Towards new understanding of the heart structure and function

Abstract

Structure and function in any organ are inseparable categories, both in health and disease. Whether we are ready to accept,
or not, many questions in cardiovascular medicine are still pending, due to our insufficient insight in the basic science.
Even so, any new concept encounters difficulties, mainly arising from our inert attitude, which may result either in unjustified
acceptance or denial. The ventricular myocardial band concept, developed over the last 50 years, has revealed unavoidable
coherence and mutual coupling of form and function in the ventricular myocardium. After more than five centuries long debate
on macroscopic structure of the ventricular myocardium, this concept has provided a promising ground for its final understanding.
Recent validations of the ventricular myocardial band, reviewed here, as well as future research directions that are pointed
out, should initiate much wider scientific interest, which would, in turn, lead to reconciliation of some exceeded concepts
about developmental, electrical, mechanical and energetical events in human heart. The benefit of this, of course, would be
the most evident in the clinical arena.

Full text

DOI: 10.1016/j.ejcts.2004.11.026
2005;27:191-201 Eur J Cardiothorac Surg
Francesc Carreras-Costa, A. Flotats, Juan Cosin-Aguillar and Han Wen
Francisco Torrent-Guasp, Mladen J. Kocica, Antonio F. Corno, Masashi Komeda,
Towards new understanding of the heart structure and function
This information is current as of October 18, 2010
http://ejcts.ctsnetjournals.org/cgi/content/full/27/2/191
located on the World Wide Web at:
The online version of this article, along with updated information and services, is
ISSN: 1010-7940.
European Association for Cardio-Thoracic Surgery. Published by Elsevier. All rights reserved. Print
for Cardio-thoracic Surgery and the European Society of Thoracic Surgeons. Copyright © 2005 by
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Invited paper
Towards new understanding of the heart structure and function
Francisco Torrent-Guasp
a
, Mladen J. Kocica
b,
*, Antonio F. Corno
c
, Masashi Komeda
d
,
Francesc Carreras-Costa
e
, A. Flotats
f
, Juan Cosin-Aguillar
g
, Han Wen
h
a
Denia, Alicante, Spain
b
Clinic for Cardiac Surgery, Institute for Cardiovascular Diseases, UC Clinical Centre of Serbia, 8th Kosta Todorovic St., 11000 Belgrade, Serbia and Montenegro
c
Alder Hey Royal Children Hospital, Liverpool, UK
d
Department of Cardiovascular Surgery, Kyoto University Graduate School of Medicine, Kyoto, Japan
e
Department of Cardiology, Cardiac Imaging Unit, Hospital Sant Pau, Barcelona, Spain
f
Department of Nuclear Medicine, Hospital Sant Pau, Barcelona, Spain
g
Cardiocirculatory Unit, Investigation Centre, University Hospital La Fe, Valencia, Spain
h
Laboratory of Cardiac Energetics, National Heart Lung and Blood Institute, NIH, Bethesda, MD, USA
Received 6 September 2004; received in revised form 25 November 2004; accepted 26 November 2004; Available online 28 December 2004
Summary
Structure and function in any organ are inseparable categories, both in health and disease. Whether we are ready to accept, or not, many
questions in cardiovascular medicine are still pending, due to our insufficient insight in the basic science. Even so, any new concept encounters
difficulties, mainly arising from our inert attitude, which may result either in unjustified acceptance or denial. The ventricular myocardial band
concept, developed over the last 50 years, has revealed unavoidable coherence and mutual coupling of form and function in the ventricular
myocardium. After more than five centuries long debate on macroscopic structure of the ventricular myocardium, this concept has provided a
promising ground for its final understanding. Recent validations of the ventricular myocardial band, reviewed here, as well as future research
directions that are pointed out, should initiate much wider scientific interest, which would, in turn, lead to reconciliation of some exceeded
concepts about developmental, electrical, mechanical and energetical events in human heart. The benefit of this, of course, would be the most
evident in the clinical arena.
q2004 Elsevier B.V. All rights reserved.
Keywords: Ventricle; Anatomy; Myocardium; Physiology
“Hos igitur quasi vestigiis odorati tradendae medicinae initium
ab humano corpore ducemus, quod et artis subjectum existit, et
omnium primum sensibus occurrit notissimum: a quo dein per
minima quaeque deducti ad ea denique mentis impulsu feremur,
quae cogitatione sola comprehendi possunt”
“We shall start the beginning of the teaching of medicine from
the human body, which is both the subject of the art of medicine
and, first of all, it comes most clearly under our senses. Then
from there, led through all the minutiae, we shall be finally
carried by an impulse of the mind, to those things which can be
understood by thinking alone.”
Jean Fernel (1497–1558), physician to King Henri II of
France [13]
1. Introduction
One of the most important scientific missions in this
century is integration of basic research with clinical
medicine. With substantial achievements at genetic, mol-
ecular and cellular levels, during the past few decades,
recent advances in elucidating myocardial structure and
function [1–5] have made a paradigm shift in research and
provided a promising ground for the new integrative knowl-
edge of the heart structure and function.
