Anatomy of the human atrioventricular junctions revisited

There have been suggestions made recently that our understanding of the atrioventricular junctions of the heart is less than adequate, with claims for several new findings concerning the arrangement of the ordinary working myocardium and the specialised pathways for atrioventricular conduction. In reality, these claims are grossly exaggerated. The structure and architecture of the pathways for conduction between the atrial and ventricular myocardium are exactly as described by Tawara nearly 100 years ago. The recent claims stem from a failure to assess histological findings in the light of criterions established by Monckeberg and Aschoff following a similar controversy in 1910. The atrioventricular junctions are the areas where the atrial myocardium inserts into, and is separated from, the base of the ventricular mass, apart from at the site of penetration of the specialised axis for atrioventricular conduction. There are two such junctions in the normal heart, surrounding the orifices of the mitral and tricuspid valves. The true septal area between the junctions is of very limited extent, being formed by the membranous septum. Posterior and inferior to this septal area, the atrial myocardium overlies the crest of the ventricular septum, with the atrial component being demarcated by the landmarks of the triangle of Koch. The adjacent structures, and in particular the so-called inferior pyramidal space, were accurately described by McAlpine (Heart and Coronary Arteries, 1975). Thus, again there is no need for revision of our understanding. The key to unravelling much of the alleged controversy is the recognition that, as indicated by Tawara, the atrioventricular node becomes the atrioventricular bundle at the point where the overall axis for conduction penetrates into the central fibrous body. There are also marked differences in arrangement, also described by Tawara, between the disposition of the conduction axis in man as compared to the dog.     

Anatomy of membranous ventricular septum in the human heart

One of the most common congenital abnormalities of the human heart is a defect in the development of the membranous part of the ventricular septum which, in this study, is designated the MVS. The WS is a thin fibrous membrane, about 1 cm long, which extends upward and to the right from the muscular ventricular septum to the adjacent part of the aortic fibrous annulus that also gives attachment to the right posterior (noncoronary) and anterior (right coronary) aortic valve cusps. It is of considerable clinical importance that there lies between the muscular ventricular septum and the MVS the atrioventricular (AV) bundle of the cardiac conduction system. The MVS has an irregular quadrangular form and has right and left surfaces. This study is based on macroscopic and histological sections of more than 30 normal and abnormal hearts. © 1994 Wiley-Liss, Inc.     

The ox atrioventricular conduction axis compared to human in relation to the original investigation of sunao tawara

It was Sunao Tawara who, in 1906, established the foundations for knowledge of the arrangement of the atrioventricular conduction axis in man and other mammals. Study of the hearts of ungulates was a central part in his investigation, which assessed other species, including man. He described several subtle differences between the mammals. We have now ourselves studied the cardiac conduction tissue of the ox heart, comparing our findings with our knowledge of the arrangement in man, and providing new insights into the differences illustrated by Tawara. It is, perhaps, surprising that these differences, although subtle, have not attracted more attention. We show that the major difference is the fact that the noncoronary aortic sinus in the ox heart is mainly supported by the myocardium of the ventricular septum, whereas in the human heart the sinus, and its leaflet, are in fibrous contiguity with the aortic leaflet of the mitral valve. It is this feature that determines the difference in the arrangement of the conduction axis between the species. We also show that the emergence of the left bundle branch on the left ventricular aspect of the muscular septum is more variable than previously described. Clin. Anat. 33:383–393, 2020. © 2019 Wiley Periodicals, Inc.     

Intramural Purkinje Cell Network of Sheep Ventricles as the Terminal Pathway of Conduction System

