Wells M J
J Exp Biol. 1980 Apr;85:111-28. doi: 10.1242/jeb.85.1.111.
The circulatory system of cephalopods is based on a trio of hearts, with two pairs of associated ganglia linked to the CNS by a pair of visceral nerves. The beat of the hearts was recorded from free-moving octopuses before and after surgical removal or disconnexion of elements of the nervous system. Severing the visceral nerves does not stop the hearts, which continue to beat in a powerful well co-ordinated manner in isolation from the CNS. The nerves seem to be concerned in raising the cardiac output in exercise, and with stopping the hearts when mantle movements cease, but they are not necessary for the initiation of maintenance of the normal rhythm. Removal of the fusiform ganglia severs all nervous connexions between the ywo gill hearts, and deprives the systemic heart of its nerve supply. The trio of hearts continues to beat as strongly as before. Removal or disconnexion of a cardiac ganglion disrupts the beat of the corresponding gill heart which now tends to contract in an ill-coordinated and rather feeble manner, though at much the same frequency as before; with both cardiacs gone the systemic heart, which contracts only when it is filled, tends to drop in frequency and the mean aortic pressure falls. The system remains rhythmic, however, and the beat may recover, to the point where aortic pressures and frequencies approach those found in intact animals at rest; even octopuses with both fusiform and both cardiac ganglia removed can survive for many hours. From the performance of the isolated branchial heart, the existence of a pulsating vesicle in each cardiac ganglion, the effects of cardiac ganglion removal and the remarkable steadiness of heartbeat frequency shown by intact animals under a variety of conditions, it is argued that the heartbeat rhythm is normally controlled by pacemakers in the branchial heart/ cardiac ganglion complexes, and perhaps, in intact animals, from within the cardiac ganglia themselves. The picture of the control of the heartbeat that emerges from the study of free moving essentially intact animals is quite different from that arising from in vitro and acute preparation studies. It suggests that the conventional wisdom about the control of the heartbeat in cephalopods (and perhaps by implication, in other molluscs) may need to be considerably revised.
头足类动物的循环系统由三个心脏组成，有两对相关的神经节通过一对内脏神经与中枢神经系统相连。在手术切除或切断神经系统的某些部分之前和之后，记录了自由活动的章鱼的心跳情况。切断内脏神经并不会使心脏停止跳动，心脏在与中枢神经系统隔离的情况下仍会以有力且协调良好的方式继续跳动。这些神经似乎与在运动时提高心输出量有关，并且在套膜运动停止时使心脏停止跳动，但它们对于启动和维持正常节律并非必需。切除梭形神经节会切断两个鳃心之间的所有神经连接，并使体心脏失去神经供应。这三个心脏仍会像以前一样剧烈跳动。切除或切断一个心脏神经节会扰乱相应鳃心的跳动，此时鳃心往往会以不协调且相当微弱的方式收缩，尽管频率与以前大致相同；两个心脏神经节都被切除后，仅在充满血液时才收缩的体心脏，其频率往往会下降，平均主动脉压也会降低。然而，该系统仍保持节律性，心跳可能会恢复，直至主动脉压和频率接近完整动物在静息状态下的水平；甚至切除了梭形神经节和两个心脏神经节的章鱼也能存活数小时。从孤立鳃心的表现、每个心脏神经节中搏动囊泡的存在、心脏神经节切除的影响以及完整动物在各种条件下表现出的心跳频率的显著稳定性来看，有人认为心跳节律通常由鳃心/心脏神经节复合体中的起搏器控制，也许在完整动物中，是由心脏神经节自身内部控制。从对基本完整的自由活动动物的研究中得出的心跳控制情况与体外和急性制备研究中得出的情况大不相同。这表明关于头足类动物（也许由此类推，其他软体动物）心跳控制的传统观念可能需要大幅修正。
