2 The role of the kidney in the maintenance of water balance

This chapter shows how the mammalian kidney is able to regulate the excretion of water independently from that of solutes. For this function, which derives from several evolutionary steps among vertebrates, it takes advantage of the diluting ability of the thick ascending limb to produce osmotic energy which is then used to concentrate solutes in the urine. This concentration is permitted by a highly sophisticated architecture of nephrons and vessels in the renal medulla, combined with special permeability characteristics of the different nephron segments and specific hormonal regulation. Two different types of loops of Henle and several well-insulated vascular compartments contribute to this process. The major nitrogenous waste product, urea, is concentrated by an indirect process involving a transfer of osmotic energy from the outer to the inner medulla. As known for several decades, concentrating function is primarily regulated by the effect of antidiuretic hormone (ADH) on water permeability of the collecting duct. However, as discovered more recently, it is also largely dependent upon the effect of the same hormone on urea permeability in the terminal collecting duct. In addition, recent investigations have revealed a much more complex hormonal regulation of the concentrating process than previously thought. ADH itself acts on many other structures in the kidney, and many other hormones and mediators, the secretion of which is not thought to be influenced by the water status, do affect urine concentration either directly or by their interaction with ADH. Rodents display a wide spectrum of morphological and functional renal adaptations improving water conservation. Their study has brought a better understanding of the significant steps and anatomical structures that contribute to the concentrating process. Finally, it is also apparent that the capacity to concentrate urine is influenced in individual animals of a given species by the availability of water, by specific feeding patterns, and by the protein content of the diet.