To identify the anatomical basis for cardiac electrical signal conduction, particularly seeking the intramural terminals of conduction pathway within the ventricles, sheep hearts were examined compared with human hearts utilizing the characteristic morphology of Purkinje cells as a histological marker. In 15 sheep and five human autopsies of noncardiac death, prevalence of Purkinje or Purkinje‐type cells were histologically examined in the atrioventricular node, its distal conduction pathway, the interventricular septum, and the right‐ and left‐ventricular free walls. Myocardial tissue cleavages were examined in the transmural sections (along cardiac base‐to‐apex axis) obtained from the septum and ventricular free walls. Serial histological sections through virtually the entirety of the septum in selected sheep were used as the basis of a three‐dimensional reconstruction of the conduction pathway, particularly of the intramural Purkinje cell network. Purkinje cells were found within the mural myocardium of sheep ventricles whereas no intramural Purkinje‐type cell was detected within the human ventricles. In the sheep septum, every intramural Purkinje cell composed a three‐dimensional network throughout the mural myocardium, which proximally connected to the subendocardial extension of the bundle branches and distally formed an occasional junction with ordinary working myocytes. The Purkinje‐cell network may participate in the ventricular excitation as the terminal conduction pathway. Individual connections among the Purkinje cells contain the links of through‐wall orientation which would benefit the signal conduction crossing the architectural barriers by cleavages in sheep hearts. The myocardial architectural changes found in diseased hearts could disrupt the network links including those with transmural orientation. Anat Rec, 2009. © 2008 Wiley‐Liss, Inc.     

人体心脏房室结、房室环及主动脉后结的巨微解剖

目的解剖分离人体心脏房室结和房室环的结构,阐述它们的形态特征及相互关系。方法通过体视显微镜解剖12例人体心脏的房室结、主动脉后结及房室环,再进行组织学观察,并绘图演示它们的结构关系。结果在二尖瓣环和三尖瓣环靠近冠状窦前缘处分别暴露了左、右房室环(12/12),直径分别为(0.69±0.12)mm、(0.78±0.13)mm。此处的左、右房室环穿行在房室隔内的心房肌与心室肌之间的间隙中,向房室结方向延伸。主动脉后结在主动脉根后方的房间隔中被探查到(7/9),它的后上方的房间隔间隙中有肌纤维与其相连,它的前下方分出左、右房室环,并且此处的左环比右环粗。在中心纤维体后方的心内膜下的深部,主动脉后结与房室结之间有直接的心肌组织连接通路,这条通路有别于另两条通路(左、右房室环)。结论主动脉后结和房室环可通过体视显微镜解剖暴露,主动脉后结与房室结之间有3条通路。     

Localisation and quantitation of autonomic innervation in the porcine heart I: conduction system

This study was prompted by the prospect of transgenic pigs providing donor hearts for transplantation in human recipients. Autonomic innervation is important for the control of cardiac dynamics, especially in the conduction system. Our objective was to assess the relative distribution of autonomic nerves in the pig heart, focusing initially on the conduction system but addressing also the myocardium, endocardium and epicardium (see Crick et al. 1999). Quantitative immunohistochemical and histochemical techniques were adopted. All regions of the conduction system possessed a significantly higher relative density of the total neural population immunoreactive for the general neuronal marker protein gene product 9.5 (PGP 9.5) than did the adjacent myocardium. A similar density of PGP 9.5-immunoreactive innervation was observed between the sinus node, the transitional region of the atrioventricular node, and the penetrating atrioventricular bundle. A differential pattern of PGP 9.5-immunoreactive innervation was present within the atrioventricular node and between the components of the ventricular conduction tissues, the latter being formed by an intricate network of Purkinje fibres. Numerous ganglion cell bodies were present in the peripheral regions of the sinus node, in the tissues of the atrioventricular groove, and even in the interstices of the compact atrioventricular node. Acetylcholinesterase (AChE)-containing nerves were the dominant subpopulation observed, representing 60–70% of the total pattern of innervation in the nodal tissues and penetrating atrioventricular bundle. Tyrosine hydroxylase (TH)-immunoreactive nerves were the next most abundant neural subpopulation, representing 37% of the total pattern of innervation in the compact atrioventricular node compared with 25% in the transitional nodal region. A minor population of ganglion cell bodies within the atrioventricular nodal region displayed TH immunoreactivity. The dominant peptidergic nerve supply possessed immunoreactivity for neuropeptide Y (NPY), which displayed a similar pattern of distribution to that of TH-immunoreactive nerve fibres. Calcitonin gene-related peptide (CGRP)-immunoreactive nerves represented 8–9% of the total innervation of the nodal tissues and penetrating atrioventricular bundle, increasing to 14–19% in the bundle branches. Somatostatin-immunoreactive nerve fibres were relatively sparse (4–13% of total innervation) and were most abundant in the nodes, especially the compact atrioventricular node. The total pattern of innervation of the porcine conduction system was relatively homogeneous. A substantial proportion of nerve fibres innervating the nodal tissues could be traced to intracardiac ganglia indicative of an extensive intrinsic supply. The innervation of the atrioventricular node and ventricular conduction tissues was similar to that observed in the bovine heart, but markedly different to that of the human heart. It is important that we are aware of these findings in view of the future use of transgenic pig hearts in human xenotransplantation.     

