Control of cardiovascular function and its evolution in Crustacea.

Work in the last decade has shown that crustacean open circulatory systems are highly efficient and controlled in a complex manner. Control occurs at several levels. Myocardial contraction is initiated in the cardiac ganglion but constantly modulated by the central nervous system, both directly via the cardioregulatory nerves and indirectly via the neurohormonal system. Heart rate and stroke volume can be controlled independently and measurements of both are needed to assess cardiac output accurately. Haemolymph outflow from many arthropod hearts is via a complex multiarterial distribution system, and the regional distribution of cardiac output is tightly controlled via cardioarterial valves at the base of each artery. These valves contain innervated muscle, and differential contraction serves to regulate the efflux of oxygenated haemolymph into a particular system. The major influence on both the evolution and control of arthropod open blood vascular systems is efficiency of oxygen uptake and delivery. This influence is illustrated by reference to a variety of crustacean and other arthropod types.