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Activation pattern of the avian left ventricle during ventricular pacing.


Objective: This study was planned to investigate ventricular myocardial myocardial /myo·car·di·al/ (-kahr´de-al) pertaining to the muscular tissue of the heart.


pertaining to the muscular tissue of the heart (the myocardium).
 excitation in birds in which Purkinje fibres penetrate into the ventricular wall and reach the epicardium epicardium /epi·car·di·um/ (-kahr´de-um) the visceral pericardium.

n. pl.
 to advance our knowledge about the evolution of the ventricular activation process in vertebrates.

Methods: A depolarization depolarization /de·po·lar·iza·tion/ (de-po?lahr-i-za´shun)
1. the process or act of neutralizing polarity.

2. in electrophysiology, reversal of the resting potential in excitable cell membranes when stimulated.
 pattern of the left ventricular free wall in seven open-chest laying hens was mapped by 14 seven-electrode plunge needles under ventricular pacing from different sites.

Results: Duration of activation of the left ventricular free wall is significantly increased during ventricular ectopic ectopic /ec·top·ic/ (ek-top´ik)
1. pertaining to ectopia.

2. located away from normal position.

3. arising from an abnormal site or tissue.

 excitation as compared with sinus rhythm sinus rhythm
A normal cardiac rhythm proceeding from the sinoatrial node.
. Its lowest increase occurs during subendocardial pacing of the middle part of the left ventricle left ventricle
The chamber on the left side of the heart that receives the arterial blood from the left atrium and contracts to force it into the aorta.
, but its greatest increase is observed during subepicardial pacing of the left ventricular base. Multifocality and mosaicity of depolarization of the left ventricular myocardium myocardium /myo·car·di·um/ (-kahr´de-um) the middle and thickest layer of the heart wall, composed of cardiac muscle.

hibernating myocardium  see myocardial hibernation, under
 are expressed in a considerably less degree during ventricular pacing in comparison with sinus rhythm.

Conclusion: Ectopic excitation of avian heart ventricles Ventricles
The two chambers of the heart that are involved in pumping blood. The right ventricle pumps blood into the lungs to receive oxygen. The left ventricle pumps blood into the circulation of the body to deliver oxygen to all of the body's organs and tissues.
 occurs mostly due to successive spreading of the activation wave from a pacing site during both ipsi- and contraventricular pacing. During ipsiventricular pacing at least, ectopic excitation of the heart ventricles with the "rich" Purkinje network behaves like one of the mammalian ventricles with the subendocardial Purkinje network. (Anadolu Kardiyol Derg 2007.7 Suppl 1, 95-7)

Keywords: myocardium, heart ventricles, pacing, bird, chicken


In theory, in multifocal multifocal /mul·ti·fo·cal/ (mul?te-fo´k'l) arising from or pertaining to many foci.

Relating to or arising from many foci.
 ("flash") activation of avian ventricular myocardium (1-3) caused by the specific distribution of Purkinje fibres that penetrate deeply into the ventricular wall and run close to the epicardium (1, 4, 5) there must not appear considerable disturbances in the myocardial activation pattern under local disturbances of conduction. However, in spite of the "rich" network of Purkinje fibres, various disturbances of electrical conduction including ventricular extrasystoles are observed in avian hearts (6-8). Thus, an avian heart is an interesting and unique model to investigate mechanisms of disturbances of electrical conduction within the heart and to advance our knowledge about the evolution of the cardiac electrical function in vertebrates. A study of myocardial excitation features under ventricular premature beats in animals with the multifocal activation pattern of the ventricular myocardium is of significant interest for comparative and comprehensive electrocardiology. The aim of the present study is to investigate a myocardial activation pattern of the chicken left ventricular (LV) free wall during ventricular pacing.