The ancient enigma of myocardial architecture is finally
solved. Discovery of the ventricular myocardial band (VMB,
Fig. 1) revealed unavoidable coherence and mutual coupling
of form and function in the ventricular myocardium, urging for
reconciliation of some exceeded concepts about electrical,
mechanical and energetical events in human heart [1–5].
Careful integration of this knowledge is not of merely
academic importance, but is also the essential prerequisite
in clinical evaluation and treatment of different heart
diseases [6]. The best example are recent modifications
and enrichments in heart failure conceptions, leading to new
therapies addressed to ‘disease and not to symptoms’ [7,8].
1010-7940/$ - see front matter q2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.ejcts.2004.11.026
European Journal of Cardio-thoracic Surgery 27 (2005) 191–201
www.elsevier.com/locate/ejcts
* Corresponding author. Tel.: C381 11 367 0609; fax: C381 11 361 0880.
E-mail address: kocica@sezampro.yu (M.J. Kocica).
URL: www.ctsnet.org/home/mkocica.
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Kresh and Armour [9] advised that ‘heart should be
considered as self-regulating functional system, which is
greater than the sum of its constitutive parts’. This general
statement should be adopted as a common standpoint for
those, who intend to participate and contribute in integra-
tive heart researches.
Today, more and more, parts of the road to excellence
are leading to collaboration between the scientists from
quite different branches. Many of them, from the mathe-
maticians to the cardiac surgeons, are already discussing
about heart, in very comprehensive manner [6–10]. But
even so, there are still many points of misunderstandings
and disagreements. Most of them are emanating from
different research technologies applied, while some others
[11,12], are still difficult to explain. To overcome these
diversities, we should first be aware that scientific theories
are validated by empirical testing against physical obser-
vations, rather than by simple judging of their logical
compatibility with the available data. But, when we reach
the point upon which our scientific perception is substan-
tiated, as Fernel suggests, ‘we shall be finally carried by an
impulse of the mind, to those things which can be
understood by thinking alone’ [13].
Thus, in order to fully understand complex three-dimen-
sional architecture of the ventricular myocardium and its
functional significance, some essential and scientifically
validated facts should be summarized and emphasized again.
2. Spatial organization of the ventricular myocardial
fibers—historical scope
The problem of the macroscopic structure of the ven-
tricular myocardium has remained unsolved since the XVI
century, when anatomy became an authentic science. The
spatial organization of the myocardial fibers has been
represented, as James Bell Pettigrew [14] wrote in 1864:
‘an arrangement so unusual and perplexing, that it has long
been considered as forming a kind of Gordian knot of
Anatomy. Of the complexity of the arrangement I need not
speak further than to say that Vesalius, Haller, and DeBlain-
ville, all confessed their inability to unravel it’.
Reviewing the scientific reports, made along the last five
centuries, for those who have made a substantial contri-
bution to the knowledge of rule-based assignment of
different trajectories followed in the space by each
ventricular myocardial fiber, one realizes that only Richard
Lower’s work [15], reported in 1669, provides still irremo-
vable anatomical fact. He describes that, in the left
ventricular wall, two groups of fibers could be distinguished.
Superficial fibers, that are coming from the base and run sub-
epicardically, suffering a reflection at the apex, after which,
they become sub-endocardial, coming back towards the
base. The second group (i.e. deep fibers), are located in the
middle of the ventricular wall thickness, and run in a
progressive transverse fashion. All successive contributions,
including those made in the first half of XX century, more or
less, limit themselves to repeat, although in different terms,
these Lower’s findings.
Thus, in 1749, Senac [16] arrives to the conclusion that
the outer fibers, corresponding to the external and internal
surfaces of the ventricular wall, run in an almost vertical
direction, meanwhile, the inner ones are extended circularly
in a near horizontal plane.
Later on, in 1792, Wolf [17] observes that within the deep
fibers could be distinguished two trajectories. The external
one—along which deep fibers run progressively more near to
the direction followed by the sub-epicardial fibers, and the
internal one—progressively inclining toward the sub-endo-
cardial ones.
In 1823, Gerdy [18], also according to the description of
the English pioneer [15], claims that the inner fibers of the
ventricular wall, coming from the ventricular base, fit with
uninterrupted succession into the outer, sub-epicardial and
sub-endocardial fibers, making an opened ‘figure of eight’.
Weber [19] states once more, in 1831, that the more
superficial fibers (i.e. sub-epicardial and sub-endocardial),
coming from the ventricular apex, irradiate to arrive the
ventricular base.
Ludwig [20] has noted, in 1849, that uninterrupted fibers
are forming a closed ‘figure of eight’ while passing around
left ventricle, changing their angle smoothly from the
epicardium to the endocardium.
In his comprehensive study, published in 1863, Pettigrew
[14] establishes the existence of seven distinct muscular
layers, which can be individualized by the progressive
change in direction of their fibers, from the epicardium to
the endocardium.
Krehl [21] conceptualized in 1891 the ‘Triebwerk’, as
nested set of fiber paths, describing a sub-epicardial and sub-
endocardial continuity at the level of the mitral orifice, and
at the apex of the left ventricle, where they invaginate.