Anatomical and neurohistological observation of the heart of the jungle bush quail, Perdicula asiatica (Latham.).

Anatomy, histology and innervation of the heart of the jungle bush quail, Perdicula asiatica have been described. The cardiac conducting system is well developed except the atrioventricular node. The sinuatrial node is located at the cephalic end of the interatrial septum and comprised of a large number of specialised muscle fibres enclosing a few small nodal arteries. A few syncytial cells could also be observed. The atrioventricular node is small, rounded and compact mass present at the ventrocaudal end of the interatrial septum. The node is not enclosed by any connective tissue sheath. Atrioventricular bundle is quite conspicuous and a special left bundle branch descends from it and extending to the left ventricle. The presence of special left bundle branch probably helps in pumping the pure blood of left ventricle with a great force. The heart of the jungle bush quail is richly innervated. Large number of nerve fibres and ganglion cells are present at the sulcus terminalis and atrioventricular sulcus. Fine nerve fibres are also present in the mass of sinuatrial node, atrioventricular node, atrioventricular bundle and its branches. Nerve cells are found to be absent in the conducting system. A nervous connection exists between the sinuatrial node and atrioventricular node. Nerve fibres are also seen in the ventricular myocardium and at the sites of aortic arches.     

Histological examination of the potential arrhythmic substrates in the setting of Ebstein’s malformation

Cardiac arrhythmias, notably Wolff-Parkinson-White syndrome, are known to represent a major issue in patients with Ebstein’s malformation of the tricuspid valve. Abnormal conducting circuits, however, can also be produced by pathways extending either from the atrioventricular node or the ventricular components of the atrioventricular conduction axis, direct to the crest of the muscular ventricular septum. We hoped to provide further information on the potential presence of such pathways by investigations of six autopsied examples of Ebstein’s malformation. All were studied by histological sectioning on the full extent of the atrioventricular conduction axis, with limited sectioning of the right atrioventricular junction supporting the inferior and antero-superior leaflets of the deformed tricuspid valve. We used the criteria established by Aschoff (Verhandlungen der Deutschen Gesellschaft für Pathologie, 14, 1910, 3) and Mönckeberg (Verhandlungen der Deutschen Gesellschaft für Pathologie, 14, 1910, 64) over a century ago to define abnormal connections across the atrioventricular junctions, as these definitions retain their validity for the identification of gross myocardial connections across the insulating tissues of the atrioventricular junctions. In one specimen, we found two discrete accessory myocardial connections across the parietal right atrioventricular junction. In all of the hearts, we found so-called nodoventricular connections, and in one heart we also observed a well-formed connection originating from the penetrating atrioventricular bundle. In addition to accessory myocardial connections across the parietal right atrioventricular junction, therefore, our histological findings demonstrate a potential role for direct connections between the atrioventricular conduction axis and the ventricular myocardium in the setting of Ebstein’s malformation.     

The structure of the atrioventricular conducting system in the avian heart

The atrioventricular conduction system in three avian species has been studied by light and electron microscopy. A morphologically definable atrioventricular node was not found in any of these. The atrioventricular bundle is a well-defined structure, the proximal portion of which is in direct continuity with the atrioventricular ring, located in the arterial sheet of the muscular valve of the right atrioventricular opening. In the zone of transition between atrioventricular ring and bundle the compactness of the bundle is loosened, but the fibers do not establish continuity with the atrial fibers. The ring consists of Purkinje-like fibers, 10–15 μm in diameter, and (peripherally) small 3–5-μm-diameter junctional fibers which are in continuity with the common atrial fibers. In the muscular atrioventricular valve the fibers of the ring are insulated from the ventricular myocardium by a connective tissue sheet of the annulus fibrosus. It is suggested that in the avian heart the atrioventricular ring may fulfill a role similar to that of the atrioventricular node of mammals.     