In adult laying hens Gallus Gallus (Caius Vibius Trebonianus Gallus) (găl`əs), d. 253 or 254, Roman emperor after 251. He fought in the eastern campaign that proved fatal to Decius.  gallus domesticus (n=7; age, 9-12 months; body mass, 1.3-1.8 kg) purchased from a poultry farm, general anesthesia Anesthesia, General Definition

General anesthesia is the induction of a state of unconsciousness with the absence of pain sensation over the entire body, through the administration of anesthetic drugs.
 was induced with sodium thiopental (120-150 mg/kg i.m.). The chicken was then placed ventral side up, tracheotomized and artificially ventilated ven·ti·late  
tr.v. ven·ti·lat·ed, ven·ti·lat·ing, ven·ti·lates
1. To admit fresh air into (a mine, for example) to replace stale or noxious air.

. The heart was exposed through a bilateral approach (2). After pericardectomy, fourteen plunge needles (diameter 0.35-0.40 mm), each containing seven electrodes (interelectrode distance 0.65, 0.80 or 1 mm), was inserted into the LV free wall according to one of schemes (Fig. 1A, B). Steel needle electrodes were inserted subcutaneously to obtain standard bipolar limb lead electrocardiograms.


Limb lead electrocardiograms and intracardiac intracardiac /in·tra·car·di·ac/ (-kahr´de-ak) within the heart.

Within the heart.


within the heart.
 electrograms were amplified and recorded by means of a custom-designed 128-channel computerized multiplexer recording system with a bandwidth of 0.05-1000 Hz at a sampling rate of 4000 Hz and an accuracy of 16 bits. Ventricular pacing was performed by means of right-angled 3-7 V impulses of 3 ms duration at a frequency of 3 Hz. The base (n=6), middle part (n=6) and apex (n=7) of the left ventricle and the base (n=3) of the right ventricle (Fig. 1C) were paced from different depths (epicardium, mid-myocardium, endocardium endocardium /en·do·car·di·um/ (-kahr´de-um) the endothelial lining membrane of the cavities of the heart and the connective tissue bed on which it lies.

n. pl.
). Body temperature was kept in the range of 41-42[degrees]C, the heart was prevented from cooling and drying by means of a warm (41-42[degrees]C) 0.85% NaCl solution. After the experiment, the heart was excised and cut to specify positions of plunge needles.

Local activation times relative to the S peak in the II limb lead electrocardiogram electrocardiogram /elec·tro·car·dio·gram/ (-kahr´de-o-gram?) a graphic tracing of the variations in electrical potential caused by the excitation of the heart muscle and detected at the body surface.  (sinus rhythm) or to the onset of a pacing impulse (ventricular pacing) were determined automatically on the basis of the minimum of the first derivative of the (IRS An abbreviation for the Internal Revenue Service, a federal agency charged with the responsibility of administering and enforcing internal revenue laws.  complex and reviewed and revised if necessary. During construction of isochronal i·soch·ro·nal   or i·soch·ro·nous
1. Equal in duration.

2. Characterized by or occurring at equal intervals of time.
 activation maps, all local activation times were referred to the lowest local activation time, which was considered to be the zero value. Duration of activation of the LV free wall was defined as the difference of the greatest and lowest local activation time.

Data are expressed as mean [+ or -] standard deviation. Paired t-test was used for comparisons. P value <0.05 was considered statistically significant.


Depending on the pacing site, duration of activation of the LV free wall was 14.8 [+ or -] 2.8 to 22.4 [+ or -] 3.3 ms being 9.3 [+ or -] 1.0 ms during the sinus rhythm (150 [+ or -] 14 beats per minute beats per minute Cardiac pacing The unit of measure for the frequency of heart depolarizations or contractions each minute–or pulse rate ) (Table 1). Its lowest (1.6 [+ or -] 0.5 times as much) and greatest (2.4 [+ or -] 0.3 times as much) increases were observed under endocardial endocardial /en·do·car·di·al/ (-kahr´de-al)
1. situated or occurring within the heart.

2. pertaining to the endocardium.


