Mechanisms of venous return and ventricular filling in elasmobranch fish

The current concept of ventricular filling in the elasmobranch fish (sharks and rays) is that a subambient pericardial pressure establishes a negative diastolic pressure gradient for the atrium and that ventricular end-diastolic volume is exclusively determined by atrial systole. In contrast, recent findings using echo-Doppler and digital imaging techniques have demonstrated two filling phases in the elasmobranch ventricle. In this study, simultaneous atrial and ventricular pressure measurements made on sharks with an open or intact pericardium establish that atrial pressure is above ventricular diastolic pressure until the onset of ventricular systole. A positive biphasic atrioventricular pressure gradient thus ensures ventricular filling during early diastole, as a result of ventricular relaxation, as well as during atrial systole. Although a reduction in pericardial pressure resulted in a decline in the atrial and ventricular pressure, a positive atrioventricular pressure gradient is conserved. The finding that atrial diastolic pressure is not lower than ventricular diastolic pressure, when combined with previous results showing that pericardial pressure is generally at or above ambient and that ventricular filling is biphasic, constitutes a strong body of evidence favoring the operation of a direct venous inflow as the mechanism by which the elasmobranch heart fills.