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J. B-h, Vol. 23, No. 9, pp. 86S-881. 1990. m2 I 929oPl u.00 + .

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SURVEY ARTICLE

POWER EQUATIONS IN ENDURANCE SPORTS


G. J. VAN INGEN SCHENAU*~ and P. R. CAVANAGH:
lFaculty of Human Movement Sciences, Free University Amsterdam, The Netherlands; and
$Center for Locomotion Studies, The Pennsylvania State University. University Park. Pennsylvania,
PA 16802, U.S.A.

Abstract-This paper attempts to clarify the formulation of power equations applicable to a variety of
endur;una activities. An accurate accounting of the relationship between the metabolic power input and the
mechanical power output is still elusive, due to such issuesas storage and recovery of strain energy and the
differing energy costs of concentric and eccentric muscle actions. Nevertheless, an instantaneous approach is
presented which is based upon the application of conventional Newtonian mechanics to a rigid segment
model of the body, and does not contain assumptions regarding the exact nature ofsegmental interactions-
such as energy transfer, etc. The application of the equation to running. cycling, speed skating. swimming
and rowing is discussed and definitions of power, efficiency, and economy are presented.

NOMENCLATURE mass moment of inertia


m mass
A mean work per cycle M. moment about the ankle joint
a acceleration hf.. i reaction moment at joint a on proximal (2) or
a an angle distal segment (I) of the joint
component of acceleration along x-axis moment applied to a segment by the envir-
2s rate of increase of entropy onment
moment at joint j
As displacement of point of force application
dvldt acceleration of the athlete moment about the knee joint
E metabolic energy mass of segment &
d rate of expenditure of metabolic energy angular velocity of the ankle joint
EEl potential and kinetic energy of total body center angular velocity of a scgmcnt subject to a mo-
of gravity ment IV,
4 mechanical energy of foot (and skate) segment angular velocity of the foot segment
e# gross efliciency ( P,/ P,) angular velocity of joint j
4 energy degraded to heat angular velocity of the knee joint
Ek translational kinetic energy angular velocity of the shank segment
E, mechanical energy of lower leg segment angular velocity of the thigh segment
E, metubolic energy liberated power
em muscle efficiency power output of a system including subject and
potential energy equipment
EP
propelling efficiency conservative power output component (used to
eP
E, rotational kinetic energy do work against conservative forces)
4 strain energy power dissipated to the environment
=L, sum of segmental energies power associated with overcoming gravity
a force or force component power input (the usable fraction of H)
cychc frquency (I / To) power dissipated in giving kinetic energy to the
i reaction force at joint a on proximal (2) or distal water in swimming
a.1
segment (I) of the joint p, power used for maintenance activities
F.ir net force due to air resistance P “S non-conservative power output component (de-
F .,l. 1 force of air friction at the center of mass of graded to heat)
segment i power output (mechanical muscle power)’
FO drag force F
T'
mean generated power
FL lift fora temperature (K)
F* external force time
F, frictional force time interval for integration
FLII force of ice friction cycle time for cyclic movements
propulsive fora velocity of the athlete as a function of time
FP velocity of the air relative IO the subject
F, force component along z-axis
acceleration due to gravity velocity of antcr of mass of segment c
6
H chemical power (rate of decrease of enthalpy of velocity of the point ofapplication of an external
roodstutT) force F,
VI velocity of point i
% velocity of the point of force application
velocity component along x-axis
Received in jTnp/ form 24 November 1989. velocity component along z-axis
7 Reprint requests to: G. J. van lngen Schenau, Faculty of % work done on a body
Human Movemqm Sciences,Dept. Functional Anatomy, v.d. zr projection on z-axis of height of point c above a
Boechontstraat 9, 1081 BT Amsterdam, The Netherlands. datum
866 Survey Article

1. INTRODUCTION survey will mainly be focused on human cycling,


rowing, running, speed-skating and swimming. Since
In studies of the mechanics and encrgetics of human this survey might be of interest for a broad audience,
and animal locomotion, the metabolic energy liber- including exercise physiologists and zoologists, only
ated (E) is often regarded as the main input variable of Newtonian mechanics are used and (planar) examples
the moving system. Speed u and its time derivative are given in order to clearly illustrate mechanical
(acceleration, du/dt), on the other hand, are seen as the concepts which otherwise may be difficult to concep-
final results of a process where equations of mechan- tualize.
ical power are used in attempts to deduce causal
relationships between & and u. The approaches which
are most frequently used in modelling mechanical 2. SUMMARY OF THE LITERATURE AND STATEMENT OF

power can roughly be divided into two categories, a THE PROBLEM

kinematic approach and a kinetic approach.


In the first method, calculations of power from 2.1. Energy liberation and perjxmance
changes of mechanical energy of body links or of the Many studies show a correlation between the lib-
center of mass of the moving system are made. This eration of metabolic power 2 and mean speed as the
approach is mainly used in studies on running and main performance determining factor in endurance
walking. sports (Zatsiorsky et a/., 1982). Such correlations were
The kinetic approach involves calculations of power often reported for running (Farrell et al.. 1979;
based on a summation of the components of power Holmer, 1978; Di Prampero, 1986; Daniels, 1985) but
used to overcome frictional losses in work against the also for swimming (Charbonnier et al., 1975; Holmer,
environment. This approach is mainly used in studies 1974; Di Prampero, 1986), cycling (Pugh, 1974; Di
of terrestrial or aquatic locomotion other than Prampero, 1986) and speed-skating (Holmer, 1978; Di
running or walking. Prampero, 1986; de Groot et al., 1988). These rela-
Though a number of studies pay attention to both tions are likely to have a solid exercise physiological
types of power. most studies seem to be based on basis, since humans and animals appear to be able to
ambiguous analysis of possible power dissipation. As deliver a certain amount of mean metabolic power
noted by Aleshinsky (1986a). studies do not seem to which (unlike engines) is strongly dependent on the
exist where the power calculations for the entire duration of the exercise. 6 further depends upon, e.g.,
athlete are based on the definition of power, as can be the amount of muscle mass involved (Tesch et al.,
found in textbooks on mechanics. For separate body 1976; de Groot et al., 1988). technique, environ-
segments, power equations have been deduced by mental and body temperature, and psychological fac-
Quanbury et al. (1975) and applied by, e.g., Winter and tors (see Astrand and Rodahl, 1977; Cavanagh and
Robertson (1978). Kram. 1985b; Morgan, 1985). Inter-individual differ-
The first purpose of this paper is to present a general ences in % are associated with factors like natural
power equation for the assembly of body segments ability and training background (Astrand and Rodahl,
applicable to all endurance sports. 1977).
The fact that most power calculations are based on Synthesizing the approach of various previous in-
a more or less arbitrary summation of the components vestigators (e.g., Tucker, 1975; Ward-Smith, 1985a).
ofpower, rather than on the application of mechanical Fig. 1 shows a schematic representation of the flow of
equations to an assembly of explicitly defined free energy during exercise. The chemical power ri repre-
body diagrams, leads to an overwhelming number of sents the rate of decrease of enthalpy of the original
definitions of external and internal power, and of foodstuff. According to the second law of thermo-
quotients between power and i usually referred to as dynamics, only a part (the rate of change of the ‘free
‘efficiency’. energy’) of this power can be used for biological
In running and walking, the problems associated functions (Astrand and Rodahl. 1977; Whipp and
with obtaining accurate definitions of power output Wasserman, 1969; Gibbs and Gibson, 1972; Woledge,
render the concept of ‘efficiency’ almost meaningless. 197 I). This useful component of power will henceforth
In other activities-such as rowing or cycling-where be called ‘power input’ P, (Tucker, 1975; Daniel, 1983)
power output against an external resistance can be and is essentially equal to the metabolic rate ri men-
determined with more precision, a useful-if generally tioned above. The other part is equal to the product of
underestimated-measure of efficiency can be deter- the rate of increase of entropy AS and temperature T
mined. There is, however, a need for unambiguous (K). In gross body movements, only a part of the
definitions and the second purpose of this survey is to power input is converted into mechanical muscle
provide such definitions. An additional purpose is to power P, which in principle might be used to perform
review the various methods presently used in the movements and to do work against the environment.
calculation of power. These methods are discussed The rest, the ‘maintenance power’ (Tucker, 1975).
first. concerns mainly the work-rate oCmuscles which oper-
Though a number of examples will be deduced from ate the card&respiratory system, and power used in
or applied to other activities and other animals, this tissues that do no (‘useful’) mechanical work (e.g., the
SurveyArticle 867