超极化激活环核苷酸门控阳离子通道4、连接蛋白43和平足蛋白在小鼠胚胎心的表达与心传导系的发生

目的 探讨小鼠胚胎心传导系的发生机制。方法 用抗心肌肌球蛋白重链（MHC）、抗超极化激活环核苷酸门控阳离子通道4（HCN4）、抗缝隙连接蛋白43（CX43）和抗平足蛋白（podoplanin）抗体,对40只胚龄9~16d小鼠胚胎心进行连续石蜡切片并免疫组织化学或免疫荧光染色。结果 胚龄9d,较强的HCN4阳性表达集中在MHC阴性的静脉窦壁,随心脏发育,HCN4较强阳性表达逐渐向窦房结转移。胚龄11d开始,CX43阴性表达显示部位特异性。CX43阴性染色经窦房结沿右心房背侧壁和左、右静脉瓣向房室管背侧壁延伸。胚龄13d,左、右静脉瓣与房间隔底部融合后,进一步延续为房室管背侧壁发育中的CX43阴性染色的房室结,继而与室间隔顶部CX43阴性的房室束相连。胚龄9~10d,在MHC阳性心肌、心包腔背侧壁脏壁中胚层心肌前体细胞及静脉窦周间充质均显示podoplanin阳性表达。胚龄11~13d,podoplanin阳性间充质细胞沿心脏外表面扩展形成podoplanin阳性间皮样心外膜。结论 心脏发育早期,主起搏点位于静脉窦壁,起搏电位的产生早于收缩功能的发生。CX43阴性心肌是发育中的心传导系心肌,在胚龄11d即可观察到心传导系早期雏形。podoplanin参与促进心肌前体细胞向心肌细胞的分化。     

Spatial distribution of "tissue-specific" antigens in the developing human heart and skeletal muscle. III. An immunohistochemical analysis of the distribution of the neural tissue antigen G1N2 in the embryonic heart; implications for the development of the atrioventricular conduction system.

A monoclonal antibody raised against an extract from the Ganglion Nodosum of the chick and designated G1N2 proves to bind specifically to a subpopulation of cardiomyocytes in the embryonic human heart. In the youngest stage examined (Carnegie stage 14, i.e., 4 1/2 weeks of development) these G1N2-expressing cells are localized in the myocardium that surrounds the foramen between the embryonic left and right ventricle. In the lesser curvature of the cardiac loop this "primary" ring occupies the lower part of the wall of the atrioventricular canal. During subsequent development, G1N2-expressing cells continue to identify the entrance to the right ventricle, but the shape of the ring changes as a result of the tissue remodelling that underlies cardiac septation. During the initial phases of this process the staining remains recognizable as a continuous band of cells in the myocardium that surrounds the developing right portion of the atrioventricular canal, subendocardially in the developing interventricular septum and around the junction of the embryonic left ventricle with the subaortic portion of the outflow tract. During the later stages of cardiac septation, the latter part of the ring discontinues to express G1N2, while upon the completion of septation, no G1N2-expressing cardiomyocytes can be detected anymore. The topographic distribution pattern of G1N suggests that the definitive ventricular conduction system derives from a ring of cells that initially surrounds the "primary" interventricular foramen. The results indicate that the atrioventricular bundle and bundle branches develop from G1N2-expressing myocytes in the interventricular septum, while the "compact" atrioventricular node develops at the junction of the band of G1N2-positive cells in the right atrioventricular junction (the right atrioventricular ring bundle) and the ("penetrating") atrioventricular bundle. A "dead-end tract" represents remnants of conductive tissue in the anterior part of the top of the interventricular septum. The location of the various components of the avian conduction system is topographically homologous with that of the G1N2-ring in the human embryonic heart, indicating a phylogenetically conserved origin of the conduction system in vertebrates.     

Relationship between the membranous septum and the virtual basal ring of the aortic root in candidates for transcatheter implantation of the aortic valve

