Virágh S, Challice C E
Anat Rec. 1981 Sep;201(1):157-68. doi: 10.1002/ar.1092010117.
The origin of the epicardium and the formation of the early blood vessels of the heart prior to the opening of the coronary arteries from the aorta have been studied in the 9-13.5 day post coitum (dpc) mouse embryo heart. The epicardium begins to appear by 9 dpc. The majority of the epicardial cells derive from the somatopleural investment of the septum transversum, from where they migrate, associated to form vesicles, to the dorsal aspect of the ventricles and atria. The epicardial cells then migrate over the lateral surfaces and the AV sulcus to the ventral aspect of the heart. In the subepicardial space around the sulci, the proliferating epithelial tissue is found, also in vesicular form, for a time. The ventrally migrating primordial epicardial tissue ensheaths lastly the truncus arteriosus, while the sinus venosus is coated with epicardium ab initio, where (and also in the SA sulcus) the epicardial cells derive in part from the cuboidal cells of the pleuroperitoneal canal and in part from the somatopleural cells. The early blood vessel formation follows in space and time the development of the epicardium. The first blood vessels appear by dpc by the invagination of the endocardium into the early sinus muscle, and at the same time in the ventricular chamber by the encasing of the endocardium, as the trabeculae become consolidated into the myocardial walls. By this process sinusoids are formed, some of which penetrate through the myocardium and which, by rapid proliferation, form an interconnected subepicardial plexus. These capillaries proliferate ventrally in the wide subepicardial space, reaching the septating truncus, in which the aorta and pulmonary artery are developing. The definitive coronary artery openings appear by 13 dpc, allowing the high pressure blood from the aorta to flow into a preexisting vascular bed.
在交配后9 - 13.5天（dpc）的小鼠胚胎心脏中，研究了心外膜的起源以及在冠状动脉从主动脉开口之前心脏早期血管的形成。心外膜在9 dpc时开始出现。大多数心外膜细胞源自横隔的体壁层，它们从那里迁移，聚集形成小泡，到达心室和心房的背侧。然后心外膜细胞迁移到心脏的侧面和房室沟，再到心脏的腹侧。在沟周围的心外膜下间隙中，一段时间内也发现了呈小泡状增殖的上皮组织。最后，向腹侧迁移的原始心外膜组织包裹动脉干，而静脉窦从一开始就被心外膜覆盖，在心外膜细胞部分源自胸腹管的立方体细胞，部分源自体壁层细胞的地方（以及窦房沟）。早期血管形成在空间和时间上与心外膜的发育同步。最早的血管在dpc时通过心内膜向内陷至早期窦肌中而出现，同时在心室腔中，随着小梁合并到心肌壁中，心内膜被包裹，从而形成血窦，其中一些血窦穿透心肌，并通过快速增殖形成相互连接的心外膜下丛。这些毛细血管在心外膜下的宽阔空间中向腹侧增殖，到达正在发育主动脉和肺动脉的分隔动脉干。明确的冠状动脉开口在13 dpc时出现，使来自主动脉的高压血液流入预先存在的血管床。