technique. or body composition on speed are mainly


II power
based on equations of P,. Examples of such predic-
/ tions are the calculation of the influence of shielding
il (Kyle, 1979; van Ingen Schenau, 1982). technique
\ - \ (Sanderson and Martindale. 1986; Boer et ol., 1988)
Pi 1 and altitude (Di Prampero et al.. 1979; Di Prampero,
maintenance ’ pm \ 1986; van Ingen Schenau, 1982) on speed. Though
power . 1 entropy good agreement with actual performances can be
L \ Prpln t L obtained, these models of course have their limita-
tions, since variables such as P,,. P, and i may be
influenced by environmental conditions and tech-
nique as well. This dependency may also obscure the
significance of changes in efficiency calculated on the
basis of P, and Pi.
P,
non- ’ Pnc I 2.2. Mechanical power calculations
conservative II P, conservative
power ’ power 2.2.1. Running. Since Fenn (193Oa, b), many re-
I, \ \ searchers have tried to deduce equations which might
describe P, and P,,, and their interaction for walking
and running. The calculation of this power (which is
mainly associated with the need for accelerating,
lifting and decelerating body segments) is largely
based on the following methods (Zatsiorsky et al..
1982):
(a) Calculations based on changes of segmental
energy of all body segments (eg., Winter, 1978;
Norman et al., 1976; Pierrynowski er ~1.. 1981:
Alexander, 1980: Cavagna and Kaneko, 1977) or of
the center of mass (e.g., Cavagna ct al.. 1976).
(b) Calculations based on summation of joint
powers dclincd as the product of joint moments and
acceleration their angular velocities (e.g., Winter, 1983).
(c) A combination of these methods (Aleshinsky.
Fig. I. The main flows of power in locomotion.
1986a. b: Quanbury et al., 1975; Robertson and
Winter, 1980).
In studies of running and walking, P, is usually
digestive tract, neurons, muscles which stabilise the considered to represent such a small fraction of P, that
trunk). From the muscle power PO, henceforth called it is frequently neglected. Physiological studies (Pugh,
‘power output’ (Tucker. 1975) a part P,, is degraded 1971) and, more recently, wind tunnel studies (Kyle,
into heat by non-conservative frictional forces inside 1979) have shown that P,, even in distance running,
the body or when musclesare doing work against each can be at least 10% of f,,. Moreover, Webb ef ul.
other (Alexander, 1980). A second part P, is used to do (1988) showed that the power associated with the
work against conservative forces(e.g., gravity, stretch- deformation of the shoe can be even larger than 10%
ing elastic elements) and can in principle be re-used as of P,.
P,, or P. (see pointing arrows) in a later phase of the However if P, is neglected, it will be clear that the
movement. P, represents power dissipated to the three methods of power calculation lead to a zero
environment (excluding power expended against mean power output in level running, if the cycle
gravity). averaged value is calculated by a simple time integral
In running, most studies on mechanical power of the instantaneous power. Since the instantaneous
concern the modelling of P,, and P,, while in the other power P,+ P,, is not zero, and since the delivered
types of locomotion, the majority of studies is aimed at positive and negative mechanical powers within a cycle
modelling P,. A few attempts have been made to are clearly associated with a certain P,, many different
model a direct relation between !? and speed u(t) in assumptions are proposed in order to get an estimate
running (WardSmith, 198Sa; Morton, 1985) in order, for mean mechanical power in walking and running.
for example. to’ predict the influence of wind on speed These assumptions concern questions about how
(Ward-Smith, il985b). As discussed by van lngen much energy might be transferred within and between
Schenau and Mollander (1987) and by Morton (1986). segments [not necessary in method (b)], how much
these models at present do not adequately describe the energy can be re-utilised from elastic structures where
involved physiological processes. Other attempts to it was stored in an earlier phase of the cycle, and
predict such influences as environmental conditions, estimates for the mechanical efficiency of the conver-
868 Survey Article

sion of Pi into P, and P,,, for both positive as well as that this product is equal to the power needed to
for negative power. It is likely that almost all present overcome drag. van lngen Schenau et al. (1985) and
methods overestimate the work done, and thus calcu- Fukunaga et al. (1981) associate this power with the
late ‘efficiencies’ which are unrealistically high. Since increase of mechanical energy of the athlete’s center of
the methods and their problems are already extens- mass. None of these studies (nor the many studies on
ively discussedin other papers (Cavagna and Kaneko, jumping where the same method is used) provide
1977; Williams and Cavanagh. 1983; Williams. 1985; convincing evidence for the applicability and validity
Cavanagh and Kram, 1985a.b; Aleshinsky. 1986a-e). of this product, which is not based on the definition of
the reader is referred to these papers and their referen- power since the velocity of the point of application of
ces;to prevent too much repetition, it is assumed in the the propelling force is not equal to the velocity of the
remaining part of this paper that the reader is familiar center of mass. In running (Fukunaga et al.. 1981) and
with the content of those papers. It should be men- jumping, the velocity of the point of application of this
tioned here that Aleshinsky (1986a) was the first force is actually zero. The significance of this approach
author to present an analysis for the entire athlete will be discussed below.
based on a systematic and unambiguous application From the above it is apparent that at present a
of basic mechanics. However, in his application of this number of different methods are being used to calcu-
method to running, he met with similar problems to late different types of power. It is not clear however
those described above for other investigators. how these various powers are related.
2.2.2. Ocher endurance sports. As stated above,
power measures for endurance sports other than
running are mainly based on analysis of possible flows 3. PROBLEMS IN APPLYING CLASSICAL MECHANICS
of energy to the environment. Di Prampero can be
regarded as an important pioneer in this field. (Di Since, apart from Aleshinsky, all referenced authors
Prampero et al.. 1971, 1974, 1976, 1979). He and most construct power equations for the entire athlete by
other authors (e.g.. van lngen Schenau. 1982, 1988) adding up possible flows of energy, the question arises
simply take the (scalar) product of opposing forces to what extent classical mechanics provides adequate
(mainly frictional forces with the environment) and the tools for an approach based on mechanical laws and
velocity of the athlete and his equipment (bicycle. definitions.
boat), which is taken as equal to the velocity of the When applying Newton’s second law ZF=ma, the
point of application of these forces. Though it may be result is unambiguous: after one has constructed a free
true that the majority of P, is used to overcome body diagram, the internal and external forces are
friction, and though this power calculation is based defined by the boundaries of the diagram, and the law
upon a reasonable application of the product of rorce provides a unique relation between the external forces
and velocity, this modelling of P, has nothing to do on the one hand and the acceleration of the center of
with a general and unambiguous application of class- mass of the system on the other hand. This is also true
ical mechanics on the entire system as advocated (and for a system which is deformed under the influence of
applied to running) by Aleshinsky (1986a-e). If the these external forces, with or without energy dissipa-
entire system of athlete and equipment (e.g., bike and tion by internal forces. Internal forces do not play any
rider or rower and boat) is taken as the free body for role in this force equation. The same is true if the time
the purposes of analysis (which implicitly is the case integral of this force equation is taken. The
in these referenced studies), all external forces (defined (impulse-momentum) equation is also uniquely de-
by the choice of the free body) should be accounted fined. In both equations, gravity does not play a
for, including the propulsive force(s). Moreover the specific role but is simply one of the external forces
question may arise how to define P, as power caus- which should be accounted for.
atively related to the metabolic rate P,. If for example Since power can be defined as a flow of energy, one
a skater suddenly stops pushing off, he can still deliver might expect a comparable relationship between the
P, for a certain time interval, but it is clear that P, has flow of energy associated with the external forces and
no relation with P, at the same time (so does the skater the rate of change of energy of the chosen free body.
deliver external power or not?). Of course we know Such a power equation or its time integral called
that in such a case the skater can overcome friction at ‘work-energy theorem’(Robertson and Winter, 1980).
the expense of his kinetic energy, but how should this ‘energy balance’ (Aleshinsky, 1986b). or ‘work-energy
power be defined? This example shows that here also is equation’ (Meriam, 1975). however, cannot be con-
a need for an unambiguous approach in order to structed by the same unambiguous methodology as is
deduce proper definitions for power. the case for a force or a force-time integral equation.
2.2.3. Other approaches. A number of authors have This has simply to do with the fact that the human
used the product of the propulsive force and the body can liberate metabolic energy. degrade mechan-
velocity of the center of mass of the system in order to ical energy into heat, and store energy in elastic
get a measure for mechanical power (Fukunaga et al., structures.
1981; Sanderson and Martindale. 1986; Tucker, 1975; If we wish to link information concerning the force
van Ingen Schenau et al., 1985). Tucker (1975) states environment in a given biological problem with these
Survey Article 869