Knowledge of the anatomy of the membranous septum, as a surrogate to the location of the atrioventricular conduction axis, is a prerequisite for those undertaking transcatheter implantation of the aortic valve (TAVI). Equally important is its relationship of the virtual basal ring. This feature, however, has yet to be adequately described in the living heart. We analyzed computed tomographic angiographic datasets from 107 candidates (84.1 ± 5.2 years, 68% women) for TAVI. Using multiplanar reconstructions, we measured the height and width of the membranous septum, and the distances of its superior and inferior margins from the virtual basal ring plane. We also assessed the extent of wedging of the aortic root between the mitral valve and the ventricular septum. Mean heights and widths of the membranous septum were 6.6 ± 2.0, and 10.2 ± 3.1 mm, respectively, with its size significantly associated with that of the aortic root (P < 0.05). Its superior and inferior margins were 4.5 ± 2.3 and 2.1 ± 2.1 mm, respectively, from the plane of the basal ring. The inferior distance, the surrogate for the adjacency of the atrioventricular conduction axis, was ≤ 5mm in 91% of the patients. Deeper wedging of the aortic root was independently correlated with a shorter inferior distance (β = 0.0569, P = 0.0258). The membranous septum is appreciably closer to the virtual basal ring than previously appreciated. These findings impact on estimations of the risk of damage to the atrioventricular conduction axis during TAVI. Clin. Anat. 31:525–534, 2018. © 2018 Wiley Periodicals, Inc.     

The development of the atrioventricular node and bundle in the ferret heart

The development of the atrioventricular (AV) node and bundle in the ferret heart was examined at the light microscopic level. The AV node develops from two primordia which were first observed in the posterior wall of the common atrium during the stage when the single heart tube convolutes. During septation of the heart, the AV nodal primordia eventually fuse and come to lie at the base of the interatrial septum. The right AV nodal primordium is located below the attachment of the right venous valve to the interatrial septum. The left AV nodal primordium maintains a position anterior to the prospective ostium of the coronary sinus. At 16 days of gestation, large pale cells were seen in the dorsal AV canal. By 21 days of gestation these AV canal cells have been replaced by AV bundle cells. At this time the bundle is continuous with both nodal primordia. At birth the AV bundle is continuous mainly with the component of the AV node that is derived from the right AV node primordium. The anulus fibrosus begins to undergo the greatest developmental change after the AV node and bundle attain their final position in the AV junction. However, the anulus does not completely separate the atria from the ventricles during the later stages of development nor at birth, so that accessory AV pathways are present in the newborn ferret heart. Both the AV node and the AV bundle also demonstrated continuity with the myocardial cells of the interventricular septum in the neonatal heart. During development there was no evidence that rings of specialized tissues at the junctions of the cardiac chambers give rise to any component of the cardiac conduction system.     

核纤层蛋白A、转录因子TBX3、缝隙连接蛋白43表达与小鼠胚胎心发育

目的 探讨小鼠胚胎心脏工作心肌和传导系心肌在形态发生和分化过程中核纤层蛋白A（lamin A）、转录因子TBX3、缝隙连接蛋白43（Cx43）的表达特点。
方法 用抗α-平滑肌肌动蛋白（α-SMA）、抗心肌肌球蛋白重链（MHC）、抗α-横纹肌肌动蛋白（α-SCA）、抗胰岛因子1（ISL-1）、抗Cx43、抗lamin A和抗转录因子TBX3,对46只胚龄8～15d小鼠胚胎心脏连续石蜡切片进行免疫组织化学及免疫荧光染色。 结果 胚龄9d,TBX3在原始心管的表达集中在房室管壁。10d始,TBX3阳性的表达逐渐从房室管壁沿着静脉瓣延续至窦房结、右心房背侧壁和房间隔。胚龄12～13d,TBX3阳性表达结构构成了中枢传导系雏形,包括窦房结、左右静脉瓣、房间隔、房室管、房室结和房室束。Cx43首先在胚龄9d的左心室腹侧壁和部分小梁心肌出现弱阳性表达,随着发育,Cx43逐渐在TBX3阴性的心房、心室工作心肌表达。Lamin A首先出现在10d房室管心内膜垫间充质细胞和左心室部分小梁心肌,随后在右心室小梁心肌出现,至胚龄15d,心室和心房小梁心肌及房室瓣均可见lamin A阳性表达,但致密心肌和中枢传导系心肌持续呈阴性表达。 结论 中枢传导系统雏形在小鼠胚龄13d形成,呈TBX3阳性,Cx43阴性的互补性表达。致密心肌和中枢传导系心肌在15d仍为lamin A表达阴性,说明此部分心肌分化成熟较晚。     

