Revisiting the anatomy of the left ventricle in the light of knowledge of its development.

Despite centuries of investigation, certain aspects of left ventricular anatomy remain either controversial or uncertain. We make no claims to have resolved these issues, but our review, based on our current knowledge of development, hopefully identifies the issues requiring further investigation. When first formed, the left ventricle had only inlet and apical components. With the expansion of the atrioventricular canal, the developing ventricle cedes part of its inlet to the right ventricle whilst retaining the larger parts of the cushions dividing the atrioventricular canal. Further remodelling of the interventricular communication provides the ventricle with its outlet, with the aortic root being transferred to the left ventricle along with the newly formed myocardium supporting its leaflets. The definitive ventricle possesses inlet, apical and outlet parts. The inlet component is guarded by the mitral valve, with its leaflets, in the normal heart, supported by papillary muscles located infero-septally and supero-laterally. There is but a solitary zone of apposition between the leaflets, which we suggest are best described as being aortic and mural. The trabeculated component extends beyond the inlet to the apex and is confluent with the outlet part, which supports the aortic root. The leaflets of the aortic valve are supported in semilunar fashion within the root, with the ventricular cavity extending to the sinutubular junction. The myocardial-arterial junction, however, stops well short of the sinutubular junction, with myocardium found only at the bases of the sinuses, giving rise to the coronary arteries. We argue that the relationships between the various components should now be described using attitudinally appropriate terms rather than describing them as if the heart is removed from the body and positioned on its apex.

The clinical anatomy of the atrioventricular conduction axis.

It is axiomatic that the chances of achieving accurate capture of the conduction axis and its fascicles will be optimized by equally accurate knowledge of the relationship of the components to the recognizable cardiac landmarks, and we find it surprising that acknowledged experts should continue to use drawings that fall short in terms of anatomical accuracy. The accuracy achieved by Sunao Tawara (1906) in showing the location of the atrioventricular conduction axis is little short of astounding. Our purpose in bringing this to current attention is to question the need of the experts to have produced such inaccurate representations, since the findings of Tawara have been extensively endorsed in very recent years. The recent studies do no more than point to the amazing accuracy of the initial account of Tawara. At the same time, we draw attention to the findings described in the middle of the 20th century by Ivan Mahaim (1947). These observations have tended to be ignored in recent accounts. They are, perhaps, of equal significance to those seeking specifically to pace the left fascicles of the branching atrioventricular bundle.

The Atrioventricular Conduction Axis Revisited for the 21st Century.

Although first described in the final decade of the 19th century, the axis responsible for atrioventricular conduction has long been the source of multiple controversies. Some of these continue to reverberate. When first described by His, for example, many doubted the existence of the bundle we now name in his honour, while Kent suggested that multiple pathways crossed the atrioventricular junctions in the normal heart. It was Tawara who clarified the situation, although many of his key definitions have not universally been accepted. In key studies in the third decade of the 20th century, Mahaim then suggested the presence of ubiquitous connections that provided "paraspecific" pathways for atrioventricular conduction. In this review, we show the validity of these original investigations, based on our own experience with a large number of datasets from human hearts prepared by serial histological sectioning. Using our own reconstructions, we show how the atrioventricular conduction axis can be placed back within the heart. We emphasise that newly emerging techniques will be key in providing the resolution to map cellular detail to the gross evidence provided by the serial sections.

Relationship between the aortic root and the atrioventricular conduction axis.

Damage to the atrioventricular conduction axis continues to be a problem subsequent to transcatheter implantation of aortic valvar prostheses. Accurate knowledge of the precise relationships of the conduction axis relative to the aortic root could greatly reduce the risk of such problems. Current diagrams highlighting these relationships rightly focus on the membranous septum. The current depictions, however, overlook a potentially important relationship between the superior fascicle of the left bundle branch and the nadir of the semilunar hinge of the right coronary leaflet of the aortic valve. Recent histological investigations demonstrate, in many instances, a very close relationship between the left bundle branch and the right coronary aortic leaflet. The findings also highlight two additional variable features, which can be revealed by clinical imaging. The first of these is the extent of an inferoseptal recess of the left ventricular outflow tract. The second is the extent of rotation of the aortic root within the base of the left ventricle. Much more of the conduction axis is within the confines of the circumference of the outflow tract when the root is rotated in counterclockwise fashion as assessed from the perspective of the imager, with this finding itself associated with a much narrower inferoseptal recess. A clear understanding of the marked variability within the aortic root is key to avoiding future problems with atrioventricular conduction.