MacCallum [22] in 1900 and later on, his teacher Mall
[23] in 1911, described separately, again according to
Lower, that the ventricular wall is made of fibers, with an
‘V’ configuration, which adapt progressively one into the
other, getting more and more opened ‘V’ shape, while going
to the middle regions of the ventricular wall. Mall also
describes two big muscular fascicles, ‘sino-spiral’ and
‘bulbo-spiral’, in their turn divided in superficial and
deep, but honestly underlying that he can not give any
Fig. 1. Five bovine hearts of similar sizes, representing the main successive
stages of the VMB dissection [Reproduced with permission of Ediciones Doyma
S.L. From: Torrent-Guasp F. La Mecanica Ventricular. Rev Lat Cardiol
2001;22(2):50].
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simple schema ‘which applies equally well to all portions of
the ventricular wall’ [23].
Later on, in 1956, Lev and Simkins [24] described three
different fascicles: one epicardial, other endocardial and a
third one, located at the middle of the ventricular wall. Lev,
according to Mall, claims that ‘the exact explanation for the
direction of the various fasciculi depends upon comprehen-
sive embryologic studies of the myocardium, which are still
to be carried out’.
Hort [25] has made, in 1957, an important contribution
with ‘micrometric evidence of shifting inter-digitations of
individual neighboring fibers’.
Daniel Denison Streeter, with his important works from
1966 to 1979 [26–29], undoubtedly deserves the attribute of
‘crossroad’ between the classical and the modern
approaches in studying the heart structure and function
relationships. With unique ability to ‘create a picture from
the puzzles’, he joined his ‘opened-up Japanese fan’
concept of the progressive change of direction in the
successive transmural planes of fibers, with reproducible
methodology described by Hort [25] and Torrent–Guasp [30]
into complete mathematical study and comprehensive
description of geodesic trajectories of the ventricular
myocardial fibers.
The result of anatomical studies, giving rise to the VMB
concept [1–5,30], provided that simple schema, about which
was claiming Mall [23], that applies equally well to all the
ventricular myocardial fibers, showing them joined together
in a coherent common general architectural plan. As
emphasized by Streeter, ‘accepting the statistical criterion
of the predominant fiber direction at given point’ [29],
earlier described by Torrent–Guasp [30], accommodates
factual difficulties, arising from complex and anisotropic
myocardial architectural design. Therefore, in the ventri-
cular myocardium, there are no anatomically reproducible
muscular fibers or distinctive layers, as thought by pre-
decessors [14–24] or erroneously interpreted by some
contemporary researchers [11,12,31,32], but only consistent
vectorial, functional trajectories and cleavage plans, which
could be very easily reproduced by previously described
dissection method [1–5,30] (Video 1—available only at on-
line version of this article).
The knowledge of that morphology, which configures a
helicoid with two spiral turns, has been the base that has led
to achieve the explanation of the way the heart performs
its double function, ejection and suction of blood [3,4],a
problem arisen 2300 years ago by the Greek physician
Erasistratus of Chios (304–250 BC).
3. Harmony of form and function
From the earliest days of anatomy [33] and physiology
[34], the form and the function were considered to be an
inseparable attributes of a whole, either form teleological
or from mechanistic (i.e. materialistic) points of view. It is
simply not possible to understand the essence
and the magnitude of natural harmony between form and
function, unless we accept the advice given by Sommer and
Johnson [35] that ‘relating structure to function in any
organ,.,leads inevitably to ontogeny and phylogeny’ and
unless we expand our intellectual scope from the molecular
to the organ level [6,36,37].
Recent brilliant review on cardiogenesis by Moorman and
Christoffels [38], integrates complex mechanisms, involved
in transition from a peristaltic tubular heart to a synchro-
nously contracting four-chambered organ. Development and
patterning of vertebrate heart is amenable to different,
evolutionary conserved and specific transcriptional networks
[39], as well as to the variety of epigenetic influences
[35–41]. New ‘ballooning model of chamber formation’
offers a logical explanation, from the genetic and functional
points of view, of the four-chambered heart design and the
organization of its electrical activity, thus overcoming the
mayor week points of previous ‘linear array’ concept, being
considered as ‘one of the most fatal assumptions’, regarding
morphology and flow direction [38].
Having on mind that evolution does not ‘conserve’
phenotype but genotype [39], it is easy to understand that
phylogenic memory does not necessarily produce morpho-
logical equivalents during ontogenesis, but rather appears in
a form of ‘fast-running’ genetic programs [36–40]. Accord-
ingly, Ernst Haeckel’s biogenetic law [42], stating that:
‘ontogeny is the short and rapid recapitulation of phylo-
geny’, does not necessarily mean that ancestral morphology
must appear in a recognizable manner during embryogen-
esis. Phylogenic and ontogenetic analyses and comparisons,
should take into account a fact, that patterning, rather than
structural appearance, is the only relevant and potentially
accessible information.
Specific spatial changes of the heart tube and surrounding
blood vessels are consequences of genetically programmed
proliferative and apoptotic events, as well as epigenetically
induced remodeling. Primordial cells originating from
primary and secondary heart fields, as well as those from
neural crest and pro-epicardium, all participate in a final
composition of the heart [35–41,43,44].
