标转化为3个综合指标(累计贡献率为83.23%);采用隶属函数法计算耐低磷综合评价值(D)值,对D值进行聚类分析,将苗期的162份春小麦种质资源划分为耐低磷型(10份)、较耐低磷型(26份)、低磷较敏感型(91份)、低磷敏感型(35份)4类。选取耐低磷型材料(5份)和低磷敏感型材料(4份),进一步进行成株期鉴定,最终筛选1份耐低磷材料wp-35和1份磷敏感材料wp-119。通过分析其酸性磷酸酶活性,发现在低磷胁迫下春小麦根系和叶片中的酸性磷酸酶活性均升高,且耐低磷材料的酸性磷酸酶活性高于磷敏感材料。本研究结果可为解析春小麦耐低磷特性、培育耐低磷品种提供种质资源和理论依据。     

Intra‐specific variation for salt tolerance in a potential oil‐seed crop,brown mustard (Brassica juncea (l.) Czern. and Coss.)

Thirty eight accessions of brown mustard (Brassica juncea (L.) Czern. and Coss.) were screened after two weeks growth in solution culture containing 120 mol m‐³ NaCl. Considerable variation for salt tolerance was observed in this set of germplasm, since some accessions showed relatively vigorous growth in saline medium.

In order to determine the consistency of degree of salt tolerance at different growth stages of crop life cycle two salt tolerant accessions, P‐15 and KS‐51 and two salt sensitive 85362 and 85605 were tested at the adult stage in 0(control), 100 and 200 mol m^‐3 NaCl. Both the tolerant accessions produced significantly greater fresh and dry biomass and had considerably higher seed yield than those of the salt sensitive accessions. Analysis of different ions in the leaves showed that salt tolerant accessions contained greater amounts of Na⁺, K⁺ and Ca²⁺ than the salt sensitive accessions, although they did not differ significantly for leaf Cl^‐. Only one salt tolerant accession P‐15 had greater leaf K/Na ratio and K⁺ versus Na⁺ selectivity compared with the tolerant KS‐51 and the two salt sensitive accessions.

From this study it was established that there is a considerable variation for salt tolerance in B.juncea which can be exploited by selection and breeding for improvement of its salt tolerance. Since the degree of salt tolerance in B.juncea does not change at different growth stages of the crop life cycle, selection for salt tolerance at the initial growth stages could provide individuals that would be tolerant at all other growth stages. Accumulation of Na⁺, K⁺ and Ca²⁺ in the leaves are important components of salt tolerance in B.juncea.     

Symbiotic effectiveness of Bradyrhizobium japonicum in acid soils can be predicted from their sensitivity to acid soil stress factors in acidic agar media

In acid soil, low pH, reduced availability of nutrients, and toxicity of Al and Mn limit plant growth and the survival and effectiveness of rhizobia. The symbiosis between legumes and rhizobia is particularly sensitive to acid soil stress. A pot experiment evaluated whether Bradyrhizobium japonicum strain growth on acidic agar media would predict ability to colonize the rhizosphere and form effective nodules in acidic soils. Three Indonesian strains of B. japonicum with similar effectiveness at neutral pH in sand culture but with different tolerance of acid soil stress factors in agar media, and an acid-tolerant commercial strain (CB1809) of comparable effectiveness, were tested in three acid soils using the Al tolerant soybean (Glycine max cv PI 416937). At 7 days after inoculation all strains had achieved large rhizosphere populations, but by day 14 the rhizosphere population of the acid-sensitive strain had decreased, while the more acid-tolerant strains increased. The acid-tolerant strains had significantly greater nodulation and symbiotic effectiveness than plants inoculated with the acid-sensitive strain. Laboratory prescreening of B. japonicum for acid, Al and Mn tolerance in acid media successfully identified strains which were symbiotically competent in low pH soils.     

Response of Rhizobium strains to acid and aluminium stress

Synchronous-culture, enrichment and isolation trials were done to determine effects of simulated soil acidity stress on growth of Rhizobium sp. (cowpea group), and to test whether tolerance of a strain is stable.In synchronous cultures, acidity and Al reduced the frequency of cell division. Non-dividing cells did not die, but those near division snowed evidence of heightened sensitivity to Al³⁺ and H⁺.Differences in tolerance within single-strain populations were evidently not genetically determined. Prolonged culture under stress (72 generations) failed to enrich a strain in putative tolerant variants. And from six strains of different tolerance, isolates from single colonies that had grown on stress medium were no more tolerant than isolates from colonies on non-stress medium. Tolerance is a consistent and stable strain property.     

Atlantic Salmon and Acidification in Southern Norway: A Disaster in the 20th Century, but a Hope for the Future?