factors, then the problem must be approached using a The evaluation of P in terms of the forces acting
work-energy principle which states that the work requires the evaluation of the work done by the force
done on a body is equal to the change in all these during a displacement of its point of application as
energies: follows:

W= AE, + AE, + AE, + AE, + AE, + AE, (I) W=F.As

where E, is potential energy, E, is translational kinetic where As is a displacement of the pint of application.
energy, E, is, rotational kinetic energy, E, is strain When evaluated in the time interval At as AtdO. the
energy, E, is energy degraded to heat, E, is liberated expression for power becomes:
metabolic en$rgy. If the time interval At is considered
P=F*v;.
as At-O. the expression for power becomes:
In the present example there are two forces acting on
P=dE,fdt+dE,Jdr+dE,/dt+dE,jdt
the body, the force F, and gravity.
+dE,/dt+dE,/dt. (2) The power associated with the external force F, is

If the athlete is choosen as the free body, it will be clear P,=F;v,


that at present there are no reliable models available
and the power associated with the force of gravity
to calculate ES and E,. This means that an approach
where equation (2) is applied to the entire athlete is not P,= -mg*v,
(yet) possible.
with the minus sign indicating that the force and the
For a rigid body, however, the right-hand side of
displacement are in opposite directions.
equation (2) can be fairly easily stated using the
It would now be tempting to consider the total
conventional expressions for potential and kinetic
power as P,+ Pz and substitute this in the left-hand
energies, sinae the last three terms are zero. The
side of equation (2). leading to the anomalous result:
formulation of the left-hand side of the equation in
terms of the forces acting. however, needs care in a v,. (F, -mg)=mg*v,.
gravitational cnvironmcnt, since the effect of gravity
This result is false since we know from Newton’s
must bc included either on the ‘work’ side of the
second law that F I = mg and that the rate of change of
equation or the ‘cncrgy’ side of the equation, but not
potential energy. mg*vz, is not zero as the equation
both (Mcriam, 1975).
would predict.
To illustralc this, consider a passive rigid body
Change in potential energy is simply the work done
shown in Fig. 2 moving under the influence of the
by or against gravity. and thus one or the other-but
external force F, in pure translation with a constant
not both-should be included in the work-energy
velocity u, in the vertical direction. Its position above
relationship.
an arbitrary datum at time t is L. The only other
Thus the correct formulation of equation (2) for this
external force acting is the force of gravity, mg.
illustrative example would be
Using equation (2) for this problem yields
F;v,=mg*v,
d(E,)/dt =d(mgz)/dt =mgu, (3)
which, since F,=mg. is true. In the remainder of this
since E, is constant, E, is set lo zero and E, is zero for
paper, we shall choose to include the effects of gravity
this rigid body.
as an energy term on the right-hand side of the
work-energy equation.
One final principle needs lo be presented before
examining the application of power equations to
various problems in human movement. Clearly all
external forces, other than gravity, should be included
in the development of an adequate model. In endur-
ance sports where forces such as friction provide the
principal resistance to motion (including drag from air
or water, roiling friction, etc.) there are no serious
problems. Power associated with these forces can be
calculated from the product of the forces and the
velocities of their points of application. But the pro-
pelling force is somewhat more obscure.
In the simple passive system shown in Fig. 2, be-
cause the environment adds power to the system, the
flow of energy could be derived according to the
Fig. 2. If a body is moved under the influence of an external
definition of power. The energy of a passive system
force F,, one shbuld include the eRect of gravity either as a
(potential) energ(yterm or as a work term in the formulation (such as a discus or shot) can clearly only be increased
of a work-dnergy equation for the body (see text). at the expense of a source external lo the system (such
810 Survey Article

as the thrower). lf we exclude paired activities (such as energy but to a phenomenon usually referred to as
tandem cycling or crew) and environmental factors muscle potentiation. Since Hill-type models (Hill,
such as a strong back wind and compliant floors, such 1938) do not account for this potentiation, more
a situation does not exist in most locomotor activities. reliable results might be expected in the future from
All mechanical power is delivered by the transforma- Huxley-type models (Huxley, 1957; Zahalac, 1981,
tion of metabolic power into mechanical power 1986). Other necessary information concerns the (in-
(Zarrugh 198 1).The following definition of locomotor stantaneous) total muscle lengths and the contractile
activities from the perspective of power therefore properties of the individual muscles (Hof et al., 1983;
seems to hold true: Bobbert et 01.. 1986; Whiting et al., 1984). For most
muscles this information is not yet available.
In locomotor activity there is no external force
All these (and other) uncertainties render the formu-
that adds positive power to the system.
lation of a generalised instantaneous power equation
Stated formally, the propulsive force F, will always for locomotion based on equation (I) impossible for
obey the relationship: this time.

