The retention and physiological role of urea in the elasmobranchii

TLDR: The high urea content (2·0 and 2·5 per cent.) that characterises the blood, body fluids and tissues of the Elasmobranchii owes its origin to the relative impermeability of the gills and integument to this substance.

Abstract: Summary. The high urea content (2·0 and 2·5 per cent.) that characterises the blood, body fluids and tissues of the Elasmobranchii owes its origin to the relative impermeability of the gills and integument to this substance, and to the circumstance that the urea is actively conserved by the elasmobranch kidney. In consequence of this physiological uraemia, the elasmobranch is osmotically superior to its environment, even in sea water, and is able to absorb at least a minimum quantity of water for the formation of a urine that is isotonic or hypotonic to the blood, in accordance with the osmotic limitations of the fish kidney. We may suppose that the uraemic state tends to develop and to be regulated more or less automatically; urea is constantly being formed by the ordinary metabolic combustion of protein; water shortage leads to oliguria and urea retention, and the accumulated urea in the blood raises the osmotic pressure of the latter to a point where water is again available by direct absorption. Water plethora (as in fresh water) leads to diuresis and increased urea excretion, which in turn lowers the osmotic pressure of the blood and in some measure, at least, reduces the rate of water absorption. Trimethylamine oxide, which imparts about one-quarter as much osmotic pressure to the blood as does urea, is also conserved by the elasmobranch; the fact that this substance is present in the urine in lower concentration than in the blood suggests that, like urea, it is actively reabsorbed from the glomerular filtrate. This physiological uraemia is apparently an archaic biochemical habit acquired early in elasmobranch evolution, since it is shared by the divergent orders of the subclass. Presumably it is a secondary mode of osmotic regulation superimposed upon the more primitive one of branchial regulation, as observed in the teleostomes. The cleidoic egg, unique (among the fishes) in the Elasmobranchii, and the viviparous mode of reproduction, are viewed as adaptations to urea retention, protecting the embryo against the loss of urea during its early development. Urea retention enables the Elasmobranchii to maintain a considerably greater rate of urine formation (water excretion) than is observed in the marine teleosts, a fact that perhaps explains why the former do not show the glomerular degeneration or the aglomerular development observed in the latter. Whereas urine formation in the marine teleosts appears to be carried on normally at a reduced level considerably below the maximum possible rate, the elasmobranchs appear to maintain a maximal (though small) degree of glomerular activity at all times. Unlike the teleosts, they appear to possess no mechanisms for reducing glomerular activity; it may be that because of their superior osmotic position, due in turn to their physiological uraemia, they have never been faced with the necessity for conserving water to an excessive degree in the kidneys, and have therefore never evolved the means for doing so.