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Architecture of fibers of the working myocardium and the sequence of excitation of heart ventricles of a pig A.S.Gulyaeva, I.M.Roshchevskaya, M.P.Roshchevsky Institute of Physiology, Komi Science Center, Ural Division, Russian Academy of Sciences, 50, Pervomayskaya st., Syktyvkar, 167982, Komi Republic, Russia. e-mail: [email protected] ABSTRACT The ventricular fibre architecture in pig has been studied by the method of level-by-level splitting of muscular bundles. In both ventricles, the muscular fascicles of the myocardium are arranged in three preferential directions through the ventricular mass. We differentiated three different layers of fibres: superficial, middle, and deep. The fibres of the superficial layer have spiral direction. The superficial fibres invaginate into the apex cordis and form the deep layer. The middle layer in both pig ventricles is represented by circumferential fibres, which are absent both on the apex of the right and left ventricles. Depending on the depth of location orientation of fibres is changed and differentiated in both ventricles. The fibers of the deep layer of the right ventricle are arranged obliquely on the free wall, while on the interventricular septum – lengthways the apical- basal heart axis. In the left ventricle the fibres of the deep layer are going spirally from the apex to the base of the heart. The sequence of depolarization of pig's heart ventricles was investigated and the comparison of orientation of the fibres of the working myocardium to the character of the excitation wave propagation was carried out. 1. INTRODUCTION Architecture of the conducting system and working myocardium determines the succession of excitation and contraction of heart ventricles. Conducting system takes the leading role in the chronotopography of the excitation process in the heart of warm-blooded animals and man. Heart ventricles of man and predators are characterized by “successive” type of activation. In ungulates (pig) the spatial organization of Purkinje fibres is a structural basis of polyfocal excitation of heart ventricles6. Information on the architecture of the working myocardium in animals of various classes is fragmentary. The heart of man2, 4 and dog7 is studied in greater detail. Spatial organization of fibres distribution in heart ventricles in pig is weakly studied. The problem of similarity or difference of the architecture of the working myocardium of pig and man is yet unsolved. In this relation the aim of our investigation is to study the architecture of the ventricular fibers structure for comparative-physiological assessment of the heart function. 2. METHODS The architecture of the working myocardium in 12 pigs of both sexes aged 8-9 months is studied. We used the method of layer-by-layer splitting of muscular fibers3. Hearts were boiled in running water for an hour. The direction of fibres of heart ventricles was photographed with the aid of digital video camera and then used the system of image analysis (Ista-VideoTest, St.Petersburg). Orientation of muscular fascicles was described relative to the longitudinal heart axis. The succession of depolarization of heart ventricles in pig was studied by way of multi-channel synchronous cardioelectrotopography. Cardiopotentials were recorded from the ventricular epicardium and intramural myocardial layers in pigs anaesthetized by sodium thiopental. Measurements were carried out by 128-channel system of simultaneous potential registration. On the basis of the obtained experimental data the three-dimensional patterns of ventricles' depolarization wavefronts were reconstructed. 3. RESULTS The criterion of layers separation was different direction of muscular fibers. Three layers were discovered: superficial (subepicardial), middle (intramural) and deep (subendocardial). 601 Superficial layer of fibres is common for both ventricles. Muscular fibres begin from fibrous skeleton at the base of the heart, spirally twist clockwise and form a curl at the left ventricle apex. Orifice of pulmonary artery is surrounded with a bundle of fibres attached to the fibrous skeleton. At the ventral side the superficial fibers of heart ventricles go obliquely from right to left continuously crossing the anterior interventriclular groove (Fig. 1a). At the dorsal side of ventricles the bundles of muscular fibers going from the posterior edge of the mitral valve go obliquely from left to right (Fig. 1b). The angle of inclination relative to the longitudinal heart axis is less. After crossing the posterior interventricular groove they intersect with fibres, beginning from the posterior edge of tricuspidal valve that go down from the base to the apex. In the area of the right ventricle apex the subepicardial fibers branch and give the beginning to the fibres of the deep layer. Middle layer of circumferential muscles is found in both heart ventricles in pig. We singled out high-lying and low-lying muscular bundles. High-lying circumferential fibers continuously go along the dorsal and lateral surface of both ventricles; while from the ventral side of the left ventricle they go deep forming the interventricular septum. At the right ventricle apex the circumferential fibres of the middle layer are not available. The transition of the superficial layers of the right ventricle to the deep ones is observed in this area. At the dorsal side the low-lying muscular fibres are not continuous. The fibres of the right and