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Cardiovascular Research 48 (2000) 47

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Cardiac work and efficiency

N. Westerhof*
Laboratory for Physiology, Institute for Cardiovascular Research, ICaR-VU, Vrije Universiteit, Van der Boechorststraat 7, 1081 BT Amsterdam,
The Netherlands

Received 11 July 2000; accepted 13 July 2000

See article by Kiriazis and Gibbs [1] ( pages 111 119) two parts, activation and basal oxygen consumption. The
in this issue. first depends on excitationcontraction coupling and is
proportional to cardiac contractility. For the quiescent
muscle the PVA and the oxygen cost of activation both are
zero. The remaining part results from all other processes
1. Work output and efficiency of the heart requiring oxygen such as ion pumps. With increased

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cardiac contractility the oxygen cost of activation (Fig. 1,
1.1. Pressurevolume relation and oxygen consumption bottom) is increased in a proportional manner but the slope
of the relation between PVA and oxygen consumption is
Many researchers have tried to relate cardiac energy not affected.
metabolism in terms of oxygen consumption or heat The relation between PVA and VO 2 holds for all types of
production to mechanical performance. One now popular contractions, from isovolumic to isobaric (unloaded)
and accepted mechanical parameter is based on the pres- contractions. Isovolumic contractions correspond to the
surevolume relations found in the intact heart where largest PVA and thus the largest oxygen consumption [3].
pressure is plotted as a function of volume over the cardiac We see that external work is not a good measure of oxygen
cycle [2]. The curve that is followed during the heartbeat consumption.
consists of the following four parts (Fig. 1, top). Starting at Many other relations between oxygen consumption and
end-diastole (right bottom corner) ventricular pressure mechanical or hemodynamic parameters have been pro-
increases while volume remains constant (isovolumic posed. The two best known are the rate pressure product
contraction), during the ejection phase volume decreases (RPP) and the tension time index (TTI).
but pressure changes little, and, after closure of the aortic The RPP is the product of systolic pressure (Ps ) and
valves isovolumic relaxation ensues, which is followed by heart rate and is used as a measure of oxygen consumption
the filling phase. The area circumscribed is external in many biochemical studies. When these studies are
mechanical work per beat. The triangular area between the performed in isolated hearts in Langendorff perfusion, i.e.
systolic and diastolic pressurevolume relations and no external work is performed, the PVA(0.5Ps (Ved 2
bounded by isovolumic relaxation is potential energy. V0 )(0.5P 2s /Es , with Ved is end-diastolic volume, and V0
The sum of the two areas, shaded in Fig. 1 top, is called and Es are the volume axis intercept and the slope of the
the pressurevolume area (PVA). systolic the pressurevolume relation (Fig. 1, top). In other
It was shown that the oxygen consumption per beat words, for constant contractility (constant Es ) the Ps relates
(VO 2 ) is linearly related to the PVA (Fig. 1, bottom). To in a quadratic way to the PVA. With an increase in
derive oxygen consumption per minute, VO 2 may be contractility but constant filling (constant Ved ) the PVA
multiplied by heart rate. The intercept with the VO 2 axis is increases in a linear way with Ps but the intercept of
the unloaded oxygen consumption, found when the heart relation between PVA and oxygen consumption is altered.
is not allowed to develop pressure so that the PVA is Thus the RPP, as a measure of cardiac metabolism should
negligible. The unloaded oxygen consumption consists of be used with care.
The use of the TTI [4], the averaged pressure over the
*Tel.: 131-20-444-8111; fax: 131-20-444-8255. ejection period, is also limited because for an isovolumic
E-mail address: westerhof@physiol.med.vu.nl (N. Westerhof). contraction the TTI is negligible (ejection time negligible).

0008-6363 / 00 / $ see front matter 2000 Elsevier Science B.V. All rights reserved.
PII: S0008-6363( 00 )00176-0
N. Westerhof / Cardiovascular Research 48 (2000) 4 7 5

verted to heat. In terms of the above diagrams (Fig. 1,

bottom [3]) the myocardial conversion efficiency is defined
as PVA / total VO 2 . The contractile efficiency or myofibrillar
efficiency is defined the inverse of the slope of the VO 2
PVA relation. This contractile efficiency is in the order of
4050%. Economy of contraction is often used when no
muscle shortening takes place (presumably approached in
isovolumic beats) and is usually defined as the amount of
energy (oxygen consumption, ATP-usage or contraction
related heat rate) used for isovolumic contraction [6].
Work and efficiency depend on the load and the heart
[7]. Isovolumic (isometric) contraction, i.e. no ejection (no
shortening) and unloaded contractions, i.e. no pressure
(force) development, do not produce mechanical work and
efficiency is therefore also negligible. This means that
when plotting work or efficiency as a function of pressure
or cardiac output [7], at the extremes work and efficiency
are negligible and that in between a maximum exists. It
was shown that the heart under control resting conditions
pumps at these maxima [7].

1.3. Heat production

Heat production of the heart has been measured [5].

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Measurements show that heat is transported by convection
by the coronary circulation and by diffusion to thorax and
ventricular lumen in about equal amounts, and that convec-
tion transport increases with increasing perfusion. These
measurements agree with the observation that the heart is
about 25% efficient. However, accurate measurements are
extremely difficult to carry out.