Epithelial-to-mesenchymal transformation of the endo-
thelial layer in a developing heart, is a nice example of
biologically and evolutionary ‘condensed genetic knowl-
edge’ [36–39,43,44]. Endothelial cells underwent such
transformation, start a series of mitotic divisions, filling
the acellular ‘cardiac jelly’ and producing the visible
protuberances denominated as endocardial cushions and
trabeculae. Similar process occurs on the other side of
cardiac jelly, in a compact myocardial layer, without
previous epithelial-to-mesenchymal transformation. Inter-
connections of those cellular bridges, surrounded by cardiac
jelly, define the final spiral patterning of the adult
ventricular myocardium (VMB). Trans-epicardial (i.e. pro-
epicardial) proliferation supports the myocardial mass by
vasculature, along with cardiac and non-cardiac (mesocar-
dial, neural crest) connective tissue, giving rise to extra-
cellular matrix, insulating tissues and anchoring fibrous
structures within the heart [35–41,43,43].
Filling the cardiac jelly with cells (i.e. ‘compaction of the
ventricular wall’) [43,44] is, maybe, the most important
mechanism, that governs the specific transmural spiral
patterning of the ventricular myocardial fibers, resulting in
formation of the double helical VMB. Accompanying visible
structural changes (genetically and epigenetically
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controlled), so often described as a distinctive phases of
embryological cardiac modeling, are probably nothing else,
but our ‘snap-shots’ of this continuing patterning process.
Looking back, into predecessors’ circulatory systems,
genetically, morphologically and functionally, we could not
resist to conclude that pumping organs (hearts) have been
developed later than tubular, vascular system. Moreover, it
is evident and emphasized in our previous reports [1–5,30],
that specialized pumping chambers have been developed
from the tubular vascular system. Shigei et al. [45], have
nicely reviewed phylogenic and ontogenetic changes within
tubular vascular system of invertebrates and vertebrates.
According to them, the appearance and development of
the ‘endothelium or endothelial tubular system’ and
‘sympathetic nerve/medial smooth muscle system’, were,
respectively, two most remarkable changes that have
occurred in the course of evolution of vertebrate circulation.
They hypothesize that the sympathetic nerve/medial
smooth muscle system may be regarded as a new neuro-
effector mechanism, developed for systemic regulation of
the endothelium-lined closed vascular system.
The complexity and difficulties in maintaining the
homeostasis, have urged for development of more sophisti-
cated control mechanisms [46,47], which may sometimes
overwhelm intrinsic functional capacities in circulatory
system. Accordingly, the heart and the tubular part of
circulatory system (i.e. arteries, capillaries, veins), now
being separated in pulmonary and systemic circulation, have
been submitted to numerous adjustments (e.g. complex
neural, humoral, rheological and other influences) during
phylogeny and ontogeny.
Structural features of each constitutive component of the
heart reflect both its electrophysiological and elasto-
mechanical performances, et vice versa.
Manasek [40] has shown that longitudinal re-arrangement
of developing myofibrils and appropriate re-shaping of
primitive cardiac myocytes, are induced by changes in
haemodynamic forces. Micro-design of a single cardiac
myocyte is, thus, amenable to fine structural adjustments,
in order to retain the optimal functional capacity in different
loading conditions. Gregorio [41] has proved that step-like
sarcomeral organization is necessary for synchronized
electro-mechanical action, which produces the uni-
directional blood flow in a hearts of all amniotic animals.
Accordingly, size, shape, connections and predominant
orientation (in a three-dimensional space) of a single-cell
sarcomeral protein molecules, determine their functional
behavior. The same is true for each individual myocardial
cell, myocardial fiber (being a series of longitudinally and
laterally connected cells), and myocardial laminar sheet
(Fig. 2)[3,6,29,40,41,48].
Microscopic and macroscopic myocardial architecture,
and particularly the existence of myocardial laminas or fiber
bundles, separable by distinct anatomic cleavage planes,
have been a controversial subject, since long ago [11,12].
LeGrice et al. [48], however, have provided acceptable
solution, documented by their comprehensive, detailed
measurements of canine ventricular myocardium. They
have reported that the cardiac muscular fibers are arranged
into distinct myocardial laminas, three to four myocytes
thick, separated from adjacent laminas by the extra cellular
collagen network. The myocytes are tightly coupled within
the same, but sparsely coupled between the adjacent
laminas. The planes of the laminas could be defined locally
by the longitudinal axis of comprising myocardial fibers and
by their spiral transmural direction on the ventricular mass
level.
Integrative knowledge of rule-based assignments [6,35–
38,40,41,43,44,48–50], on different organizational levels
within ventricular myocardium (Fig. 2), is essential pre-
requisite for understanding that blunt anatomical dissec-
tions, following predominant fiber direction [1–5,9,26–30],
are revealing their unique functional rather than eclectic
anatomical personality. In addition, understanding of this
principle would, hopefully, prevent erroneous comparisons
of the VMB with skeletal muscle model [11,12], and enable
those with individual difficulties to follow described VMB
blunt dissection protocol (Video 1—available only at on-line
version of this article), to adopt this easy-to-learn technique
(Fig. 3).
Since the mechanical result of the contraction of any
muscle (i.e. its laminar sheet, fiber, cell), always depends of
its disposition in the space—the direction defined by such
muscle, will have the greatest influence on its particular
action, when being activated. This interdependence of
myocardial form and function, described in VMB concept,
has been recently validated on intact, beating hearts, in
numerous investigations.