F;v,<O (4) 4.1. A model consisting of rigid links


where vp is the velocity of the point of application of Apart from movements involving the shoulder joint
F,. As will be shown later, this relationship needs (which allows considerable translation), human move-
further discussion when power equations are applied ments can to a large extent be described as the sum of
to different types of locomotion. rotations performed at joints between the body seg-
The following conclusions may be drawn from the ments. This means that the part of the work liberated
above: in muscular action which is used for limb movement
(I) For an endurance athlete as a system (including and to do work against the environmental forces is
equipment). a power equation based on the rate of predominantly being reflected by thesejoint rotations.
change of mechanical energy of the system and the Many researchershave applied the method of analysis
powers associated with the displacement of external as proposed by Elftman (1939a.b) in order to assess
forces should include internal power as dissipated into the power associated with thesejoint rotations. In this
heat, stored in elastic structures and generated by the method the body is modellcd to consist of an intcrcon-
contractile elements of the muscles. netted chain of rigid links. The mechanical intlucnce of
(2) For rigid bodies, equation (2) can unambigu- segment i on the adjacent segment i+ I is calculated
ously be applied provided that the special character of using inverse dynamics. In planar motions all these
gravity is taken into account (Meriam. 1975). Such influences are summarised in a hypothetical reaction
equations have previously been deduced from and force F,:, and moment M,,,. The analysis usually
applied to individual human limb segments by, e.g., begins wtth a distal segment where the external forces
Quanbury et al. (1975) and Winter and Robertson and moments are known (e.g., ground reaction forces)
(1978). and all intersegmental forces and moments are sub-
sequently calculated (Winter, 1983; Zatsiorsky et ul.,
J. AN INSTANTANEOUS POWER EQUATION 1982; Ericson et al., 1986). Note that rigid links do not
generate power and do not degrade power into heat.
A power equation which includes the rate of change This means that power is supplied to or absorbed from
of all energies mentioned in equation (1) is difficult to a segment by the hypothetical intersegmental forces
construct (Norman and Komi, 1987). At the present and moments, and by environmental forces such as air
state of knowledge there are no models available resistance (Winter and Robertson, 1978; Robertson
which predict reliable measures for positive and neg- and Winter, 1980). As discussed above, equation (2) is
ative power from active elements of individual muscles applicable to each rigid segment. For running, Robert-
and their efficiencies. This is not only due to the son and Winter (1980) showed that the flows of energy
indeterminacy problem (Miller, 1979) since even the associated with the external forces and moments of a
application of direct dynamics (e.g., Hatze, 1977.1978) segment are indeed in balance with the rate of change
suffersfrom severe uncertainties. Apart from problems of the mechanical energy of the segment. The only
in evaluating the reliability of proposed cost functions, exceptions were found for the foot during foot strike
this is, e.g., a result of the lack of information about the and push-off where significant deformations of the
compliance of the series elastic elements of the muscle foot occur.
tendon complexes involved. As can be concluded from As advocated by Zarrugh (1981). a power analysis
the work of Cavagna er al. (1968,1985). Alexander and for the entire athlete should include both the rate of
Bennet-Clarke (1977). Morgan (1977). Morgan et al. change of segmental energies and the joint powers. In
(1978). Walmsley and Proske (1981). Jaric et al. (1985) his original contributions to this journal, Aleshinsky
and van Ingen Schenau (1984). the capacity for (1986a.b) was the first to succeed in deducing such a
muscles and tendons to store elastic energy is often power equation showing an unambiguous relation
overestimated, while the effects of pre-stretch are, in between the rate of change of segmental energies on
part, not due to storage and re-utilisation of elastic the one hand and the sum of power from environ-
Survey Article 871

mental sources and joint power on the other hand. application and M,., by the angular velocity w, of the
Aleshinsky’s deduction was based on a summation of foot. E, is the mechanical energy of foot and skate:
the power equations constructed for each rigid link.
E,=$m,vf+m,g:,+~lwf (6)
These power equations were derived by differentiation
of the segmental energy of the links (Aleshinsky, with m, the mass of foot and skate, v, the velocity of its
1986a. b). To avoid the complicated mathematics ne- center of gravity, rC the vertical position of this
cessary in that approach, we take here the left-hand segmental center of gravity and 1 the moment of
side of equation (2) as a starting point. Figure 3 shows inertia of foot and skate relative to the center of
an example of three links with the external forces and gravity. For the lower leg the power equation is:
moments per link (except gravity). The example con-
M..,%+F,.,*v,+F ,,r.z*v4+Ft.,-vs+Mt.t~,
cerns the leg of a skater but the deduction would
essentially be the same if it concerned a skier, cyclist or =dE,/dt (7)
a runner. where the symbols used are comparable to those of the
According to equation (2). the power equation for foot. These equations can be constructed for all other
the foot is as follows: rigid links although not all external forces and mo-
ments per link have to be known. This may be the case.
for example, in cycling where the trunk is supported
=dE,/dt (5) by the saddle and where a reaction force and moment
where F, is the reaction force from the ice, F,,, and F,,, can act on the hands. When compared to the work
the ice friction and air friction forces respectively. F,., presented by Quanbury et al. (1975). Winter and
and Al,,, the’joint force and moment. The forces are Robertson (1978). and Robertson and Winter (1980)
multiplied by the velocities of the points of force the new step made here is that we now add the

POWER SOURCES PER SEGMENT

thigh

foot + skate

Fig. 3. II the body of a skater is modelled to consist of interconnected rigid links, :he sources of power per
link are the external moments and forces (except gravity) per link.
872 Survey Article

equations for the separate links in order to obtain one be judged as a firm mechanical basis for existing
equation for the entire athlete. If one realises that approaches. Equation (9) does not solve most of the
F,.1-v~+F..z-v>==0 but that M..,w( + M.+. problems that were previously described for running.
= M,o,, it is clear that a summation of all these power This is due to the (hypothetical) nature of the joint
equations yields the following power equation for the moments and joint power. Sincejoint moments are net
system: moments (sum of all contributing structures), the
origin of positive power and the ultimate location of
Fp~~p+F,,,~up+F,i,~~.i,+~M,o,=d~~,,~ldt (8)
destination of negative joint power cannot be assessed.
where M, and oj are joint moment and angular Positive power can originate from active as well as
velocity per joint respectively. F,,, * v,~,. which repres- elastic elements of active muscles. Negative joint
ents the total power lost to air-friction, is approx- power can reflect power which is degraded into heat
imately equal to the main air-friction force averaged by friction in joints or other passive structures or by
over the segments times the mean velocity of the eccentric actions of active muscles, but it can also be
segments with respect to the air (uair),and ZE,,, is the stored in elastic structures and later be re-utilised as
sum of the segmental energies. If, as already discussed positive joint power. Positive joint power does not
above, Z M,u, is to be judged as the origin of power in account for power which is simultaneously degraded
locomotion, all the other expressions can be judged as into heat by friction or when agonists do work against
locations of power dissipation. To express the causal their antagonists which cross the same joint. Bi-
relation for this and other locomotor activities, all articular muscles may distribute power over the adja-
powers associated with external forces F, are sum- cent joints which are crossed by these muscles. As
marized and placed on the right-hand side, leading to convincingly shown previously (van lngen Schenau et
the general instantaneous power equation: al., 1987; Bobbert and van Ingen Schenau, 1988; van
Ingen Schenau, 1989) one cannot simply assign posit-
ZM,u,=dZE,,,Jdt-ZF;v,. (9) ive or negative joint power to the actions of muscles
For the most general case, this equation can be which cross a joint. For example, the bi-articular
extended with an expression X Mp, reflecting power gastrocnemius can oppose the knee extensors in such a
associated with environmental moments which might way that no, or even negative, power is generated in
act on a segment which has an angular velocity w, the knee joint, since the net moment is zero (or
(Aleshinsky. 19861, b). Such contributions however negative). In reality, the knee extensors may lift the
are scarce since a pure moment requires a combina- calcaneus via the gastrocnemius muscle (Gregoire et
tion of pushing and pulling forces.This might occur in, al., 1984; van Ingen Schenau ef al., 1987; Bobbert et al.,
e.g., the hands in cycling. However in that situation, w, 1986). In other words, the joint power that appears
will be negligible. It should be emphasised that in this computationally to have its source at the ankle can, to
power equation the contributions of the hypothetical a large extent, be liberated in the knee extensor
joint forces are cancelled out. The meaning of this is muscles. Since the extremities contain many import-
that thesejoint forcescannot generate power; they can ant poly-articular muscles, such a redistribution of
only redistribute power among the segments (see power over the joints will often occur. This does not
Winter and Robertson (1978) and Robertson and mean that bi-articular musclescause erroneous results
Winter (1980) for further discussion of this power when calculating the powers expressed by equa-
concept). Although a different deduction is used here, tion(9). Bi-articular muscles can cause an inter-
equation (9) is essentially equal to the power equation compensation of joint powers which, however, does
deduced by Aleshinsky (see equation(20) in not affect the summed joint power. Errors in the
Aleshinsky, 1986b). calculation of the joint powers will mainly be caused
As stated before. each contribution F;v, from the by the necessary assumptions concerning the rigidity
environment should be negative or zero. This means of the links and the position of the (moving) axes of
that the expression -IF, * v, can also be defined as rotation.
the sum of positive powers which flow to the environ- It should be emphasized here that the redistribution
ment. Expressed in words, equation (9) says that, at of joint powers by action of poly-articular muscles
each instant, the summed joint power is equal to the reflects a process which is previously defined as a
rate of change of the energy content of the segments transport of energy (e.g., van lngen Schenau, 1989).
plus the power which Rows to the environment. As This expression should not be confused with the
such, it shows the causal relation between the three transport of energy between actual segments as de-
main existing approaches mentioned above. scribed by, e.g., Quanbury er al. (1975). Winter and
Robertson (1978). and Robertson and Winter (1980). If
one is interested in the flow of energy to actual
s. PROBLEMS IN APPLYING THE POWER EQUATION segments,one should use the separate equations [here,
equations (5) or (7)]. After summation of these sep-
5. I. General remarks arate equations, the information about flows of energy
Though it appeared to be possible to unambigu- between segments is lost.
ously deduce an instantaneous power equation based On the other hand it should be stressed that the
on a linked segment model, this result should mainly action of bi-articular muscles obviously also influ-
Survey Article 873