Evidence for concealed fasciculo-ventricular connections as revealed by His bundle pacing.

BACKGROUND: It is almost 100 years ago since Mahaim described the so-called paraspecific connections between the ventricular conduction axis and the crest of the muscular ventricular septum, believing such pathways to be ubiquitous. These pathways, however, have yet to be considered as potential pathways for septal activation during His bundle pacing. MATERIALS: So as to explore the hypothesis that specialised septal pathways might provide the substrate for septal activation during His bundle pacing, we compared the findings from 22 serially sectioned histological datasets and 34 different individuals undergoing His bundle pacing. RESULTS: We found histologically specialised pathways connecting the branching component of the atrioventricular conduction axis with the crest of the muscular ventricular septum in almost four-fifths of the histological datasets. In 32 of 34 patients undergoing His bundle pacing, the QRS complex closely resembled published images of known conduction through fasciculo-ventricular pathways. In only two patients was a delta wave not seen at any pacing voltages. Capture of these connections varied according to pacing voltage, a finding which correlated with the distance of the pathways from the site of penetration of the ventricular conduction axis. Ventricular activation times remained normal in the presence of the delta wave at higher pacing voltage but were prolonged at lower voltages. CONCLUSIONS: Our histologic findings confirm fasciculo-ventricular connections, initially described by Mahaim as being paraspecific, are likely ubiquitous. Analysis of 12-lead electrocardiograms leads us to conclude that fasciculo-ventricular pathways, concealed during sinus rhythm, become manifest with His bundle pacing.

Vulnerability of the ventricular conduction axis during transcatheter aortic valvar implantation: A translational pathologic study.

The ventricular components of the conduction axis remain vulnerable following transcatheter aortic valvar replacement. We aimed to describe features which may be used accurately by interventionalists to predict the precise location of the conduction axis, hoping better to avoid conduction disturbances. We scanned eight normal adult heart specimens by 3T magnetic resonance, using the images to simulate histological sections in order accurately to place the conduction axis back within the heart. We then used histology, tested in two pediatric hearts, to prepare sections, validated by the magnetic resonance images, to reveal the key relationships between the conduction axis and the aortic root. The axis was shown to have a close relationship to the nadir of the right coronary leaflet, in particular when the aortic root was rotated in counterclockwise fashion. The axis was more vulnerable in the setting of a narrow inferoseptal recess, when the inferior margin of the membranous septum was above the plane of the virtual basal ring, and when minimal myocardium was supporting the right coronary sinus. The features identified in our study are in keeping with the original description provided by Tawara, but at variance with more recent accounts. They suggest that the vulnerability of the axis during transcatheter valvar replacement can potentially be inferred on the basis of knowledge of the position of the aortic root within the ventricular base. If validated by clinical studies, our findings may better permit avoidance of new-onset left bundle branch block following transcatheter aortic valvar replacement.

Equal force generation potential of trabecular and compact wall ventricular cardiomyocytes.

Trabecular myocardium makes up most of the ventricular wall of the human embryo. A process of compaction in the fetal period presumably changes ventricular wall morphology by converting ostensibly weaker trabecular myocardium into stronger compact myocardium. Using developmental series of embryonic and fetal humans, mice and chickens, we show ventricular morphogenesis is driven by differential rates of growth of trabecular and compact layers rather than a process of compaction. In mouse, fetal cardiomyocytes are relatively weak but adult cardiomyocytes from the trabecular and compact layer show an equally large force generating capacity. In fetal and adult humans, trabecular and compact myocardium are not different in abundance of immunohistochemically detected vascular, mitochondrial and sarcomeric proteins. Similar findings are made in human excessive trabeculation, a congenital malformation. In conclusion, trabecular and compact myocardium is equally equipped for force production and their proportions are determined by differential growth rates rather than by compaction.

Mahaim Revisited.