The first, indirect visualization of the VMB fiber trajec-
tories in intact bovine heart was done by Lunkenheimer et al.
[51]. Using a series of computerized tomography images,
obtained after trans-coronary produced pneumo-myocar-
dium, he was able to trace directly only the connective
tissue scaffold of the ventricular musculature, arranged in a
series of differently orientated spirals. Although this,
basically, double-contrast technique, was not able to
visualize myocardial muscular compartment (due to its
desiccation and partial fragmentation by the pressurized
Fig. 2. Left: Morphological substrates, at different organizational levels,
explaining the principle of predominant longitudinal action (arrow-lines)
within the heart: (A) molecules of sarcomeral proteins (thin and thick
myofilaments); (B) sarcomere; (C) cardiomyocytes and myocardial fibers; (D)
myocardial laminar sheets. Right: Myocardial tissue block (schematic) with
different possible dissection cleavage planes: (x and y) laminar trajectories,
(z) linear trajectory; each of them respecting the predominant principal fiber
direction at given point. Bottom: Unraveled VMB.
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interstitial air), it has left us a solid ‘negative image’ of its
spatial orientation, which, as the author said: ‘.confirms
and extends Torrent–Guasp’s double loop concept’ [51].
Recent advances in imaging techniques have provided
even more reliable methods for myocardial structural and
functional analyses. Material anisotropy of living tissue has
been mapped using diffusion-tensor magnetic resonance
imaging (DTMRI). It has been established that water diffusion
anisotropy, measured by MRI, faithfully parallels histological
anisotropy. DTMRI studies on intact, human beating heart,
have confirmed that myocardial fibers within the ventricular
mass, are arranged in layers of counter-wound helices,
encircling the ventricular cavities [52] (Fig. 4).
Displacement encoding stimulated echo (DENSE) MRI,
provides a high spatial density of displacement measure-
ments in the myocardium, via stimulated echoes, while the
image is always acquired at the same time point in the
cardiac cycle (Fig. 5). The spatial and temporal resolution of
the data acquired using this technique is superior to any
other imaging modality involving myocardial tagging and
tracking. In recent report, Saber et al. [53] explains that it
was possible to determine both myocardial fiber orientations
and ventricular wall dynamics in vivo, based on the VMB
concept, by applying DENSE-MRI technique.
Many lines of evidence suggest that this fiber configur-
ation serves to equalize stresses and strains across the thick-
walled ventricle, allowing both active and passive tissue
components to operate in optimal mechanical regimes
during the cardiac cycle [1,3,4,48–50,54–56].Horovitz
et al. [57] claimed that: ‘Anatomical findings as well as
theoretical considerations indicate that the myocardial
fibers lie along minimal length geodesics of the left
Fig. 3. Upper tray: A silicone rubber mould of the VMB (anterior and basal
aspect), designed and made for the educational purposes in 1990, by
Francisco Torrent-Guaspq. Lower tray: Schematic drawings of the VMB
silicone rubber model. (A and B) anterior aspect; (C and D) lateral aspect;
(1) right segment of the basal loop; (2) left segment of the basal loop;
(3) descending segment of the apical loop; (4) ascending segment of the apical
loop.
Fig. 4. Left: Schematic presentation of the successive CT scan planes,
obtained after trans-coronary produced pneumo-myocardium [51]. Right:
Drawing and the anatomical specimen (bovine heart), depicting the principal
counter-wound spiral orientation of the myocardial fibers in a transversal
plane, equidistant from the ventricular base and apex. For the comparisons
with CT images, please refer to Ref. [51].
Fig. 5. Left: Three-dimensional motion tracking of the left ventricle of the
human heart, during contraction (DENSE-MRI). The downward movement of
the base of the heart toward the apex and the twisting motion of the apex are
clearly visible [Reproduced with kind permission of Dr Han Wen, NHLBI
Laboratory of Cardiac Energetics. Original image available on Internet on
following URL: http/dir.nhlbi.nih.gov/labs/lce/Imaging-Physics.html]. Right:
VMB (bovine heart) with clearly visible fiber orientations, which are
concordant with trajectories traced by DENSE-MRI (left).
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ventricular wall.’ It is also a well known fact, that
myocardial cells are optimized to shorten along their long
axis. In a true physiologic situation, the radius of curvature,
sarcomeral length, fiber orientation, wall thickness, and
electrical activation vary widely with location within the
ventricular wall and within different time frames, during the
cardiac cycle [54–56,58,59]. Even so, the ventricles, being
considered as remarkable product of natural engineering,
have the unique ability to translate 15% linear sarcomeral
shortening into ejection fractions of greater than 50% and
wall thickening greater than 30% [54]. Such high efficiency
coefficient could not be explained without taking into
account a specific three-dimensional architectural plan,
explained by VMB concept [1–5,30]. Unraveling helicoid,
configured by VMB, allow us to distinguish four segments
whose respective fibers adopt different directions within left
ventricular mass. These four segments, when successively
activated during the cardiac cycle, can be further function-
ally individualized, through the different actions they per-
form during the contraction. Those actions, as explained in
our previous reports [1–5,30], provide the healthy heart with
strong capacities (defined by specific spatial arrangement of
the VMB segments) to ensure its own efficient emptying
(systolic function) and filling (diastolic function), by means
of successive muscular contraction along the VMB. Further-
more, the true understanding of the VMB biomechanics,
allows clear insight in the ventricular torsion mechanics and
systolic wall thickening, a phenomena clearly visible by
different imaging techniques, but hardly explainable by
classical knowledge of ventricular structure and function
[60–62].
