Printed in Great Britain THE FUNCTIONAL DESIGN OF THE INSECT EXCRETORY SYSTEM

Insects have a slowly operating excretory system in which the passive rate of movement of haemolymph solutes into a slowly secreted primary excretory fluid is restricted by a reduction in the area available for passive transfer. They may have come to possess such an energy-saving system as a result of their evolution as small animals in osmotically and ionically stressful environ-ments. Although the possession of a waxy cuticle is a major element in their ability to live in such environments, insects have a very high surface-area/ volume ratio and this is likely to have conferred a selective advantage on individuals able to withstand unusually variable extracellular conditions. Among their major adaptations evolved to allow them to tolerate such conditions are the lack of a blood-borne respiratory pigment to be affected and the development of a system whereby their most sensitive tissues are protected by the regulatory activities of special covering epithelia. Because of these features it follows that there has been less evolutionary pressure for rapid excretory control of the haemolymph composition. With an excretory system that only slowly filters the haemolymph, less energy expenditure is involved in the production of the primary excretory fluid and in reabsorption of useful substances from it. In addition, insects are able to maintain in circulation high concentrations of substances such as amino acids, trehalose, and lipids. They can also eliminate excess fluid at very high rates with the loss of only trace amounts of haemolymph solutes. It is argued that terrestrial insects owe much of their success to their ability to recover virtually all the water from the slow flow of primary excretory fluid. The hindgut cells that are responsible for this recovery are aided by their cuticular lining which protects them from contact with the very high concentrations of potentially interfering compounds in the excretory material