人胚心静脉窦和传导系统的早期发育

目的 探讨早期人胚心静脉窦及传导系的发生发育机制. 方法 用抗α-平滑肌肌动蛋白(α-SMA)、抗α-横纹肌肌动蛋白(α-SCA)和抗结蛋白(DES)抗体对29例C10～C16期人胚心连续切片行免疫组织化学染色. 结果 人胚发育C12～C13期,系统静脉汇集形成的静脉窦出现于心包腔尾端原始横膈间充质中,静脉窦壁间充质细胞逐渐分化为α-SCA阳性的静脉窦心肌细胞.C14期,心包腔的扩张使静脉窦进入心包腔内,参与了右心房的形成.DES阳性传导系心肌的分化始于C10期心房室管右侧壁,随发育逐渐向室间沟心肌扩展,发育为房室传导系的希氏束、左右束支及心室腔面的小梁心肌.在心房,DES表达首先出现于C11期心房背侧壁,在C13期,可见静脉窦左背侧壁α-SCA、α-SMA、DES阳性心肌带与左心房底部、房室管背侧壁相延续,这条心肌带可能参与了人胚心静脉窦至房室管传导系的发育.C14～C16期,DES强阳性染色从窦房结经左、右静脉瓣及心房的背、腹侧壁延伸至房室管右侧壁,可能是原始的心房传导通路. 结论 心包腔尾端原始横膈间充质是人胚静脉窦心肌发生区,原始横膈间充质细胞逐渐分化为心肌细胞,添加到人胚心管静脉端,形成心静脉窦心肌.人胚心传导系心肌的分化始于房室管,随心管发育逐渐向动、静脉端扩展,在C16期,已分化为形态清晰可辨的DES阳性胚胎心传导系.     

Cardiac expression of polysialylated NCAM in the chicken embryo: correlation with the ventricular conduction system.

The neural cell adhesion molecule (NCAM) and its polysialic acid moeity (PSA) affect cellular interactions during the development of the nervous system and skeletal muscle. NCAM has also been identified in the embryonic heart of various species including humans. However, knowledge regarding the role of NCAM and its function-modulating PSA in cardiogenesis is limited. The distribution of NCAM and its PSA in the ventricular myocardium of chicken embryos was determined by indirect immunofluorescence staining. The NCAM polypeptide was found throughout the cardiac myocardium. In contrast PSA was located in discrete regions in stage 20 to 44 embryos (during and after septation). Myocardium at the subendocardial regions of the atrioventricular canal and ventricular trabeculae were PSA positive by stage 20. At later stages, transverse sections of the postseptation heart just below the level of the atrioventricular interface revealed a PSA-positive bundle of myocardium in the septum. This bundle was continuous with two branches at a more apical level which in turn were continuous with the PSA-positive subendocardial myocardium lining the left and right ventricles. This pattern of PSA in the myocardium was similar to that of the ventricular conduction system configuration defined in the adult heart. Electron micrographs of the subendocardium of the ventricular septum revealed PSA positivity on myofibril-containing cells with the ultrastructural location of Purkinje fibers. At later stages (35-44) a subset of cells within PSA-positive regions was stained by an antibody against an isoform of the myosin heavy chain found in adult Purkinje fibers. These cells and surrounding tissue lacked PSA in the adult heart. Thus polysialylated NCAM may be modulating cell-cell interactions during the development of the ventricular conduction system.     

Role of the left interventricular sulcus in formation of the interventricular septum and crista supraventricularis in normal human cardiogenesis

Serial sections of normal human embryos were studied and three-dimensional images reconstructed to determine the early development of the interventricular septum. The position of the interventricular septum is determined in stage 9 of normal development by the formation of the left interventricular sulcus. As a result of unknown properties of the cells of the myocardial layer, the left interventricular sulcus persists while the right disappears, producing the initial lateral asymmetry of the primary heart tube. By stage 14, the left interventricular sulcus forms a spiral which is continuous with the developing interventricular septum. The dorsal limb of the spiral passes to the right between the atrioventricular canal and the origin of the outflow tract, and is lost in the wall of the trabeculated right ventricle. It appears that this dorsal limb of the spiral is the precursor of part of the cirsta supraventricularis. The midportion of the sulcus, the bulboventricular groove, becomes the socalled fibrous continuity between the aortic and mitral valves. The ventral limb of the spiral passes caudally in the anterior interventricular groove and then dorsally and cranially toward the dorsal cushion of the atrioventricular canal. The ventral limb of the spiral is continuous with the crest of the muscular interventricular septum, which develops by apposition of tissue from the expanding right and left ventricles. From stage 14 to stage 19, the muscular interventricular septum, the atrioventricular endocardial cushions, and the ventricular end of the spiral ridges of the outflow tract appose and fuse. Subsequent formation of the membranous interventricular septum completes the physical separation of the right and left ventricles.     