ences the net moments as used in the separate equa- and shoe. For deformations of the shoe. the same
tions. This means that the warning concerning the conclusion was drawn by Webb er al. (1988) in a
actual origin of power as made with respect to equa- calorimetric study where the power input PO was
tions (8) and (9) is also valid if one uses the separate compared with heat production. A considerable
equations as performed by Winter and Robertson amount of power may also be associated with running
(1978) and Robertson and Winter (1980). on loose sand or running on a treadmill the belt speed
Equation (9) is only useful when instantaneous of which is distinctly influenced by the runner. If the
flows of energy are to be studied. For many applica- speed fluctuations of the belt are small (e.g., < 5%). the
tions it is necessary to calculate a measure for mean flow of energy to the treadmill can be neglected (Webb
power production. This means that equation (9) has to et al.. 1988) and no mechanical difference exists
be integrated with respect to a time interval z and between level and overground running (van lngen
divided by 7;:: Schenau, 1980).
5.22. Cgcliny. In cycling, different applications of
T* Tt

(l/T,)
J 0
M,w,dt=(l/T,)
J 0
-ZF;v,dt

7.1
equations (12) and (13) are possible. When
equation
extremity,
(12) to a segmental
the measurement
analysis
of direction
applying
of the lower
and magni-
+(1/T;:)
J 0
(dZE,,,/dt)dt. (10) tude of the force applied
(Soden and Adeyefa,
on the feet by the pedal
1979; Ericson er al., 1986) should
be included. A second (and relatively simple) approach
Since, in cyclic movements. the cycles are repeated
possible in cycling is to use such a force measurement
each time interval To with the cycle frequency/= I / To,
to calculate the power delivered to the pedal.
equation (10) can be replaced by:
For the instantaneous power it should be noted that
TO in ergometer cycling(no air friction), the flow ofenergy
fJTO
0
Z M,w,.dt =/
J0
-XF;v,dt +fAE,,. (II) to the pedal should be equal
power minus the rntc of change of segmental energy
to the summed joint

Apart from the potential and kinetic energy of the [see equation (9)]. Recently this equality was valid-
body center of gravity, the mean change of segmental ated by cxpcrimcntal data (van lngcn Schcnau ct al.. in
energies during a cycle is zero. This means that AE,, press). Applying equation (13) to the entire system
reflects the change in potential and kinetic energy of (athlete and bicycle) will lead to a slightly lower result
the center of gravity of the athlctc. The left hand side of since frictional losses in chain, gearing. etc. are not
equation (11) can be used as a measure of mean accounted for. A few notes should be made concerning
generated power P,: this last application.
TO At lirst glance, the propelling force on the rear wheel
PM=/- A=/
J 0
Z M,co,dt (12) seems not to obey equation (4). Since its point of force
application is always at the same position relative to
where A is the mean work per cycle. This P, is also the bicycle frame and thus seems to have the same
equal to the mean power reflected by the right-hand velocity as the frame, it stems as if the propelling force
side of equation (1 I). However, it is often convenient adds power to the system. This paradox however is
to take not only the athlete but also his equipment due to the unique property of a wheel, and can easily
(bicycle, boat,‘ski) as a system. In that case one can use: be explained if it is realised that the velocity of the part

p:=/
J
of the tire which is in contact with the road is always
TO
Z-F;v,dt+AE,;f zero (when slip does not occur). The particles of the
(13)
0 tire which are in contact with particles of the road
where P,- Pi equals the power lost in the transmis- exert forces on each other with no relative velocity. A
sion system (in friction with the oar pin, chain, gearing, wheel thus can be imagined as a centipede with an
etc.). F, now reflects environmental forces which are inlinite number of legs that consecutively push elf
external to the entire system (propelling force on the against a fixed point of the road (Fig. 4). The main
rear wheel, water friction on the boat etc.). The dilierence with running is that in contrast to running,
applicability of these equations is now discussed for a the process of lifting these legs from the ground and
number of locomotor activities. * bringing them to the next point of contact hardly
requires any power. Given the other environmental
5.2. Terrestrid lucomotion forces, rolling and air resistance (Di Prampero et al.,
5.2.1. Running. In running. equations (12) and (13) 1979; Faria and Cavanagh. 1978; Raine, 1970;
represent only a fraction of P, and are of little use Zatsiorsky et al., 1982) a complete power equation
except in sprinting situations where a significant frac- can be formulated for different situations (level,
tion of PO is used to accelerate the body. uphill or downhill, with or without wind).
Equations (9) and (12). however, can be used to Though the frictional forces are described suffi-
achieve insightsinto the order of magnitude of power lost ciently in the literature, a note should be made on the
against environmental forces. As can be deduced from influence of (side-) wind. In the case of a wind velocity
Robertson and Winter (1980). a considerable amount Av, which has a different direction from the velocity v
of power may be dissipated by deformations of foot of the cyclist, one should first determine the magnitude
874 Survey Article

Fig. 5. Since the skate is gliding forwards during push-off.