1. situated or occurring within the heart.

 pacing of the LV middle part and epicardial epicardial

pertaining to the visceral pericardium (epicardium) or to the epicardia.

epicardial receptors
receptors in the left ventricle adapted to respond to stretch and chemical stimulants.
 pacing of the LV base, respectively. Under sinus rhythm, activation propagated in radial and tangential directions from foci appearing within the LV free wall myocardium during 2-3 ms after the onset of its depolarization (Fig. 2A); a general sequence of activation common to all birds was identified, with interindividual differences in specific details of the patterns. In contrast, multifocality and mosaicity of the activation pattern were expressed in a considerably less degree during ectopic excitation, and the LV free wall was mostly excited due to successive spreading of the depolarization wave from a pacing site (Fig. 2B, 2C). Activation patterns were similar during ventricular pacing from different depths (epicardial, mid-myocardial, endocardial) of each pacing area.


Within the LV free wall, activation spread from the base to the apex and from the left to the right during pacing from the LV base. The opposite sequence was observed during pacing from the LV apex and the right ventricular base (Fig. 2B, C). Discussion

The non-multifocal activation pattern of the ventricular myocardium during ventricular pacing might signify that Purkinje fibres do not participate in myocardial activation. This supposition is proved by following facts: 1) similar activation patterns of the left ventricle during ventricular pacing from different depths; 2) similar activation patterns of the LV free wall during ipsiventricular (LV apical apical /ap·i·cal/ (ap´i-k'l) pertaining to an apex.

1. Relating to the apex of a pyramidal or pointed structure.

) pacing and contraventricular (right ventricular basal) pacing; 3) activation within the left ventricle spreads from the apex to the base during apical pacing in both chickens and animals with other ventricular activation patterns (3, 9, 10, 11).

On the other hand, participation of the Purkinje network in ventricular activation of the avian heart during ventricular pacing can not be fully denied for following reasons: 1) the obtained values of duration of activation during ventricular pacing (Table 1) are 2-4 times less than values (50-60 ms) calculated taking into account the conduction velocity in the contractile contractile /con·trac·tile/ (kon-trak´til) able to contract in response to a suitable stimulus.

Capable of contracting or causing contraction, as a tissue.
 myocardium of -0.4 m/s and the length of the heart ventricles of 20-25 mm in chickens; 2) in so me cases, foci of activation were located ahead of the main depolarization front and their merging for one millisecond One thousandth of a second. See space/time and ohnosecond.

(unit) millisecond - (ms) One thousandth of a second, one thousand microseconds. A long time for a modern computer.
 were observed in the ventricular wall during pacing; 3) duration of activation of the LV free wall is minimal during pacing of subendocardial layers of the middle part of the LV free wall that are the most "rich" in Purkinje fibres as compared to the located on a level with them intramural intramural /in·tra·mu·ral/ (-mu´r'l) within the wall of an organ.

Occurring or situated within the walls of a cavity or organ.
 and subepicardial layers (Table 1). A probable cause of insignificant participation of the Purkinje network in myocardial activation during ventricular pacing is that an electrical impulse spreads along Purkinje fibres in the retrograde direction as on a cable without interaction with neighbouring myocardial cells. Probably, during ectopic excitation most of the ventricular myocardium is depolarized by the myogenic myogenic /my·o·gen·ic/ (-jen´ik)
1. pertaining to myogenesis.

2. originating in myocytes or muscle tissue.

my·o·gen·ic or my·o·ge·net·ic
 way (muscle fibre to muscle fibre) with simultaneous activation of some parts of it through the Purkinje network. Earlier, suggestions about the combined way of myocardial activation during ventricular ectopic excitation were made for ungulates ungulates, ungulata

animals with hooves; cattle, sheep, goat, pig, horse and many wild and other domesticated species.
, in particular sheep (9), goats (12), reindeers (Shmakov DN, unpublished), having the "rich" Purkinje network (13-16). Taking into account the similarity of ventricular activation patterns in chickens (2) and wild avian species (1, 3), it can be suggested that activation of the ventricles during ectopic excitation in the latter occurs in the same manner as in chickens.