Myocardial fiber architecture is also a key determinant of
both normal and pathological electrical properties of the
myocardium, strongly influencing initiation and spread of
the ventricular dysrhythmia [63–66].
Along with the mechanical aspects of the ventricular
performance, attention has been refocused again, on the
electrical events, giving rise to excitation–contraction
coupling along the VMB. Those events, and particularly
their anatomical and molecular substrates, still remain
controversial [1–5,56,67].
Examining electrophysiological and functional data, in
order to test the hypothesis of activation sequence within
the ventricular myocardium, James Cox has calculated the
delivery of the impulse through-out the VMB. He had
carefully plotted out how the electrical impulse was
delivered to the myocardium, by the specialized conduction
system, and correlated those findings with the velocities of
conduction in thin and thick areas of the heart. It came out
that impulse delivery precisely mimicked predicted
sequence of activation [67].
It has to be emphasized here, that patterns of
contractile activities within ventricular myocardium (i.e.
impulse deliveries), may or may not precisely correspond
with patterns of electrical excitations, as recorded by
microelectrodes or voltage sensitive dyes. This fact is
essential for the explanation of different results, obtained
by ventricular electrical mapping studies, and those that
are analyzing sequences contractile activities within ven-
tricular mass [63–66].
It has been clearly demonstrated by fast Fourier analyses
of ventricular MUGA scans, that muscular contractile
activity, during the cardiac cycle, progresses (in a peristal-
toid manner), along successive VMB segments [68]. Since the
excitation necessarily precedes contraction, the most
logical pattern of ventricular electrical activation should
follow both spatially and temporarily, previously described
sequence of its mechanical action.
Recent analyses of temporal shortening at the sites of
sono-micrometric crystals, implanted in selected regions
of intact animal heart, have validated proposed sequence of
contraction along the VMB [69]. Apart from initial encoura-
ging results [56,67,68], more studies are needed to confirm
that impulse delivery (i.e. excitation–contraction coupling),
is following the sequence of activation along the VMB.
Current anatomical and electrophysiological knowledge
neither fully accept nor deny this pattern of electro-
mechanical coupling within ventricular myocardium. The
most controversial topics are related to the organization and
function of, so-called, ‘specialized conduction system’
within heart. There have been numerous attempts to explain
the origin, development, organization and function of the
conduction system, both in health and in disease. It is
evident fact, from the pertinent literature, that cardiac
myocytes, conventionally distinguished as working myocar-
dium and specialized conducting system cells, share the
same embryologic origin [69,70]. Phylogenic and ontogen-
etic studies have shown the early appearance of the
unidirectional, slow peristaltic waves of contractions along
the heart tube, prior to any distinguishable molecular
or other phenotypic differentiation of existing myocytes
[69,70]. Patten concluded long ago, from his experimental
studies that: ‘the whole of the primary myocardium,
constituting the wall of the myocardial tube, was acting as
a conducting tissue.’ [71] Recent reports on fetal myocyte
transplantation, and consecutive reestablishment of myo-
cardial pacemaking activity, seems to be in accordance with
previous statement [72]. During the further developmental
stages, it appears that certain myocyte populations matu-
rate faster than the others, giving a rise to the anisotropy of
conduction velocities, which is, on the other hand, necessary
for coordinated activation of different segments within
developing heart. This maturation, among other aspects,
was shown to be related to specific ion channels, and
paradoxically, those cells that we call ‘specialized’ are in
fact less mature than cells belonging to the working
myocardium [37,38,40,69–71]. Pacemaker activity, while
present in embryonic ventricular myocytes is lost in adult
ventricular myocytes because it is normally held back or
repressed by the presence of inward-rectifier potassium
channels. Inhibiting those channels in working cardiac
myocyte, by adenoviral gene transfer, Miake et al. [72]
were able to produce a cells with higher spontaneous
depolarization rates.
There were many attempts to define some universal
conductive genotype and phenotype, but this task still
remains to be accomplished [37,38,40,69–71]. We find it
interesting to take into consideration some epigenetic
influences, which may play a certain role. Since the electrical
propagation through the developing heart (being basically
physical event) follows the pathway of the lowest resistance
F. Torrent-Guasp et al. / European Journal of Cardio-thoracic Surgery 27 (2005) 191–201196
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(according to Ohm’s law), than the cumulative electromotive
force, could be dependent on spatial organization of
myocardial cells. In other words, the structural patterning
of the developing heart, may have an influence on further
functional (and structural) differentiations toward slow and
fast conducting compartments.
4. From the laboratory to bedside
To close the circle, the new morpho-functional concept
of the VMB has to be widely integrated in clinical practice. In
its present extent, the VMB concept does offer solutions to
some ongoing clinical controversies.
During the past two decades, a new branch of cardiology
has been developed, on the basis of scientific efforts to
understand, define and treat diastolic heart failure [73–75].