the propulsive force FP of a speed skater can only lie in a
plane which is at right angles to the gliding direction.
Fig. 4. A wheel acts as a centipede with an infinite number of
legs that consecutively push offagainsta fixed point on the
road. of human swimming show a summation of powers
associated with the opposing (frictional) forces (see
of the sum (v + Av) before the air friction force F,,I can Toussaint ef al., 1983; de Groot and van Ingen
be determined [proportional to (v + Av)*]. The power Schenau, 1988, for references). We know, however,
required to overcome air friction then equals Far;v. A from experience, that during the strokes both in
calculation based on ‘components’ of air friction force swimming and rowing the push-of force is displaced.
based on v and Av respectively will lead to wrong This means that the power associated with this
results. If for example the wind Av is at right angles to v push-off force should bc accounted for according to
(pure side-wind), the latter method will lead to a power equation (9).
estimate which is equal to cycling with the same Since propulsion in water can only be achieved by
velocity in the absence of wind. The correct approach giving water momentum or ‘driving water backwards’
however will show that a side-wind always requires (Alexander. 1975) the water acquires kinetic cncrgy as
more power. a result of push-off. This phenomenon is well known
5.2.3. Spvrd skuting. In contrast to cycling where from studies of fish swimming (Webb, 1971, 1975;
one might speak of a ‘stcpwise’ displacement of the Lighthill, 1975; Alexander and Goldspink, 1977). Even
point of application of the push-off force, the proper in fish with relatively large surfaces which are involved
gliding tcchniquc in speed skating requires a push-ofi in generating propulsion, this amount of ‘wasted
while the skate continues to glide forwards (Haase. power’ (Alexander and Goldspink, 1977) is con-
1953; Djatschkow, 1977; van lngen Schenau er ul.. siderable. Expressed as fraction of P, henceforth called
1985; Boer cl (I/., 1986). The solution of this paradox is ‘propelling eficiency e,,’ (Alexander and Goldspink,
simple: it is essentially impossible for a forward gliding 1977; Lighthill. 1975). values for (Past) fish swimming
system to exert any force on the underlying surFAce in are between 65 and 75% (Bone, 1975; Webb, 1975;
the direction opposite to this gliding direction. So Videler and Hess, 1984). The power associated with
skaters can (and do) push oiTonly in a plane which lies generating propulsive forces in human swimming
at right angles to the gliding direction of the push off would be relatively easy to determine if the limbs
skate (Fig. 5). This results in the typical ‘sine wave’-like would simply move in a backwards direction using the
trajectory of the body center of gravity (see van Ingen drag on the limbs as the propulsive force. The pro-
Schenau ef al.. 1987 for further references). Though pelling efficiency in such a technique, however, would
essentially ditTerent from cycling, skating thus also result in a waste of power of more than 50% of P, (de
obeys equation (4), since the scalar product of force Groot and van lngen Schenau. 1988). Since the work
and velocity of its point of application is zero. The of Counsilman (1968). practitioners know that in
nature of the frictional forces is described in the particular the hands should not follow such a short
referenced literature citations and is comparable to the trajectory, but they should make as many side and
friclional forces in cycling. upwards excursions as possible. Schleihauf( 1979), and
Schleihauf et 01. (1983) showed that this technique is
5.3. Aquatic locomotion associated not only with the need to push offagainst
Though the inclusion of power associated with the water which was not previously accelerated
propelling force might seem rather trivial in locomo- (Counsilman, 1968; Toussaint et al., 1983), but also
tion where the push-off takes place against the entire with the creation of lift forces such as those that occur
earth, the significance of this approach based on in wings of birds and in hydrofoils (Lighthill, 1975;
equation (9) may become more convincing when ap- Alexander and Goldspink, 1977). The wasted power
plied to aquatic locomotion. The overwhelming associated with generating lift forces on the limbs is
majority of power equations applied for analysis considerably less than the power lost in generating
Survey Article 875

drag forces(Lighthill, 1975; Alexander and Goldspink, towed subject (e.g.. Di Prampero et al., 1974: Clarijs.
1977; de Groot and van Ingen Schenau, 1988). Lift, 1978). Since movements may disturb a steady-state
however, cannot be generated without drag. Much motion of the water relative to the body, this sugges-
future research in human swimming must therefore be tion seems reasonable (Daniel, 1983, 1984). A number
focused on the relation between technique and ep. The of authors have used indirect methods to determine
exact analysis of this propulsion mechanism is com- this so called ‘active drag’ (Di Prampero et al., 1974:
plicated from a hydrodynamic point of view (see Pendergast et al., 1977; Rennie et at., 1975; Clarijs.
Cheng and hurillo, 1984, for a three-dimensional 1978). In a more direct approach by measuring pro-
analysis). But even an experimental arrangement ac- pulsive forces Schleihauf(l979). Schleihauf er aI. (1983)
cording to $chleihauf (1979) and Schleihauf et al. and Van der Vaart et al. (1987) found values which are
(1983). who experimentally determined the lift and not significantly different from drag on passively
drag forces on hand and arm, will not allow a simple towed swimmers (see Van der Vaart et al. (1987) for
calculation of power. At first glance, one would tend to further discussion).
take the scalar product of the resultant of both forces Sanderson and Martindale (1986) reported a value
and the velocity of the hand. This however would of 75% for ep in rowing. Despite the absence of
ignore the so-called ‘induced power’, which is associ- significant lift forces in rowing, this value is still higher
ated with the creation of lift forces (Alexander and than that calculated for swimming, probably because
Goldspink, 1977; Tucker, 1975; Weis-Fogh. 1975). of the larger surface area of the oar blades. As with
One can envisage this effect by realising that the cycling, different parts of the overall system can be
hydrodynamics involved in creating lift are also re- modelled with a free body diagram using equation (9).
sponsible for’s deviation of the relative velocity of the (12) or (13).
water with respect to the hand in a direction opposite Taking the man-boat-oar system, only air friction
to the direction of the lift force (Fig. 6). This means (small), drag. and propulsive forces determine P,. The
that one cannot take the velocity of the hand relative net power output P, of the rower can be measured
to the undisturbed water as the velocity of the point of from the power input of the oar (Sanderson and
application of the resultant force on the water. Wardle Martindale, 1986). One should take care, however,
and Vidclcr ~(1980) show four different approaches with force measurements on the oarlock. Celentano et
which arc used to model the hydrodynamic processes at. (1974) used the force on the oarlock as the pro-
involved in creating propulsion in llsh swimming. pulsive force of the boat. This force however is only an
Apart from a few hypothetical examples (de Groot and external force if the boat (excluding man and oar) is
van lngcn Schcnau, 1988) based on hydrofoil theories, taken as a free body. In such an application, the force
no studies have yet been performed which relate from the seat and the forces on the foot stretcher
technique to the wasted power or to e,, in human should be accounted for as well. Neglecting inertial
swimming. The first indirectly determined values for ep elIects of the oar, the force on the oarlock may,
in freestyle swimming are reported to be approxim- however, be used to calculate the power input to the
ately 65% (Toussaint et al., 1988). oar (Fukunaga et al., 1986). It should be noted that,
There is considerable debate over the nature of particularly in rowing but essentially for all other
opposing forces in swimming. It has been suggested endurance sports too, one should not take the mean
that frictional forces (in swimming mostly referred to drag force and the mean velocity of the boat (or
as ‘drag’) experienced by swimmers moving their arms athlete) when calculating the power to water and air
and legs may ‘bedifferent from the drag on a passively friction. Instead, the time varying forces within the
cycle should be used to calculate cycle averaged
power, according to equation (131 The mean force and
velocity can lead to a significant underestimation of
the actual mean power (Nigg. 1983; Sanderson and
Martindale. 1986).
When comparing the examples presented of ter-
restrial and aquatic locomotion, it is clear that the
Vl differences in friction may explain the differences in
speed which are achieved in exercises of comparable
duration (Di Prampero, 1986). Next to the (small)
differences in power input which are due mainly to a
difference in total muscle mass involved in producing
P,. the differences in speed in terrestrial locomotion
Fig. 6. The pr pulsion force P, on the hand during swim- are mainly due to a difference in P,, which is associ-
ming is genera9 ed by moving the hand through the water ated with the creation of propulsive forces.
(relative velocit u,). Depending on theshape of the hand and In running, as well as in the classical diagonal stride
the angle betwL n the hand and the direction of u,. this
cross-country skiing. the push-offtakes place against a
relative flow d not only induce a drag force F, but also a
lift force F,. J-z generation of this lift force causes a change fixed point on the earth. This means that the athlete’s
of directidn of the relative velocity or the water. speed cannot exceed the maximal horizontal compon-
876 Survey Article

aquatic locomotion this expression does not account


for ‘wasted’ power at push-off.
.
6. DEFINITIONS

Based on the foregoing deduction of an unambigu-


ous power equation. it is now possible to reconsider
the various definitions that are commonly used.

