Freshwater acidification: effects on the early life stages of fish

Summary Over the last 20–30 years there has been a very extensive international research effort into freshwater acidification and the decline or loss of soft-water fisheries. The research includes many laboratory studies on physiology and survival of fish under the conditions caused by acidification. The causes of acidification have become increasingly well understood and accepted. In many countries, industrial and environmental management strategies are being implemented with the eventual aim of halting or reversing acidification in vulnerable areas. Attention has shifted away from research into the effects of acidification on fish, and the number of new papers on this subject is declining. Now appears a good time to review this research.As is often the case in research of this kind, it is difficult to compare work from different laboratories. Especially in earlier work, experimental conditions may not be fully relevant to the field (e.g. in the choice of ambient calcium concentration), experimental media may be inadequately specified (e.g. trace metal concentrations) and analytical techniques may be open to question (e.g. pH measurement). This may explain, in part, why agreement is lacking on many aspects of the physiological effects on the early life stages of fish, as well as the adult stages to some extent. If research is to continue in this field, then adoption of standard soft, acid water media would be of significant benefit. Nethertheless, some very general conclusions on fish early life stages can be reached.There is some evidence of impaired gametogenesis and spawning, but in this area there is still a need for systematic research. After fertilization, the chorionic membranes afford protection from the effects of low pH and especially the accompanying toxic trace metals, notably aluminium. Indeed, dissolved aluminium can ameliorate acid toxicity at this stage in development, possibly by helping to decrease membrane permeability and reduce H⁺ uptake. The main hazard seems to be impaired development (perhaps related to a decline in pH of the perivitelline fluid), often leading to a lack of ability to escape from the chorionic membranes on hatching.After hatching, vulnerability increases. Dissolved aluminium, and other trace metals associated with acidification, become more important. Some studies indicate continuing protection from low pH afforded by low concentrations of aluminium. The causes of death can still only be inferred: much attention has concentrated on mineral uptake and balance, and there is anecdotal evidence of respiratory impairment, although not as unequivocal as it is for postlarval fish. Comparison with the physiological effects on postlarval fish is not necessarily helpful. There seems to be at least one important difference. Unlike postlarvel fish, larvae do not find aluminium at around pH 5.5 (when respiratory effects are probably most important) the most hazardous combination; rather, the greatest danger to larvae seems to be when very low pH (<5.0) and trace metals (including aluminium) combine in some kind of joint toxic effect.There is a great increase in vulnerability once dependence on the yolk has come to an end. This period coincides with the movement from the incubation site in the substratum into the open water, where the relevant water chemistry is likely to fluctuate between greater extremes. However, it is earlier, during autumn rainstorms and spring snowmelt, that pH and concentrations of protective calcium ions usually reach their lowest and metals like aluminium their highest values. In waters where acidification is most likely to occur, the early life stages of many freshwater fish species will be close to their survival threshold and only a slight decline in water quality might result in the loss of a complete year class, which would affect recruitment and eventual population status.With gradual acceptance of the causes and effects of freshwater acidification, the research effort in relation to fish is declining or shifting towards attempts to re-establish fisheries in vulnerable areas. As outlined above, only very general conclusions can be reached on individual life stages, and the relative susceptibilities compared. However, in many respects the sequence of events in the loss of fisheries is not fully understood, and information is lacking on how events at significant stages of the life cycle affect subsequent survival, development and recruitment. Plenty of work is still required.