Environmental hypoxia as a metabolic constraint on fish: the case of Atlantic cod, Gadus morhua

Hypoxia is known to provoke a wide range of effects on aquatic animals. Here we use laboratory and field data on Atlantic cod, Gadus morhua, to illustrate that many of these responses can be explained within the metabolic scope (MS) framework, i.e. taking into account the directive and limiting effects of dissolved oxygen (DO) on the ability of animals to acquire energy for growth and activity. A MS model for cod shows that scope for activity (swimming, feeding, etc.) is proportional to DO and becomes nil, jeopardising survival, when DO is < approximately 20% air saturation. Laboratory studies have confirmed this lethal threshold and demonstrated that growth and food ingestion were significantly reduced below 70% sat. This loss of appetite has been linked to a reduction of the peak value and an increase in duration of postprandial metabolism, in agreement with the MS model. Dwindling MS during hypoxia imposes an upper limit to swimming performance. Cod may also opt to reduce spontaneous swimming activity to spare oxygen for other activities such as digestion. In the Kattegat, the Baltic Sea, and the Gulf of St. Lawrence, eastern Canada, cod completely avoid waters where their MS is near zero. Furthermore, cod density increases exponentially with DO up to approximately 70% sat in the Gulf of St. Lawrence. Although hypoxia results in other direct and indirect effects as well, the MS framework allows modelling of many of the responses to hypoxia for individual cod that ought to be reflected at the population and community levels. The MS framework is also useful to compare species responses. We show that the impact of hypoxia on MS is similar, when expressed as a proportion of MS in normoxia, in cod, European sea bass (Dicentrarchus labrax), the common sole (Solea solea) and turbot (Psetta maxima). Data are required for other species to evaluate how general these findings are.