As explained in our previous work [3], the concept of active
diastole has been offered a time ago as possible solution.
Until detailed explanation of VMB biomechanics was avail-
able, it was hard to understand the phenomenon of the
active ventricular filling. Contemporary experimental and
clinical investigations unequivocally support the attitude
that only powerful suction force, developed by the normal
ventricles, could be able to produce an efficient filling of
the ventricular cavities [3,73]. The most important part of
ventricular diastole (i.e. the rapid filling phase), during
which it receives more than 70% of the stroke volume,
belongs to the active muscular contraction of the VMB
ascendant segment [3]. Therefore, it is of utmost import-
ance for the clinical practice to realize, that normal
ventricular myocardium possesses strong intrinsic capacities
(due to specific spatial arrangement of the VMB segments),
to ensure its own efficient emptying (systolic function) and
filling (diastolic function), by means of successive muscular
contraction along the VMB [3]. By understanding this
principle, there would be no longer doubts, whether there
is: ‘Diastolic heart failure or heart failure caused by subtle
left ventricular systolic dysfunction? [76]. As Yip et al. [77]
concluded: ‘time for a redefinition’ has come.
We are all aware that hallmark of this century is an
increasing number of patients with hart failure. A lot of them
will need some kind of surgical treatment. So far, several
surgical ventricular restorative procedures [78–82] were
developed on the basis of VMB concept. Vaage has nicely
concluded in his recent editorial [10], that basicscience ‘helps
and promotes the development of cardiothoracic surgery as a
modern specialty with a dynamic and scientific profile’.
Therefore, instead of empirical, we should try to develop
new surgical procedures using more rational approach.
Trying to find an optimal non-transplant option, many
surgeons were trapped in purely mathematical evaluation of
their surgical strategies. These ‘Laplacian’ procedures
(although some of them being very elegant), are commonly
neglecting some essentially biological principles (e.g. tissue
composition, viability, inflammation), governing behavior of
normal and diseased hearts [83–85]. To develop more
physiological heart failure surgery—the first thing that we
have to realize is that the heart is not a ‘soap bubble’. In
another word—to deal with a heart failure—we have to
understand the function of the normal heart. To do so—we
have to understand its structure [1–5,7–9,54–57,86].
Apical loop of the VMB (Fig. 6) is the principal motor of
the heart [1–4,56,78–82]. Cutting through this loop, as
Batista proposes, not only reduces the left ventricular
volume, but impairs both its systolic and particularly
diastolic performance [83–85]. But if we preserve the apex,
as we have suggested long ago [78,79], and as Komeda et al.
[80] have done in large experimental animals, the mayor
disadvantages of the left ventricular reduction surgery could
be avoided. Understanding of the VMB concept is of critical
importance in performing a variety of restorative surgical
procedures, all of them addressed toward unfavorable left
ventricular remodeling [80–85].
We reasonably expect to see a lot of new clinical
implementations in the nearest future [86], as already
seen for congenital heart defects [87].
5. Important research directions
The VMB concept has emerged from more than 50 years
long research with more than 1000 anatomical dissections of
the hearts, belonging to different species [1–5,30,88].
Although the first publication with detailed anatomical
study has appeared more than 20 years ago [88], a serious
attention of the wider scientific community was not evident,
until NIH and NHLBI have organized the workshop: ‘Form and
Function: New Views on Disease and Therapy for the Heart’
(Bethesda, MD, 2002). Since then, this new anatomical and
functional concept has become seriously considered in many
researches [6,52,53,67,80–82,87], but still, there are many
questions to be answered.
The architectural plan of the ventricular myocardial
fibers, represented by VMB, which describes two spirals in
the space during its trajectory from the pulmonary artery to
the aorta, defining a helicoid which delimitates two ventri-
cular cavities, raises many interesting questions related to
phylogeny and ontogeny of the heart. In spite of perplexing
amount of information, obtained from contemporary
Fig. 6. Special dissection, depicting spatial orientation of the descending
segment (DS) and the ascending segment (AS) fibers, in the apical loop of the
VMB. The ‘window’ cut through the AS shows the septal crossing of AS and DS
fibers. Ao, aorta; PA, pulmonary artery.
F. Torrent-Guasp et al. / European Journal of Cardio-thoracic Surgery 27 (2005) 191–201 197
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molecular and genetic analyses [36–46], it remains unclear
the true origin of evident anatomical, histological and
functional differences between atria and ventricles of the
heart (Fig. 7), on the one side, and the apparent histological
and functional similarities between veins and atria, or
between arteries and ventricles, on the other side.
The form and the function of the heart (as well as of any
other structure) are inevitably interdependent. The same,
we believe, is true at each phylogenic and ontogenetic level.
Regarding previous, one of the most challenging questions is
related to the transition from the aquatic to the terrestrial
breathing. Precisely, the only pumping chamber in the
circulatory system of the fishes functionally belongs to its
venous part (bringing the venous blood to the gills). The
absence of pumping chamber in the arterial part of their
circulation is explained by low metabolic rates (low
resistance to blood flow) and the ‘whole body movements’
both being sufficient for adequate blood supply of the
systemic capillary network [38,39,89–93].