6.1. Power
As indicated in the Introduction, there are a number
of different definitions of power and efficiency in the
Fig. 7. External forces acting on a cyclist. Note that in the
literature of human and animal locomotion. The same
reference frame used the system has potential energy which
depends on both coordinates. power is expressed as internal by one author and
external by another. In particular in running and
walking there is no concensusabout these expressions
ent of the velocity, which can be real&d between the (e.g., Cavagna and Kaneko, 1977. vs Winter, 1978).
body center of gravity and the sole of the foot Most authors, however, use reasonable arguments for
(Djatschkow, 1977). This velocity is, to a large extent, their definitions. It does not seem possible to relate
dependent on the extension velocity of the joints of external and internal power simply to the free body
the lower extremity. In cycling and speed skating, the diagrams which are used in the different approaches.
speed of the athlete is not constrained by the extension Based on the free body diagrams for the rigid links
velocity of the leg since the point of application of used by Aleshinsky (1986a) and by the present
the propelling force is displaced during a leg extension. authors, one might argue that all power comes from
This phenomenon partly explains why competitive sources external to these links. But a definition based
cross-country skiers use a skating-like technique on such a number of different free body diagrams does
nowadays. not hold as soon as the derived equations are added in
At the end of this discussion of the application of order to achieve the power equation for the entire
power equations, one additional approach should be athlete. Following the arguments of Winter (1979).
considered. Imagine a system (such as a cyclist) which only the second expression on the right-hand side of
moves uphill under the influence of a propulsive force equation (9) deserves the designation of external po-
F, and is opposed by friction forces F, (Fig. 7). An wer. The first expression should be defined as internal
additional equation for P can be deduced if the rate of power. Though such a definition would be closely
change of energy of the center of mass is calculated. In connected to the environmental (external) forces, it
the coordinate system indicated in Fig. 7. the poten- would change as soon as one replaces the rate of
tial energy can be expressed as E,= mg sin 01.x change of potential energy by the work done by
+ntgcosa~z, while the kinetic energy equals ~rno~ gravity. The same is true for kinetic energy, which can
with m the mass of the system and u, its velocity in the be replaced by work done by (external) inertial forces.
x-direction, if the velocity in the z-direction is zero So it can be concluded that there are no decisive
(u, = 0). arguments against a definition of external power
The time derivative of the total energy than equals: according to equation (9) if one intends to couple the
expressions ‘internal’ and ‘external’ to a well-defined
dE/dt=mgsinzo,+mgcosau,+mu,a,
free body. It should further be noted that Winter’s
=mgsinau,f(F,-Fr-mgsina)u, definition can lead to situations where the system does
positive external work on the environment without
=Fpu,-Ptu,.
simultaneous muscle actions. A cyclist who, for exam-
According to equation (9). the power is then: ple, passively descends a hill, would deliver external
work (against friction), and negative internal work.
P,=Fp-vp-F,*v,+F,u,-Fro,.
Though both work expressions would cancel out in
In terrestrial locomotion, u,=O. The magnitude of Winter’s proposed expression for total mechanical
- F,* v, is F,- vX, since F, and v, have opposite work (Winter, 1979). the same zero work output can
directions. Then be achieved by a more straightforward approach using
equation (9). (12) or (13). As already discussedabove,
P,= Fg,.
the summed joint power is the power which is most
This means that the instantaneous power which is closely related to muscle power. So it is proposed here
available to overcome friction and to increase the to define this power as external power. All power P,
energy of the center of mass can simply be calculated -P, is then internal power. This, at least, is an
from the propulsive force and the velocity of the mass unambiguous definition which fits the majority of
center of gravity, which may serve as an additional previous definitions for activities other than walking
tool in many applications. Note however that in and running. Note, however, that this power depends
Survey Article a77

upon the chosen free body which at least includes the For various animal muscles. experiments on iso-
athlete but which may also be extended to the athlete lated muscles show efficienciesfrom 14 to 40% (Gibbs
and his equipment (bicycle, boat. skate, ski. etc.) using and Gibson, 1972; Wendt and Gibbs, 1973; Heglund
equation (13). and Cavagna, 1987; de Haan et al., 1989).
The majority of the mechanical power as calculated However, the highest values are obtained by only
using different methods for level running and walking accounting for the positive work done in the concen-
is thus part of the internal power. The best expression tric phase of muscle action (e.g., Heglund and Cav-
to use here is probably ‘mechanical power’. Other agna, 1987). It should be noted that one might
expressions which are unambiguously defined in the even find efficiencies up to 100% or higher if only
literature and’which are already used in this paper positive work is taken into account. Imagine for
concern (Fig. 1): example that the contractile element acts as a force
(1) power input or metabolic power P,; generator (without detachment of cross-bridges) in a
(2) power output PO or muscle power; ‘stretch-shortening cycle’ of a muscle tendon complex
(3) maintenance power P,: with short muscle fibers and a long compliant tendon.
(4) conservative power P,; In such actions the positive work (from re-utilisation
(5) non-conservative power P,, (e.g., Tucker, 1975). of elastic energy) can be considerable with little meta-
Note that P, can be part of both external and bolic cost. If, however, the efficiency is measured on
internal power: while P,, is always part of the internal the level of the contractile elements, the (actual) muscle
power. Other definitions which are recommended efficiency seems not to exceed the 25% in actions
concern basically swimming: starting from an isometric state as well as in
(6) wasted power (e.g., Alexander and Goldspink, stretch-shortening cycles (de Haan et al., 1989).
1977) P, (k: kinetic energy given to the water). This is For negative work one can also speak about effi-
all power associated with F,; ciency, but it should be noted that here this expression
(7) induced power (e.g., Lighthill. 1975). the power has a totally different meaning. At the level of con-
associated with lift forces in the generation of F, (part tractile machinery, an eccentric muscle action simply
of the wasted power); acts as a brake (Stainsby et al.. 1980) converting
(8) profile power (Tucker, 1975; Weis-Fogh. 1975) mechanical work into heat. This means that the
the power associated with drag forces in the gencta- cxprcssion ctlicicncy now provides a measure for the
tion of F, (also part of the wasted power); cost to control the brake. It has even been suggcstcd
(9) parasitic power (Tucker. 1975; Wcis-Fogh. that during txcentric muscle actions, the cross-bridges
1975). the power associated with F,, the opposing can attach and develop tension without metabolic
frictional forces. This definition is also recommended costs (Curtin and Davies, l974), the costs of an
for terrestrial locomotion. eccentric action being mainly due to the process of
muscle activation. The magnitude of this efficiency of
6.2. EJiciency negative work will (at equal metabolic costs for the
The many diiflerent delinitions of ellicicncy that can activation process) be strongly dependent on the force
be found in the literature are, on the one hand, level and range of lengthening of the contractile
associated with the dilferent types of mechanical element, and can ofcoursc in principle be considerably
power and, on the other hand, with different estimates larger than 100%.
of metabolic power input. Again, there has been much It should be emphasized that one cannot relate
discussion concerning the various definitions (e.g., phasesof negative summed joint work to this negative
Whipp and Wasserman, 1969; Donovan and Brooks, muscle work and to the efficiency of negative work.
1977; Stainsby cr al., 1980; Fish, 1984; Cavanagh and Particularly during locomotion, much of this negative
Kram. 1985a,b) and reasonable concensus has emer- joint work will be stored as elastic energy in the series
ged concerning some of them. This is especially true elastic elements of the muscles, and re-utilised as
for muscle efficiency e,, which is defined as the positive work.
product of the phosphorylative coupling efficiency At the level of separate joints one can also expect
and the contraction coupling efficiency. This muscle that negative work at one joint appears as positive
efficiency e, = P,/( P,- P,) (see Fig. I) reelects the work in another joint through actions of bi-articular
efficiency of thelcontractile machinery and is estimated muscles (van lngen Schenau. 1989).
not to exceed the e,= 30% (Astrand and Rodahl,
1977). This estimation is based on thermodynamical
6.3. Gross eficiency
considerations (Stainsby et al., 1980; Astrand and
Rodahl, 1977). In contrast to measurements of effici- Expressed in terms of power components as pro-
ency on the level of total body movements, efficiency posed in this paper, this widely used expression may
data on isolated muscles or muscle fibers are relatively unambiguously be dcflned as
scarce. Such dlata. however, are important, since
es= P./P,.
clearly the effidency of total body movements has to
be lower than the efficiency measured on the level of The quotients between other possible measures for
the contractile machinery. power discussed above and power input Pi might be
878 survey Article