Role of the left interventricular sulcus in formation of interventricular septum and crista supraventricularis in normal human cardiogenesis.

Serial sections of normal human embryos were studied and three-dimensional images reconstructed to determine the early development of the interventricular septum. The position of the interventricular septum is determined in stage 9 of normal development by the formation of the left interventricular sulcus. As a result of unknown properties of the cells of the myocardial layer, the left interventricular sulcus persists while the right disappears, producing the initial lateral asymmetry of the primary heart tube. By stage 14, the left interventricular sulcus forms a spiral which is continuous with the developing interventricular septum. The dorsal limb of the spiral passes to the right between the atrioventricular canal and the origin of the outflow tract, and is lost in the wall of the trabeculated right ventricle. It appears that this dorsal limb of the spiral is the precursor of part of the cirsta supraventricularis. The midportion of the sulcus, the bulboventricular groove, becomes the so-called fibrous continuity between the aortic and mitral valves. The ventral limb of the spiral passes caudally in the anterior interventricular groove and then dorsally and cranially toward the dorsal cushion of the atrioventricular canal. The ventral limb of the spiral is continuous with the crest of the muscular interventricular septum, which develops by apposition of tissue from the expanding right and left ventricles. From stage 14 to stage 19, the muscular interventricular septum, the atrioventricular endocardial cushions, and the ventricular end of the spiral ridges of the outflow tract appose and fuse. Subsequent formation of the membranous interventricular septum completes the physical separation of the right and left ventricles.     

The AV junction region of the heart: A comprehensive study correlating gross anatomy and direct three‐dimensional analysis. Part I. Architecture and topography

There is little detailed knowledge of the architecture of the AV junction region, the cytoarchitecture of the AV node or of its atrial connections. In the present study, the gross anatomy and topography of intracardiac structures in 21 adult canine hearts were photographically compared in whole and dissected hearts and tissue blocks and serial histologic sections made in three orthogonal planes. There are seven major new findings: 1) A coronary sinus fossa exists at the crux of the heart. It separates the right medial atrial wall (MAW) superoposterior region from the left atrium, its floor is the coronary sinus, and it carries the medial atrionodal bundle and proximal AV bundle on its right wall. 2) The posterior MAW forms two isolated bridges of myocardium as it surrounds the coronary sinus ostium, is isolated from the sinus venarum with crista terminalis and interatrial septum—by the floor of the inferior vena cava, and the narrow bridges link the posterior atrial wall to the mid MAW. 3) The tendon of Todaro has both epicardial and endocardial exposures, terminates in the superoposterior MAW and its medial aspect is adjacent sequentially to the medial atrionodal bundle and proximal AV bundle. 4) Only ordinary myocardium contacts the anulus fibrosus. 5) The ventricular septum's shoulder is humped shape posteriorly, is completely overlaid by anular myocardium and the medial leaflet and is joined by struts of papillary muscle. 6) The membranous septum joins the anterior ventricular septum to the crista supraventricularis, forms part of the posterior noncoronary and right aortic valve sinus walls and encases the right bundle branch. 7) The specialized conduction tissues, the superior, medial and lateral atrionodal bundles, the proximal AV bundle, AV node, distal AV bundle and right bundle branch are subjacent to MAW epicardium outside the right atrium, share regular intracardiac relationships with topographic landmarks and the medial atrionodal bundle, terminal superior atrionodal bundle, the proximal AV bundle and AV node are aligned to the medial leg of Koch's triangle. Thus, atrial myocardium of the AV junction region is that of the MAW. The floor of the inferior vena cava forms a natural barrier to impulse transmission along the full extent of the posterior MAW. The specialized tissues are outside of the MAW. Anatomic landmarks form reliable topographic landmarks for the specialized AV junction region tissues. A knowledge of the association of the specialized conduction tissues with specific regions of the MAW is useful in localizing the tissues and along with the coronary sinus fossa provides several extracardiac approaches. Anat Rec 256:49–63, 1999. © 1999 Wiley‐Liss, Inc.