left ventricle form the interventriclular septum and corresponding free walls. In the right ventricle the fibres go transversally to the longitudinal heart axis. In the left ventricle the low-lying fibres penetrate deep and twist as a spiral. Spires of the spiral approach in the medial part of the interventriclular septum and go away in the direction of the apex and the base of the free wall reminding in form the truncated cone (Fig.2). At the left ventricle apex the middle layer of fibres is also absent. The fibres of the superficial and low-lying middle layer participate in the formation of the deep layer. Deep muscular bundles from the side of the interventricular septum are oriented along the longitudinal heart axis, while from the side of the free wall – obliquely (Fig. 3a). Reaching the ventricle base the bundles of muscular fibres attach to the fibrous ring of the tricuspidal valve. In the left ventricle the muscular fibres of the superficial and low-lying middle layers combine and, twisting inside, give the beginning to the deep fibres. From the apex to the base the fibres of the deep layer spirally rise relative to the longitudinal heart axis (Fig. 3b) and attach to the fibrous ring of the mitral valve. The excitation wave in heart ventricles in pig spreads from multiple foci of depolarization located in subendocardial and intramural myocardial layers. From multiple intramural foci of depolarization the excitation wave spreads successively. The multi-focused excitation of heart ventricles in pig results in depolarization of myocardium during a short time period. The excitation wave in the intramural layers of heart ventricles spreads not only from endocardium to epicardium, but also in the reverse direction – from subepicardial and intramural zones of depolarization to endocardium. Multiple foci of depolarization occurring on subepicardium are caused by both the excitation wave coming to subepicardium from deeper layers and the activation from the subepicarial Purkinje fibres. There are plenty of areas of subepicardial activation located on both heart ventricles. 4. DISCUSSION Analysis of the direction of superficial fibers has shown that on the anterior surface of pig’s heart the ventricles are oriented as in man from right to left, but differ by the angle of inclination relative to the longitudinal heart axis4. On the posterior surface the difference in the direction of fibers is more clearly expressed: in man the fibres on the posterior surface go continuously from left to right. The difference in the direction of muscular fibres maybe related to different form of heart ventricles. It is known that in pig greater part of the heart is formed the left ventricle, which means that the interventricular septum is located more to the right, while the apex is composed of entirely left ventricular musculature. In man the left ventricle is less dominant, as a result the interventricular septum occupies a more central position within the section, while the heart apex is formed of both right and left ventricles1. It was shown that the middle layer in both ventricles of pig’s heart is represented by the circumferential fibres. However, we have different information on the availability of middle layer in the right ventricle. Some authors distinguish a thin sheet of fibres between the superficial and the deep layers2, others do not discover it5. 602 Our investigation has shown that the left ventricle of pig’s heart has more complexly twisted spires of the spiral, their form reminding the truncated cone, unlike that of man. We revealed that in pig’s heart deep muscular bundles of the left ventricle and deep fibres of the free wall of the right ventricle are oriented obliquely from the apex to the base, and from the side of the interventricular septum of the right ventricle – longitudinally, while in both ventricles of man’s heart deep fibres are oriented along the longitudinal heart axis2, 5. The carried out investigation has shown that pig’s heart ventricles are characterized by essential differences in the orientation of the working fibres of superficial, middle and deep layer as compared to that in man. ACKNOWLEDGMENTS The work is supported by the grant of the scientific school of the academician M.P.Roshchevsky Sci.Sch.-759.2003.4, Russian Science Support Foundation for I.M.Roshchevskaya. REFERENCES 1. Crick S.J., Sheppard M.N., Ho S.Y., Gebstein L., Anderson R.N. Anatomy of the pig heart: comparisons with normal human cardiac structure. J. Anat., 1998, V. 193, p. 105 – 119. 2. Fernandez-Teran M.A. and Hurle J.M. Myocardial fiber architecture of the human heart ventricles. Anat. Rec., 1982, V. 204, p. 137 – 147. 3. Puff A. Der funktionelle Bau der Herzkammern. Georg Thieme Verlag. Stuttgart. 1960, 88 p. 4. Sanchez – Quintana D., Garcia – Martinez V., Climent V., Hurle J.M. Morphological changes in the normal pattern of ventricular myoarchitecture in the developing human heart. Anat. Rec., 1995, V. 243, № 4, p. 483 - 495. 5. Sanchez – Quintana D., Climent V., Ho S.Y., Anderson R.H. Myoarchitecture and connective tissue in heart with tricuspid atresia. Heart, 1999, V. 81, p. 182 – 191. 6. Shmakov D.N., Roshchevsky M.P. Activation of myocardium. Syktyvkar, Publishing hous of Institute of Physiology, Komi Science Center, Ural Division Russian Academy of Sciences, 1997, 166 p. 7. Streeter D.D., Spotnitz H.M., Patel D.P. Fiber orientation in the canine left ventricle during diastole and systole. Circ. Res., 1969, V. 24, № 3, p. 339 – 347. 603