Chapter 9: Osmoregulation and Acid-Base Balance
Stephen D. McCormick, Eric T. Schultz, and Colin J. Brauner
Maintaining relatively constant levels of internal cellular ions is critical to the normal function of all animals. For many organisms this is achieved primarily by regulating the ion and acid-base composition of the blood within narrow limits. This understanding of the importance of “le milieu interior,” first espoused by Claude Bernard in the mid-1800s and later described as “homeostasis” by Walter Cannon, is a cornerstone of modern physiology. “It was Bernard’s view that we achieve a free and independent life, physically and mentally, because of the constancy of the composition of our internal environment” (Smith 1961:1). Direct contact between the gills and water makes ion, water, and acid-base balance especially challenging and important to fish and, in turn, makes fish important subjects for understanding the evolution and control of all of these homeostatic processes.
Several strategies exist within fishes for regulating ion concentrations in the blood relative to external (environmental) salt concentrations. Hagfishes, which are one extant group representing the ancestral jawless condition of vertebrates, are restricted to seawater (SW) and have an osmoconforming strategy in which the internal (blood) and external osmotic concentrations are very similar (Currie and Edwards 2010), but important differences do exist (Sardella et al. 2009). Lampreys are the other group of extant jawless fishes and either live wholly in freshwater (FW) or are anadromous. Lampreys have an osmoregulatory strategy in which the internal concentrations of ions are approximately one-third that of SW (Reis-Santos et al. 2008). Their underlying mechanisms of ion transport and osmoregulation appear to be nearly identical to those of the more recently evolved ray-finned fishes (Figure 9.1), which have adopted a similar osmoregulatory strategy. Elasmobranchs and coelacanths in SW retain high levels of urea in their plasma and are osmoconformers (Figure 9.2), whereas in the relatively rare instances elasmobranchs are found in FW, they are hyperosmoregulators, maintaining plasma ion levels in excess of environmental levels via mechanisms similar to FW ray-finned fishes (Ballantyne and Robinson 2010).