Ontogeny and evolution of salinity tolerance in anadromous salmonids: Hormones and heterochrony

Use of estuaries and oceans by salmonids varies greatly, from no use in nonanadromous species, to movement toward the sea soon after hatching and emergence in some Pacific salmon. This variation is accompanied by large differences in the ontogeny of salinity tolerance among salmonids. Some species acquire increased salinity tolerance early in development, whereas others develop this characteristic much later, indicating there is a heterochrony (change in timing) in the development of salinity tolerance in salmonids. The basic physiological mechanisms for ion regulation in seawater (such as increased gill chloride cells, gill Na⁺,K⁺-ATPase activity, membrane permeability, and drinking rate) are common to all salmonids. What determines the differences in salinity tolerance among the salmonids is not the basic mechanisms for salt secretion but the environmental and ontogenetic control of these mechanisms. In salmonids such as pink and chum salmon that enter seawater soon after emergence, acclimation to seawater may be controlled largely by internal (ontogenetic) information. In smolting salmonids that acquire increased salinity tolerance 1–2 yr after hatching, photoperiod is the dominant environmental cue. In nonsmolting species that migrate 2–3 yr after hatching, salinity itself may be the primary stimulus for salt secretory mechanisms. Physiological changes triggered by developmental and environmental cues are mediated by endocrine factors. Treatments with cortisol, growth hormone, and insulin-like growth factor I have been shown to increase seawater tolerance of salmonids, whereas prolactin is inhibitory. Differences in developmental patterns of endocrine activity (such as secretion, binding proteins, and receptors) are hypothesized to be responsible for the differences in timing (heterochrony) of increased salinity tolerance among and within salmonid species.