This example raises several concerns. First, is it really low
metabolic demand and whole body movement a consistent
explanation for the ‘un-pumped’ blood flow in the arterial
system of the fishes or, maybe, a kind of vestigial vascular
peristaltic waves may be the additional blood propelling
force? Second, concerning the function (pumping the
venous blood toward the gills), the entire heart of the fishes
is in fact the venous pump. Morphologically, it looks like a
heart tube of the higher, terrestrial breathing species (e.g.
birds, mammals) [38,39,46,89–93]. This similarity in mor-
phology, with obviously different (more complex) function
is confusing. Available explanations (more or less) [36–39,
46,92] could offer some insight into morphological trans-
formation of the serially connected chambers within heart
tube—into parallelly connected chambers of the final organ.
From the functional point of view—it remains unclear how
does the segmented organ which pumps exclusively venous
blood (i.e. the heart of the fishes)—acquires anatomical and
functional connection with the systemic, arterial circulation
in higher species, having in mind that it’s entire ‘arterial
pole’ (i.e. aortic sac) is actually deriving venous blood to the
gills? [38,39,46,89–96] Phylogenic and ontogenetic studies
and meta-analyses, performed along with VMB researches,
have opened completely new fields, now being conceptual-
ized in several ongoing research projects. One of them is
related to electrical activity of the heart and blood vessels
[5] and is supposed to analyze possible presence of the
vestigial vascular peristaltic activity.
It is widely accepted that electrical coupling between
myocardial cells is accomplished by cell-to-cell transmission,
via the gap junctions. Recent advances in understanding the
subtle mechanisms of spreading electrical activation, by
proton (H
C
) hopping between hydrogen bonded water
molecules have stressed the importance of the interstitial
tissue composition in that process [67]. Changes in the
electrical field, transmitted by Purkinje network, may alter
the ionization of water molecules bonded to the matrix
proteoglycans and this may, in turn, produce successive
proton hopping along those molecules. Such kind of mono-
valent cation mobility is remarkably faster than Na
C
or K
C
diffusion, responsible for cell-to-cell electrical conduction.
Having in mind again, that electrical activation precedes the
mechanical contraction, it has been observed that the
earliest contractions in human heart appear 35 ms after
initial excitation, and that most of the fibers are entering
contraction within 105 ms. Taking into account the 98% limit
of the QRS duration of 116 ms in normal human subject, and
the maximal possible velocity of the cell-to-cell conduction
of 0.3 mm/ms (along the longitudinal axes), it comes out that
maximal length of the unraveled VMB should not be greater
than 3.5 cm. In a reality, just as a comparison, the average
length of its ascending segment (in adult human heart) is
13 cm [30,88]. Therefore, proton hopping theory of electrical
conduction, may bring some new light on excitation–
contraction coupling at any given heart rate [67].
The importance of the mitral apparatus to left ventricular
function has been suggested in many clinical and exper-
imental studies of mitral valve replacement. True reasons
for better ventricular function with preserved ‘annulo-
papillary continuity’ still remain unclear [97,98]. Develop-
ment of new experimental research technologies has
allowed us to design an experimental protocol which would
(hopefully) prove our thesis about the role of the papillary
muscles and chordae tendineae within the mitral apparatus
and their influence on the ventricular function.
Finally, based on VMB anatomical and functional concept
[1–5,30,78–80,86,88] we have designed an external cardiac
supporting device (‘brace prosthesis’), which is meant to
preserve both systolic and diastolic functions of the heart
(Fig. 8) and prevent a series of events accompanying process
of ventricular remodeling [99,100]. This investigation is
currently in the experimental phase, and we hope that we
would be able to report some results in a nearest future.
6. Conclusions
‘No man-made structure is designed like a heart.
Considering the highly sophisticated engineering evidenced
in the heart, it is not surprising that our understanding of it
comes so slowly.’ [29]. The VMB concept, developed over
the last 50 years, has gathered many people willing to
contribute to the knowledge of heart structure and function.
Fig. 7. Bovine heart with completely separated atria and ventricles. Both atria
could be easily ‘lifted up’ from the ventricles without producing any
myocardial damage.
F. Torrent-Guasp et al. / European Journal of Cardio-thoracic Surgery 27 (2005) 191–201198
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All of them, led by the mysterious call of unknown, are still
aware that our mutual final goal is yet to be achieved.
Whether ‘the helix and the heart affect nature, the heart,
and the human’ [101], we really don’t know, but one thing is
evident—we are still together on the road towards new
understanding of the heart structure and function.
Acknowledgements
The first author dedicates this work to the memories of
his dear friends, Pedro Zarco Gutie
´rrez and Daniel Denison
Streeter, for their invaluable contributions to the VMB
concept, as well as for their support and understanding
throughout his life and career. God bless them.
Appendix. Supplementary material
Supplementary data associated with this article can be
found, in the online version, at doi:10.1016/j.ejcts.2004.11.
026
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DOI: 10.1016/j.ejcts.2004.11.026
2005;27:191-201 Eur J Cardiothorac Surg
Francesc Carreras-Costa, A. Flotats, Juan Cosin-Aguillar and Han Wen
Francisco Torrent-Guasp, Mladen J. Kocica, Antonio F. Corno, Masashi Komeda,
Towards new understanding of the heart structure and function
This information is current as of October 18, 2010
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