The name Ivan Mahaim is well-known to electrophysiologists. However, alternative anatomical substrates can produce the abnormal rhythms initially interpreted on the basis of the pathways he first described. These facts have prompted suggestions that Mahaim should be deprived of his eponym. It is agreed that specificity is required when describing the pathways that produce the disordered cardiac conduction, and that the identified pathways should now be described in an attitudinally appropriate fashion. The authors remain to be convinced that understanding will be enhanced simply by discarding the term 'Mahaim physiology' from the lexicon. It is fascinating to look back at the history of accessory atrioventricular junctional conduction pathways outside the normal accessory atrioventricular conduction system, and their possible role in rhythm disturbances. It took both the anatomist and the clinical arrhythmologist quite some time to understand the complex anatomical architecture and the ensuing electrophysiological properties. Over the years, the name Mahaim was often mentioned in those discussions, although these pathways were not the ones that produced the eponym. The reason for this review, therefore, is to present relevant information about the person and what followed thereafter.

How does the cardiac impulse pass from the sinus to the atrioventricular node?

More than a century has passed since Tawara demonstrated the presence of the insulated pathways that extend from the "knoten" at the base of the atrial septum to their ramifications at the ventricular apexes. Having initially doubted the existence of the atrioventricular bundle until reading the monograph produced by Tawara, Keith, together with Flack, soon revealed the presence of the sinus node. Shortly thereafter, Thorel suggested that a special system might be found within the atrial walls, connecting the newly discovered atrial nodes. This prompted the convening of a special session of the German Pathological Society in 1910. The consensus was that no tracts existed within the atrial walls, with Aschoff and Mönckeberg establishing criteria to be met by those proposing recognition of "specialized" atrial conducting pathways. None of those who subsequently proposed the presence of such pathways have discussed their findings on the basis of the criteria established at the meeting of 1910. It remains the case, nonetheless, that drawings continue to be offered by cardiological experts showing narrow pathways within the atrial walls that parallel the arrangement used to show the ventricular conduction pathways. A similar drawing adorns the front cover of Heart Rhythm Journal. We are unaware of any evidence supporting the presence of pathways as illustrated existing within the overall walls of the atrial chambers. In this review, we summarize the evidence that shows, instead, that it is the aggregation of the working atrial cardiomyocytes within the atrial walls that underscores preferential anisotropic interatrial conduction.

Anatomy of the left atrial appendage for the interventional cardiologist.

Exact knowledge of the anatomy of the left atrial appendage (LAA) is crucial for LAA isolation by catheter ablation and for interventional LAA occlusion in patients with atrial fibrillation. This review outlines the current anatomical understanding of LAA morphology from ostium to distal lobes, myocardial fiber orientation and wall structure, and adjacent structures such as the left upper pulmonary vein with the Coumadin ridge, the circumflex artery with its side branches, the aortic root, pulmonary artery, and the pericardial space. Insight into these details will facilitate these interventions and reduce the risk of complications.

Anatomical knowledge for the ablation of left and right atrial flutter.

The different forms of atrial flutter (AFL) and atrial macroreentrant tachycardias are strongly related to the atrial anatomy in structurally normal atria, and even more so in patients with dilated chambers or with previous interventions. Atrial anatomy, macro- and microscopic tissue disposition including myocardial fibers, conduction system and connective tissue is complex. This review summarizes knowledge of atrial anatomy for the interventional electrophysiologist to better understand the pathophysiology of and ablation options for these complex arrhythmias, as well as to perform catheter ablation procedures safely and effectively.

Diversity and complexity of the cavotricuspid isthmus in rabbits: A novel scheme for classification and geometrical transformation of anatomical structures.

The aim of this study was to describe the morphology of the cavotricuspid isthmus (CTI) in detail and introduce a comprehensive scheme to describe the topology of this region based on functional considerations. This may lead to a better understanding of isthmus-dependent flutter and fibrillation and to improved intervention strategies. We used images of the cavotricuspid isthmus from 52 rabbits of both sexes with a median weight of 3.40 ± 0.93 kg. The area of the CTI was 124.25 ± 42.14 mm2 with 53.28 ± 21.13 mm2 covered by pectinate muscles connecting the terminal crest and the vestibule. Isthmus length decreased from inferolateral (13.09 ±2.14 mm) to central (9.85 ± 2.14 mm) to paraseptal (4.88 ± 1.96 mm) resembling the overall human geometry. Ramification sites of pectinate muscles were identified and six levels dividing the CTI from posterior to anterior were introduced. This allowed the classification of pectinate muscle segments based on the connected ramification level. To account for the high inter-individual variations in size and shape, the CTI was projected onto a normalized reference frame using bilinear transformation. Furthermore, two measures of complexity were introduced: (i) the ramification index, which reflects the total number of muscle segments connected to a ramification site and (ii) the complexity index, which reflects the type of ramification (branching or merging site). Topological analysis showed that the complexity of the pectinate muscle network decreases from inferolateral to paraseptal and that the number of electrically uncoupled parallel pathways increases in the central section between the terminal crest and the vestibule which introduces potential reentry pathways.