Due to acidification, 18 Norwegian stocks of Atlantic salmon are extinct and an additional 8 are threatened. In the two southernmost counties, salmon is eradicated. Due to the high acid sensitivity, production of salmon was greatly reduced as early as 1920, several decades before acid rain was recognized as an environmental problem. International agreements on reduced atmospheric emissions will reduce acidification effects in Norway substantially during the coming 20 to 50 years. However, the extreme acid sensitivity of salmon makes the destiny of this species in Southern Norway uncertain. Liming is an effective measure to protect and restore fish populations in acidified waters. Liming of acidified salmon rivers has become important in Norway in recent years which in combination with reduced emissions will be an important contribution to protection of the Atlantic salmon species. In this paper we give an overview of the effects of acidification on Norwegian salmon and discuss different aspects of mitigation measures; the expected effect of international agreements on reduced atmospheric emissions, the expected effect of liming on salmon production and the possibilities of re-establishing self sustaining salmon stocks in limed rivers.     

Acid Toxicity Levels in Nova Scotian Rivers Have Not Declined in Synchrony with the Decline in Sulfate Levels

The Atlantic salmon (Salmo salar) resource of eastern Canada is impacted by acid rain in the Southern Upland (Atlantic Coast) area of Nova Scotia. Salmon runs in this area have become extinct in 14 rivers, are severely impacted in 20 rivers, and lightly impacted in 15 rivers. Water chemistry and fish communities in nine Southern Upland salmon rivers were studied from 1982 to 1996 as part of the effort to monitor the effects of the emission controlprograms in Canada and the United States. There hasbeen no statistically significant change in total ioncontent of Southern Upland river water, but there wasa significant decline in sulfate levels that was balanced by an increase in organic anions, and declines in calcium and magnesium that were balanced by increases in sodium and potassium. A geochemical scenario is proposed to account for these chemical changes. River water pH levels showed no overall linear trend, but at borderline toxicity sites the year-to-year variations in pH were correlated withchanges in juvenile salmon population densities. Tenfish species were collected, but none showed anysignificant overall time trend in population density.Fish species diversity was positively correlated with pH.     

Tolerance of durum wheat lines to an acid,aluminum‐toxic subsoil

Durum wheat, Triticum durum Desf., is reportedly more sensitive to aluminum (Al) toxicity in acid soils than hexaploid wheat, Triticum aestivum L. em. Thell. Aluminum‐tolerant genotypes would permit more widespread use of this species where it is desired, but not grown, because of acid soil constraints. Durum wheat germplasm has not been adequately screened for acid soil (Al) tolerance. Fifteen lines of durum wheat were grown for 28 days in greenhouse pots of acid, Al‐toxic Tatum subsoil at pH 4.5, and non‐toxic soil at pH 6.0. Aluminum‐tolerant Atlas 66 and sensitive Scout 66 hexaploid wheats were also included as standards. Based on relative shoot and root dry weight (wt. at pH 4.5/wt. at pH 6.0 X 100), durum entries differed significantly in tolerance to the acid soil. Relative shoot dry weight alone was an acceptable indicator of acid soil tolerance. Relative dry weights ranged from 55.1 to 15.5% for shoots and from 107 to 15.8% for roots. Durum lines PI 195726 (Ethiopia) and PI 193922 (Brazil) were significantly more tolerant than all other entries, even the Al‐tolerant, hexaploid Atlas 66 standard. Hence, these two lines have potential for direct use on acid soils or as breeding materials for use in developing greater Al tolerance in durum wheat. Unexpectedly, the range of acid soil tolerance available in durum wheat appears comparable to that in the hexaploid species. Hence, additional screening of durum wheat germplasm for acid soil (Al) tolerance appears warranted. Durum lines showing least tolerance to the acid soil included PI 322716 (Mexico), PI 264991 (Greece), PI 478306 (Washington State, USA), and PI 345040 (Yugoslavia). The Al‐sensitive Scout 66 standard was as sensitive as the most sensitive durum lines. Concentrations of Al and phosphorus were significantly higher in shoots of acid soil sensitive than in those of tolerant lines, and these values exceeded those reported to cause Al and phosphorus (P) toxicities in wheat and barley.     

Varietal diversity of upland rice in sensitivity to aluminium

A rapid and simple nutrient addition technique was used for evaluating Al tolerance of six local upland rice (Oryza sativa L.) cultivars (BG35, BR21, DA25, DA26, DA14, and DA22) from Bangladesh and three IRRI rice, IR46, IR97, and IR45, cultivars from the Philippines. The plants were grown for 21 days with Al (0 μM, 140 μM, 280 μM or 560 μM) at pH 4.1. The roots were more affected by Al than the shoots. In rating cultivars for Al sensitivity, relative shoot weight (RSW) was found to be the best parameter due to the severe damage of the roots, irrespective of Al sensitivity. The cultivars were rated as Al tolerant (BG35, BR21, DA25, and DA26), mid‐tolerant (DA14, DA22, and IR46) and sensitive (IR97 and IR45) . More Al was retained in the roots of tolerant cultivars than in the mid‐tolerant or sensitive cultivars. In shoots, the Al concentration of tolerant cultivars was less than in the mid‐tolerant or in the sensitive cultivars and the inhibition of growth was proportional to Al concentration irrespective of Al tolerance. Therefore, the variation among cultivars in Al sensitivity could be related to the capacity of roots to retain Al from transport to the shoots.     