Fig. 1. Top: diastolic and systolic pressure volume relations of the heart 2. Papillary muscle
and the force length relations of (linear) heart muscle for a single
contractile state. The intercept with the volume axis is V0 . The sum of Papillary muscle is used as a model of the whole heart
external work and potential energy, pressure volume area (PVA) or force with the main advantages that (blood) perfusion is not
length area (FLA) is a measure of cardiac oxygen consumption. Bottom:
relation between oxygen consumption or enthalpy with PVA or FLA. The
required and geometry is relatively simple. When a (com-
intercept with the vertical axis can be obtained by unloaded contraction plex) geometric model of the left ventricle is used, wall
(no pressure or force developed). The slope of the line is independent of stress (F, tension) can be related to ventricular pressure (P)
contractility. The unloaded oxygen consumption consists of two parts: and muscle length (L) to ventricular volume (V ). The PVA
activation related oxygen consumption, which is proportional to contrac- becomes force length area (FLA, see Fig. 1), which can be
tility, and a basal part, the oxygen consumption used for all other
related to oxygen consumption per contraction.
In the tradition of the measurement of heat in skeletal
muscle many studies have been performed on isolated
When the TTI is calculated as the pressuretime integral cardiac muscles using thermopiles [8,9]. The temperature
for isovolumic beats it is a measure of cardiac economy of the muscle is measured, and heat production derived.
(see below). Many aspects of the heat production and their meaning
have been published [8,10,11]. However, for the present
1.2. Efficiency and economy discussion it is sufficient to state that the measurement of
temperature and mechanical performance allows the calcu-
External (mechanical) efficiency is defined in analogy to lation of enthalpy, a measure of oxygen consumption
that of motors and equals the ratio of external work and [1,12].
oxygen consumption. Cardiac efficiency is normally about The model relating PVA and VO 2 , as suggested by Suga
2025% [5]; the remainder of the oxygen used is con- and coworkers [2,3], based on (isolated) blood perfused
6 N. Westerhof / Cardiovascular Research 48 (2000) 4 7

heart studies, did not seem to agree with the accepted performance and efficiency are studied in the isolated
concepts on (skeletal) muscle contraction. Fenns experi- cardiac muscle of aging rat. The basal metabolism (Fig. 1),
ments [13], on skeletal muscle, disproved the notion that measured in the quiescent muscle, was found to be
with each contraction, a predetermined amount of energy decreased with age. This could have resulted from fewer
was liberated either in the form of work or heat, or their membrane pumps in the somewhat hypertrophied
combination. This all or none phenomenon is not sup- myocytes where cellular areavolume ratio is decreased.
ported by the model relating FLA with VO 2 , because the Active metabolism (Fig. 1) was not different between the
PVA depends on the load. In other words the FLA concept muscles from young and old rats.
is not supporting the long discarded all or none behavior With senescence both shortening and force generation
of muscle energetics. The major argument that appeared to decreased, leading to a smaller external work. Also energy
disprove the concept of FLA, was also given by Fenn [13] consumption was decreased, but mechanical efficiency (in
who reported that, for skeletal muscle, a contraction with terms of work over enthalpy) remained the same. Contrac-
shortening (thus smaller FLA) generated more heat (used tile efficiency and economy did not change either with age.
more oxygen) than an isometric contraction (with larger Thus force and shortening change with age but all in-
FLA). However, this so-called Fenn effect is difficult to dicators of efficiency stay the same. The authors found
show in cardiac muscle, and is only found under very similar results in muscles of the rabbit with long-term
special experimental conditions [11]. volume overload [12].
Most studies on isolated cardiac muscle did not use the In guinea-pigs the effects of aging were found to be a
pattern of contraction the muscle encounters in the heart shift myosin heavy chains (MHC) from the alpha to the
and thus also did not allow for determination of the FLA. beta type resulting in a slower but more economical
This approach was started relatively late [14,15] but makes myocardium [18]. The difference may result from the
it possible to study if the findings in the heart are based on difference in experimental animal because in the rat mostly
cardiac muscle behavior. It took until 1990 [16] to show b-MHC is present. In pressure overloaded hearts [17] and
that the FLA of isolated heart muscle was a good measure in the transition to failure in the rat [12] efficiency and

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of relaxation heat (oxygen consumption). This was shown economy appear to be increased as well. Aging is thus in
also by Gibbs et al. [12]. In other words there appeared to terms of contraction speed not unlike the overloaded
be basic differences between skeletal muscle and heart condition but efficiency and economy seem to stay the
muscle in terms of the Fenn effect. It also implies that the same while in overload these parameters increase.
PVA approach to cardiac metabolism finds its basis in the It is possible to study energetics in human muscle strips
behavior of the cardiac muscle. from biopsies [10]. This approach gives the opportunity to
study patient groups in terms of mechanics and efficiency.
This type of study should be encouraged.
3. Cardiac aging
3.2. Clinical aspects
Because of the interaction of regulatory systems in the
intact human and animal, it is extremely difficult to obtain With the use of ultrasound techniques (Doppler and
a dependable measures of intrinsic contractile and ener- echo) much information on cardiac pump function can be
getic behavior. Therefore studies on isolated cardiac obtained. Also magnetic resonance imaging gives much
muscle, where conditions can be accurately controlled, are functional information. The mechanical parameters re-
required to obtain the effects of aging on cardiac muscle quired to obtain work, in combination with oxygen con-
mechanics and energy. sumption and the derivation of efficiency, are presently not
The changes taking place in the cardiovascular system part of the standard diagnostic procedures. This is in large
with age have been extensively discussed by Lakatta [17]. part because a series of contractions with different filling is
Although studies on the mechanics and efficiency of required to obtain at least some accuracy in the de-
cardiac muscle in normal and diseased conditions have termination of the PVA. Also oxygen consumption is rather
been performed [6,12], very little has been done with complex to measure in the human. The use of magnetic
respect to aging. It has been reported that with age the rate resonance spectroscopy and positron emission tomography
of contraction is decreased, electrical performance is may change this.
altered, changes take place in the isoenzymes of the
contractile apparatus and calcium handling is affected
[17,18]. References

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