Elasmobranch pericardial function. 1. Pericardial pressures are not always negative

Acute studies have led to the generalization that negative pericardial pressure is necessary for optimal cardiac function in elasmobranchs. We chronically instrumented horn sharks with pericardial catheters to test the hypothesis that ejection of pericardial fluid through the pericardioperitoneal canal (PPC) during routine handling could have accounted in part for previous measurements of exclusively negative pressures (–0.3 to –9.1 cm H₂O) in elasmobranchs. Maximum and minimum pericardial pressures measured immediately following routine handling (acute pressures) were more negative than those measured in resting horn sharks at intervals from 1 to 27 days following handling (chronic pressures). Chronic pericardial pulse pressure was less than acute. Entirely positive pericardial pressures were observed on occasion. Handling of chronically catheterized horn sharks resulted in ejection of 21 per cent (range=10–26, n=5) of the initial pericardial fluid volume through the PPC and reduced pericardial pressure. Operating pericardial fluid volume of horn sharks averaged 2.0 ml.kg^–1 (range=1.6–2.6, n=9). The PPC opened after 4.3±0.2 ml.kg^–1 (x±S.E.) of elasmobranch saline had been slowly infused into the pericardium, corresponding to an average pressure of 1.3±0.2 cm H₂O (n=10). The presence of the PPC plus a comparatively large pericardial fluid volume allows horn sharks to regulate pericardial pressure. Our analysis of pericardial pulse pressure, which can be an index of cardiac activity, suggests in contrast to previous studies that the elasmobranch heart can have relatively high stroke volumes at pericardial pressures near ambient. Thus, for venous return in resting or even moderately active elasmobranchs, it is more important that pericardial pressure be pulsatile than at a mean level which is negative.     

Elasmobranch pericardial function 2. The influence of pericardial pressure on cardiac stroke volume in horn sharks and blue sharks

The importance of negative pericardial pressure to heart function in clasmobranchs has been questioned due to the discovery of positive pericardial pressures in healthy horn sharks (Heterodontus francisci). We therefore implanted electromagnetic flow probes on the ventral aorta of horn sharks and blue sharks (Prionace glauca) and assessed stroke volume and pericardial pressure as pericardial fluid volume (PFV) was varied to test the hypothesis that elasmobranchs are capable of maintaining a relatively large stroke volume at pericardial pressures near and above ambient. Stroke volume was maximum between zero and 25% maximum PFV (maximum PFV: the volume of pericardial fluid required to open the pericardioperitoneal canal), where pericardial pressure was most negative. At 50% maximum PFV (pericardial pressure near or slightly above ambient) stroke volume was 70% and 98% of its maximum in horn sharks and blue sharks, respectively. At a larger PFV, stroke volume declined drastically, reaching zero where both PFV and pericardial pressure were maximum. Thus, at a pericardial pressure apparently associated with resting or mild activity, stroke volume is a relatively large proportion of the apparent maximum. Increased circulatory demands associated with strenuous activity may induce ejection of pericardial fluid through the perieardioperitoneal canal, which then lowers pericardial pressure. The lowered pericardial pressure causes an increase in stroke volume, indicating that control is in part effected by changing pericardial pressure.     

Regulation of cardiac function in the horn shark by changes in pericardial fluid volume mediated through the pericardioperitoneal canal

The California horn shark (Heterodontus francisci), instrumented with a pericardial catheter and a ventral-aortic flow probe, was studied to determine the effect of complete pericardial chamber evacuation on the time course for restitution of pericardial fluid-volume and pressure, and the effects of both fluid removal and its restitution on cardiac output. Prior to evacuation, pericardial pressure was –0.02±0.02 kPa, and cardiac output was 18±2 ml min^–1 kg^–1. Evacuation reduced pericardial pressure to –0.73±0.14 kPa, and increased cardiac output to 23±4 ml min^–1 kg^–1. The time course for restoration of post-evacuation pressure is described by a non-linear asymptotic function. A large percentage of the pericardial pressure and volume recovery occurred within the first hour, while, complete restoration of pre-withdrawal conditions required about 11 h. Pericardial pressure-volume relationships, determined by incremental infusion of small volumes of elasmobranch saline into the pericardium, confirm previous findings that the operating pericardial pressure in the horn shark is at or near ambient pressure and that both pericardial fluid volume and cardiac stroke volume influence horn shark pericardial pressure.