Vertebrate salt glands: short- and long-term regulation of function

Excess salt loads in most non-mammalian vertebrates are dealt with by a variety of extra-renal salt-secreting structures collectively described as salt glands. The best studied of these are the supra-orbital nasal salt glands of birds. Two distinct types of response to osmoregulatory disturbances are shown by this structure: a progressive adaptive response on initial exposure to a salt load that results in the induction and enhancement of the secretory performance or capabilities of the gland; and the rapid activation of existing osmoregulatory mechanisms in the adapted gland in response to immediate osmoregulatory imbalance. Not only is the time-frame of these two types of response very different, but the responses usually involve fundamentally different processes: e.g., the growth and differentiation of osmoregulatory structures and their components in the former case, compared with the rapid activation of ion channels, pumps etc. in the latter. Despite marked differences in the nature and time-frame of these responses, they both are apparently triggered by neuronally released acetylcholine, which acts at muscarinic receptors on the secretory cells to induce an inositol phosphate-dependent increase in cytosolic-free calcium concentrations ([Ca2+]i). Therefore, the question arises as to how the cells produce the appropriate distinct response using a single common signal (i.e., an increase in [Ca2+]i). Examination of the features of this signaling pathway in the two conditions described, reveals that they each are uniquely tuned to generate a response with the characteristics appropriate for the cells' requirements. This tuning of the signal involves often rather subtle changes in the overall signaling pathway that are part of the adaptive differentiation process.