The data obtained reveals that multifocality and mosaicity of activation of the LV myocardium in chickens are expressed in a considerably less degree during ventricular pacing in comparison with sinus rhythm. Ectopic excitation of the heartventricles in birds occurs mostly due to successive spreading of the activation wave from a pacing site during both ipsi- and contraventricular pacing. The results convince that during ipsiventricular pacing at least, ectopic excitation of the heart ventricles with the "rich" Purkinje network behaves like one of the mammalian ventricles with the subendocardial Purkinje network.


This study was supported by the Russian Foundation for Basic Research (grant N 06-04-48022), the Grant (MK-5347.2006.4) of President of the Russian Federation, and the program of support for basic research performed in the Ural Branch of the Russian Academy of Sciences Russian Academy of Sciences (Russian: Росси́йская Акаде́мия Нау́к,  in association with the Far Eastern Branch of RAS (1) See network access server.

(2) (Remote Access Service) A Windows NT/2000 Server feature that allows remote users access to the network from their Windows laptops or desktops via modem. See RRAS and network access server.
 during 2005-2007.


(1.) Shmakov DN, Kliushina IV, Roshchevskii MP. Sequence of the spread of excitation in bird heart ventricles. Fiziol Zh SSSR SSSR Society for the Scientific Study of Religion
SSSR Society for the Scientific Study of Reading
SSSR Smallest Set of Smallest Rings (chemistry)
SSSR Sojus Sowjetskich Sozialistitscheskich Respublik (USSR; Russian) 
 Im I M Sechenova 1979; 65: 872-80.

(2.) Kharin SN, Shmakov DN, Antonova NA, Roshchevskii MP. Formation of cardioelectric field on the body surface at a period of activation of ventricular myocardium in the chicken Gallus domesticus. J. Evol Biochem Physiol 2001; 37:161-9.

(3.) Roshchevsky MP, Shmakov DN. Excitation of the heart. Moscow: Nauka; 2003.

(4.) Truex RC, Smythe MG. Comparative morphology of the cardiac conduction tissue in animals. Ann NY Acad Sci 1965;127:19-33.

(5.) Prosheva V, Rapota I. Structural and functional organisation of atrioventricular atrioventricular /atrio·ven·tric·u·lar/ (-ven-trik´u-ler) pertaining to both an atrium and a ventricle of the heart.

adj. Abbr.
 conducting system in the avian heart. In: Shubert E, editor. Electrocardiology'87. Proceedings of the 14th International Congress on Electrocardiology; 1987 August 17-20; Berlin, Germany. Berlin: Academic-Verlag;1988. P.127-30.

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emanating from or pertaining to electrocardiography.

electrocardiographic monitoring
maintenance of a more or less continuous surveillance of a patient's cardiac status by means of electrocardiography.
 observation on spontaneously occurring arrhythmias in chickens. J Vet Med Sci 1996; 58: 953-61.

(7.) Olkowski AA, Classen HL. High incidence of cardiac arrhythmias in broiler broiler

a young (about 8 weeks old) male or female chicken weighing 3 to 3.5 lb.
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(9.) Hamlin RL, Smith CR, Redding Redding, city (1990 pop. 66,462), seat of Shasta co., N central Calif., on the Sacramento River; inc. 1872. A principal tourist center for a mountain and lake region, it also has lumbering, food-processing, and diverse manufacturing.  RW. Time-order of ventricular activation for premature beats in sheep and dogs. Am J Physiol 1960,198: 315-21.

(10.) Shmakov DN, Mostivenko KK, Roshchevskaia IM. Effect of localizati on of the ectopic excitation site on the sequence and duration of de polarization in the ventricular epicardium in dogs. Ross Fiziol Zh Im IM Sechenova 2000; 86: 681-7.