defined by a more trivial expression ‘mechanical effici- far in excess of the thermodynamically determined
ency’, indicating that this efficiency is not associated maximal muscle efficiency (see Asmussen and Bonde
with an explicitely defined free body (as can be done Petersen, 1974; Pugh. 1971; Zacks, 1973). Since the
with contractile clement(s) or the entire athlete). examples concern efficiencies of external power, these
Clearly, the gross efficiency will always be lower discrepancies cannot be explained by internal storage
than the muscle efficiency discussed above. In this and re-utilisation of the elastic energy of muscles and
respect it should be realized that, particulary in not- tendons.
highly-trained subjects, one can expect a contribution There appears to be enough evidence to completely
of anaerobic power already at relatively low levels of reject the validity ofany base-line subtraction. There is
exercise, which is not accounted for in the denom- only one unambiguous definition possible for gross
inator if P, is estimated on the basis of oxygen body movements at the present state of knowledge:
consumption. This leads to an underestimation of P, the gross efficiency. This is the only quotient which is
and an overestimation of the gross efficiency. based on valid, well defined powers.

6.4. Baseline subtractions 6.5. Economy


It is obvious that the gross efficiency will always be As argued by Cavanagh and Kram (1985a);
(much) smaller than e,. Values reported in the liter- Morgan (1985); Williams (1985) and Daniels (1985).
ature are about 22-23% for cycling (Pugh, 1974; the concept of gross efficiency or a mechanical effici-
Steinacker et al., 1984). 17-19% for rowing(Steinacker ency is of little use in the study of the mechanics and
et al., 1984: Fukunaga et al., 1986), 21% for cross- energetics of walking and running (the gross efficiency
country skiing (Niinimaa et al.. 1978) and 19-2f% for is close to zero in level walking and running). There-
speed skating (de Groot et al., 1988). For human fore these authors advocate the application of ‘eco-
swimming. much lower values are reported (e.g., Craig nomy* defined for a particular activity (e.g.. a certain
ef al.. 1985; 6-g%). These values however do not speed) as the rate of submaximal oxygen uptake either
include the wasted power Pk. If the total power P, is in absolute units or relative to body weight
accounted for, one may expect a gross efficiency in (ml kg-’ min-I). While this allows both direrent ac-
swimming close to or even higher than the values of tivities (such as uphill running) and diffcrcnt indi-
about 15% as reported for armcranking(Powers et al.. viduals performing the same activity to be compared.
1984). it can not make any statement with respect to effici-
Given the difTerencesin the involvement of muscle ency. One individual may be less economical than
mass in generating P,, in maintenance power P,. and another, but may actually be more elXcicnt bccausc he
given the large differences in P,, which may exist is actually performing considerably more work. Elow-
between the diflerent movements, these variations in ever, economy is a fairly unambiguous concept which
reported gross efficiency can be understood. Part of might be applied for other types of locomotion as well.
the variations will also be due to difTerencesin e, as a Note however that this expression is also used in
result of differences in, e.g., the force and velocity muscle mechanics to represent the force-time integral
patterns of the contractile elements. Though the num- divided by energy input over the integration time,
ber of unknown variables is far in excess over the mostly applied in isometric conditions (Goldspink.
number of equations which can be deduced by 1978).
measuring P, at rest, in unloaded movements (P, I 0)
and in dimerent loaded movements (P,>O), many 7. CONCLUDING REMARKS
physiologists still perfoim questionable attempts to
identify e, from measurements of P, and P, in gross The literature reviewed here exhibits a considerable
body movements. These attempts are known in the diversity in the approach to the formulation of a
literature as base-line subtractions (e.g., Gaesser and power equation for endurance activities. The method
Brooks, 1975; Donovan and Brooks, 1977).The idea is that we have suggestedcan be seen as somewhat of a
that, e.g., the power input in unloaded cycling equals bridge between the energy approach of Winter and his
all power except the power input necessaryto produce associates(Winter, 1979, 1983; Robertson and Winter,
P, in loaded cycling. So by subtracting P, in unloaded 1980) and the theoretical approach of Aleshinsky
cycling from the power input in cycling at an external (1986a-e). The former method involves assumptions
power P,. one attempts to deduce a measure for concerning energy exchange while the latter approach,
muscle efficiency. although computationally rigorous, is fairly complex
Many arguments can be and are formulated against in form and lacks application to specific activities. The
these approaches (e.g., Stainsby et al., 1980; Cavanagh application of the present approach to a number of
and Kram. 1985a). The main argument is that it is terrestrial and aquatic activities indicates the utility of
unlikely to be true that the maintenance power would the method. It should be realized that there is unlikely,
be independent of the workload. Moreover the so- in the near future, to be an entirely satisfactory
called work or apparent efficiencies [ = P,/( PIa,.,,drd solution to the problem of power output calculation in
- PL,““kmd.d )I and delta efficiencies (=AP,/AP,) locomotor activities, since a number of intrinsic fac-
(Gaesser and Brooks, 1975) lead to values which are tors remain difficult to quantify. It is, however, hoped
Survey Article 879

that the clarification of certain concepts related to Cavagna, G. A. and Kaneko. M. (1977) Mechanical work and
force, power. eflkiency and economy during move- efficiency in level walking and running. J. Physiof. 268,
ment presentad in this review will assist in the further 467-481.
Cavagna. G. A., Dusman. B. and Margaria. R. (1968) Positive
development of this sometimes ambiguous field of
work done by a previously stretched muscle. 1. uppl.
study. Physiol. 24. 21-32.
Cavagna. G. A., Mazzanti, M., Heglund. N. C. and Citterio,
Acknuwledgemehts-The comments and suggestions of the C. (1985) Storage and release of mechanical energy by
members of tl@ research group ‘Cyclic movements’ of active muscle: a non-elastic mechanism? J. exp. Biof. 115.
the faculty of Human Movement Sciences are gratefully 79-87.
acknowledged. Cavagna. G. A.. Thijs, H. and Zamboni, A. (1976) Thesources
of external work in level walking and running. 1. Physiol.
262.639-657.
Cavanagh. P. R. and Kram. R. (1985a) The efficiency of
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