Resting and maximal heart rates in ectothermic vertebrates.

Resting and maximal heart rates (HR) in ectothermic vertebrates are generally lower than those in endotherms and vary by more than an order of magnitude interspecifically. Variation of HR transcends phylogeny and is influenced by numerous factors including temperature, activity, gas exchange, intracardiac shunts, pH, posture, and reflexogenic regulation of blood pressure. The characteristic resting HR is rarely the intrinsic rate of the pacemaker, which is primarily modulated by cholinergic inhibition and adrenergic excitation in most species. Neuropeptides also appear to be involved in cardiac regulation, although their role is not well understood. The principal determinants of resting HR include temperature, metabolic rate and hemodynamic requirements. Maximal HRs generally do not exceed 120 b min-1, but notable exceptions include the heterothermic tuna and small reptiles having HRs in excess of 300 b min-1 at higher body temperatures. Temperature affects the intrinsic pacemaker rate as well as the relative influence of adrenergic and cholinergic modulation. It also influences the evolved capability to increase HR, with maximal cardiac responses matched to preferred body temperatures in some species. Additional factors either facilitate or limit the maximal level of HR, including: (1) characteristics of the pacemaker potential; (2) development of sarcoplasmic reticulum as a calcium store in excitation-contraction coupling; (3) low-resistance coupling of myocardial cells; (4) limitations of force development imposed by rate changes; (5) efficacy of sympathetic modulation; and (6) development of coronary circulation to enhance oxygen delivery to myocardium. In evolutionary terms, both hemodynamic and oxygen requirements appear to have been key selection pressures for rapid cardiac rates.