(11.) Sedmera D, Reckova M, de Almeida A, Sedmerova M, Biermann M, Volejnik J, et al. Functional and morphological evidence for a ventri cular conduction system in zebrafish and Xenopus hearts. Am J Physiol Heart Circ Physiol 2003; 284: H1152-60.

(12.) Durrer D, van der Tweel LH. Excitation of the left ventricular wall of the dog and goat. Ann NY Acad Sci 1957; 65: 779-803.

(13.) Abramson DI, Margolin S. Purkinje conduction network in the myocardium of the mammalian ventricle ventricle /ven·tri·cle/ (ven´tri-k'l) a small cavity or chamber, as in the brain or heart.ventric´ular

ventricle of Arantius  the rhomboid fossa, especially its lower end.
. J Anat 1936; 70: 250-9.

(14.) Kliushina IV. Distribution of cardiac conducting myocytes and elect rophysiological peculiarities of myocardium in birds and ungulates. In: Comparative Electrocardiology. Proceedings of the International Symposium; 1979 June; Syktyvkar, Russia. Leningrad: Nauka; 1981. p. 128-32.

(15.) Oosthoek PW, Viragh S, Lamers WH, Moorman AF. Immunohistoche mical delineation of the conduction system. II: The atrioventricular node and Purkinje fibres. Circ Res 1993; 73: 482-91.

(16.) Ansari A, Ho SY, Anderson RH. Distribution of the Purkinje fibres in the sheep heart. Anat Rec 1999; 254: 92-7.

Sergey Kharin, Nadezhda Antonova, Dmitry Shmakov

Laboratory of Cardiac Physiology, Institute of Physiology of the Komi Science Centre, Ural Branch of the Russian Academy of Sciences, Syktyvkar, Russia

Address for Correspondence: Dr. Sergey N. Kharin, Institute of Physiology of the Russian Academy of Sciences, 50 Pervomayskaya st., Syktyvkar, 167000, Komi Republic, Russia Phone/Fax: +7(8212)240085 E-mail:
Table 1. Duration (ms) of ectopic excitation of the left ventricular
free wall myocardium in chickens

Site of the
ectopic beat              Epicardium            Mid-myocardium

The apex of the         19.0 [+ or -]       19.4 [+ or -] 5.1 (a)
  left ventricle            5.4 (a)
The middle part of     19.3 [+ or -] 2.2   19.2 [+ or -] 3.6 (a, b)
  the left ventricle       (a, b, c)
The base of the        22.4 [+ or -] 3.3    21.4 [+ or -] 4.7 (a)
  left ventricle            (a, c)
The base of the        17.3 [+ or -] 2.2    20.0 [+ or -] 4.0 (a)
  right ventricle           (a, d)

Site of the
ectopic beat           Endocardium

The apex of the
  left ventricle       18.3 [+ or -] 3.8 (a)
The middle part of
  the left ventricle   14.8 [+ or -] 2.8 (a)
The base of the
  left ventricle       20.3 [+ or -] 5.1 (a)
The base of the
  right ventricle      18.4 [+ or -] 1.2 (a, d)

Mean [+ or -] standard deviation.

(a) p < 0.01 compared to sinus rhythm (9.3 [+ or -] 1.0 ms);
(b) p < 0.05 compared to endocardial stimulation;
(c) p < 0.05 compared to stimulation of the base
of the right ventricle;
(d) p < 0.05 compared to midmyocardial stimulation of the
middle part of the left ventricle

(Reproduced from Comparative Biochemistry and Physiology, 145A (4),
Kharin S., Antonova N., Shmakov D., Left ventricular myocardial
activation under ventricular paced beats in chickens Gallus
gallus domesticus, 540-545, [c] 2006 Elsevier Inc., with
permission from Elsevier.)
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Title Annotation:Original Investigation
Author:Kharin, Sergey; Antonova, Nadezhda; Shmakov, Dmitry
Publication:The Anatolian Journal of Cardiology (Anadolu Kardiyoloji Dergisi)
Article Type:Clinical report
Geographic Code:4EXRU
Date:Jul 1, 2007
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