Miniseries 2-Septal and paraseptal accessory pathways-Part IV: Inferior paraseptal accessory pathways-lessons from surgical and catheter ablation.

Surgeons and electrophysiologists performing accessory pathway ablation procedures have used the term 'posteroseptal' region. This area, however, is neither septal nor posterior, but paraseptal and inferior; paraseptal because it includes the fibro-adipose tissues filling the pyramidal space and not the muscular septum itself and inferior because it is part of the heart adjacent to the diaphragm. It should properly be described, therefore, as being inferior and paraseptal. Pathways in this region can be ablated at three areas, which we term right inferior, mid-inferior, and left inferior paraseptal. The right- and left inferior paraseptal pathways connect the right and left atrial vestibules with the right and left paraseptal segments of the parietal ventricular walls. The mid-inferior paraseptal pathways take a subepicardial course from the myocardial sleeves surrounding the coronary sinus and its tributaries. Our review addresses the evolution of the anatomical concept of the inferior paraseptal region derived from surgical and catheter ablation procedures. We also highlight the limitations of the 12-lead electrocardiogram in identifying, without catheter electrode mapping, which are the pathways that can be ablated without a coronary sinus, or left heart approach.

Miniseries 1-Part III: 'Behind the scenes' in the triangle of Koch.

AIMS: To take full advantage of the knowledge of cardiac anatomy, structures should be considered in their correct attitudinal orientation. Our aim was to discuss the triangle of Koch in an attitudinally appropriate fashion. METHODS AND RESULTS: We reviewed our material prepared by histological sectioning, along with computed tomographic datasets of human hearts. The triangle of Koch is the right atrial surface of the inferior pyramidal space, being bordered by the tendon of Todaro and the hinge of the septal leaflet of the tricuspid valve, with its base at the inferior cavotricuspid isthmus. The fibro-adipose tissues of the inferior pyramidal space separate the atrial wall from the crest of the muscular interventricular septum, thus producing an atrioventricular muscular sandwich. The overall area is better approached as a pyramid rather than a triangle. The apex of the inferior pyramidal space overlaps the infero-septal recess of the subaortic outflow tract, permitting the atrioventricular conduction axis to transition directly to the crest of the muscular ventricular septum. The compact atrioventricular node is formed at the apex of the pyramid by union of its inferior extensions, which represent the slow pathway, with the septal components formed in the buttress of the atrial septum, thus providing the fast pathway. CONCLUSIONS: To understand its various implications in current cardiological catheter interventions, the triangle of Koch must be considered in conjunction with the inferior pyramidal space and the infero-septal recess. It is better to consider the overall region in terms of a pyramidal area of interest.

Miniseries 2-Septal and paraseptal accessory pathways-Part I: The anatomic basis for the understanding of para-Hisian accessory atrioventricular pathways.

AIMS: Although the anatomy of the atrioventricular conduction axis was well described over a century ago, the precise arrangement in the regions surrounding its transition from the atrioventricular node to the so-called bundle of His remain uncertain. We aimed to clarify these relationships. METHODS AND RESULTS: We have used our various datasets to examine the development and anatomical arrangement of the atrioventricular conduction axis, paying particular attention to the regions surrounding the point of penetration of the bundle of His. It is the areas directly adjacent to the transition of the atrioventricular conduction axis from the atrioventricular node to the non-branching atrioventricular bundle that constitute the para-Hisian areas. The atrioventricular conduction axis itself traverses the membranous part of the ventricular septum as it extends from the node to become the bundle, but the para-Hisian areas themselves are paraseptal. This is because they incorporate the fibrofatty tissues of the inferior pyramidal space and the superior atrioventricular groove. In this initial overarching review, we summarize the developmental and anatomical features of these areas along with the location and landmarks of the atrioventricular conduction axis. We emphasize the relationships between the inferior pyramidal space and the infero-septal recess of the subaortic outflow tract. The details are then explored in greater detail in the additional reviews provided within our miniseries. CONCLUSION: Our anatomical findings, described here, provide the basis for our concomitant clinical review of the so-called para-Hisian arrhythmias. The findings also provide the basis for understanding the other variants of ventricular pre-excitation.