Acid deposition and effects in Nordic Europe. Damage to fish populations in Scandinavia continue to apace

Although the decline in fish populations due to acidicwater in Norway started as early as in the 1920's the most rapid losses appeared during the 1960–70's. Until 1978, the populations of Atlantic salmon had disappeared from the southernmost part of Norway, and in these areas, more than half of the brown trout populations had been lost. Today, in spite of no increase in acid depositions, the fishery problems seems to continue at the same speed. Data based on interviews of the local fish authority shows that lakes still holding a fish population in the late 70's, have experienced a 30% loss of brown trout populations and a 12% loss of perch in the period 1978–1983. This trend have been confirmed by testfishing in lake systems having long data series. Salmon rivers on the western coast of Norway have experienced several episodes of fish kills due to rapid changes in water quality. These fish kills have mainly affected smolts of Atlantic salmon. Spawning migrating salmon on entering their acidified home river have also been affected. In Sweden, several salmon populations along the western coast have been lost due to acidification with no positive trends so far in the 1980's. Areas in central Sweden and in some high mountain areas are still experiencing a continuous and increasing acidification with detrimental effects on invertebrates and fish. In Finland, an increase in acidic deposition during the last decades have occurred, leading to acidification in the most sensitive freshwater systems. Although some acidified freshwater lakes are reported to have lost their fish stocks, few data on fish population effects are available.     

Differential Tolerances of Amaranthus Strains to High Levels of Aluminum and Manganese in Acid Soils

Species of Amaranthus are grown extensively as leafy green vegetables in tropical Africa and Asia and as high yielding grain crops in Western South America, Central America, Northern India, Western Nepal, and Pakistan. The crop is often grown on acid, marginal soils, under subsistence conditions, where liming even the soil plow layer may not be economically feasible. Hence, the identification or development of strains with high tolerance to acid soils would be beneficial. Aluminum and Mn toxicities are the most important growth‐limiting factors in many acid soils. The objective of our research was to determine the tolerances of selected Amaranthus strains to high levels of these elements in acid soils.

Fifteen strains, representing five species, were grown in greenhouse pots of an acid, Al‐toxic Tatum soil limed to pH 4.8 and 5.8. Strains differed significantly in tolerance to the acid soil. Relative yields (pH 4.8/pH 5.8%) ranged from 50.1 to 6.3% for tops and from 54.5 to 5.7% for roots. Four strains of A. tricolor L. (vegetable type) were significantly more tolerant than six strains of A. cruentus L. (seed and vegetable type). Strains of A. hypochondriacus L. and A. caudatus L. studied were intermediate in tolerance.

Twelve strains, representing four species, were grown on an acid, Mn‐toxic Zanesville soil at pH 4.6 and 6.3. Strains also differed significantly in tolerance to this acid soil; however, overall growth was better and strain differences were smaller than on Al‐toxic Tatum soil at pH 4.8. On Zanesville soil the relative top yields (pH 4.6/pH 6.3%) ranged from 74.1 to 18.6%. The most tolerant group included three strains of A. tricolor and one strain of A. hypochondriacus, but four strains of A. cruentus were also fairly tolerant. The sensitive end of the scale included one strain of A. cruentus and two strains of A. hypochondriacus.

In general, strains that were most tolerant to the Al‐toxic Tatum soil were also among the most tolerant to the Mn‐toxic Zanesville soil. Likewise, those most sensitive to the high Al soil were most sensitive to the high Mn soil. But some strains that were sensitive to excess Al in Tatum soil were fairly tolerant to high Mn in Zanesville soil.

Results suggest that superior strains of Amaranthus can be selected or developed for use on acid soils.     

Effects of lead on the soil bacterial microflora

The sensitivity of soil bacteria towards Pb was investigated. Soil suspensions from fourteen different soil types with a high or low Pb content were plated out on soil extract agar containing various concentrations of PbCl₂.In agar with a high Pb content, higher bacterial counts were found with suspensions from Pb-containing soils than with those of soils with a low Pb content. In the Pb-containing soils, proportionally more gram-negative rods were present while coryneform bacteria decreased. In an additional experiment, in which Pb was added to a sandy soil, more Pb-tolerant bacterial strains were found 3 years later than in the same soil without Pb.When pure cultures of the bacteria isolated from the soils were tested in liquid media for Pb tolerance, a higher proportion of tolerant strains was found in Pb-containing soils. Among strains of gram-negative bacteria isolated from these soils a higher proportion of tolerant strains was found than in corneform bacteria.It was concluded that as a consequence of Pb pollution of soil a selection of Pb-tolerant bacteria may take place.