REVIEW ARTICLE

R. Hugh Morton
The critical power and related whole-body bioenergetic models
Accepted: 13 October 2005 / Published online: 12 November 2005
Springer-Verlag 2005
Abstract This paper takes a performance-based ap-
proach to review the broad expanse of literature relating
to whole-body models of human bioenergetics. It begins
with an examination of the critical power model and its
assumptions. Although remarkably robust, this model
has a number of shortcomings. Attention to these has
led to the development of more realistic and more de-
tailed derivatives of the critical power model. The
mathematical solutions to and associated behaviour of
these models when subjected to imposed exercise can
be applied as a means of gaining a deeper understanding
of the bioenergetics of human exercise performance.
Keywords Anaerobic work capacity Energy
Endurance Exercise Metabolism Performance
Introduction
Over recent decades there appears to have been a drift
towards a more micro view of human bioenergetics,
focussing more towards the cellular and even molecular
end of the spectrum. Although the macro view has not
been neglected, it is nevertheless timely to examine
models of human bioenergetics from the whole-body
perspective. The human body is a highly complex
organism and has been the subject of intense scientic
scrutiny for several centuries. Despite considering a
broad picture of only one aspect of its functioning, I do
not make any special attempt to be comprehensive in
this review. Rather, I attempt to blend the essential
theoretical elements of these models and their mathe-
matical solutions with the practical and performance-
related interpretations of their properties. In so doing, I
hope their value as an investigative research vehicle or as
a framework for practical aspects of the study of human
performance, or even as a teaching tool for undergrad-
uate or postgraduate students, can be evaluated.
The review begins with an examination of the simple
critical power model and its assumptions. When sub-
jected to scrutiny, this model is remarkably robust, but it
nevertheless has a number of shortcomings. Remedial
attention to these has led to the development of more
realistic and more detailed derivatives of the critical
power model. The mathematical solutions to these
models under a variety of exercise scenarios provide
fertile ground for gaining a better understanding of the
bioenergetics of human exercise performance.
This review is not intended to update the literature on
critical power (Hill 1993; Vandewalle et al. 1997) nor that
on the relationship between power output and endurance
time (Billat et al. 1999a; Morton and Hodgson 1996) nor
its relationship to whole-body fatigue (Walsh 2000), but
rather to set the critical power model and its derivatives in
the wider bioenergetic modelling context.
The critical power model
Origins, assumptions and development
It was Monod and Scherrer (1965) who rst observed a
hyperbolic relationship between the level of constant
power output (P) and corresponding time to exhaustion
(t) in a single muscle group. This relationship, when
transformed knowing that total work performed (W) is
given by the product Pt, can be expressed in a linear form.
The intercept of this line was termed the anaerobic work
capacity (AWC) and its slope termed the critical
power (CP). Mathematically the two equations:
W AWC CP t 1a
and
t AWC=(P CP) 1b
R. H. Morton
Institute of Food, Nutrition and Human Health,
Massey University, Private Bag 11-222,
Palmerston North, New Zealand
E-mail: h.morton@massey.ac.nz
Tel.: +64-6-3504265
Fax: +64-6-3505781
Eur J Appl Physiol (2006) 96: 339354
DOI 10.1007/s00421-005-0088-2
are equivalent. There are in fact four other equivalent
equations:
P AWC=t CP, 1c
a transformation rst proposed by Whipp et al. (1982)
t (W AWC)=CP 1d
and
P W CP=(W AWC), 1e
W AWC P=(P CP) 1f
although only the rst three are in common use. Given
that t is the most natural dependent variable, and P the
most natural explanatory variable, Eq. 1b is the pre-
ferred version from an exercise performance viewpoint,
though a similar case could be made for Eq. 1f.
Moritani et al. (1981) extended those ideas for the rst
time to whole-body exercise on the cycle ergometer. The
four essential assumptions of the whole-body CP concept
as it became known (Hill 1993) are the following:
1. There are only two components to the energy supply
system for human exercise, termed aerobic and
anaerobic.
2. The aerobic supply is unlimited in capacity, but is
rate limited. This limiting parameter is the critical
power, CP.
3. The anaerobic supply conversely is not rate limited,
but is capacity limited by the parameter AWC.
4. Exhaustion, and by implication termination of exer-
cise, occurs when all of AWC has been utilised.
From these assumptions it can be deduced that for
constant power P CP, the notion of exhaustion is
vacuous and t is undened or innite, whereas for
P>CP, Eq. 1b and its equivalents follow directly. Over
the last 20 or more years, a large number of studies have
found good ts of data to Eqs. 1a, 1b, 1c, at least within
certain bounds.
It is important nevertheless to realise that implicitly
embedded in these assumptions are a number of others,
specically:
5. Aerobic power is available at its limiting rate CP the
moment the exercise begins and remains so right up
until the end of the exercise.
6. The power domain over which the model applies is all
of CP<P<.
7. The time domain over which the model applies is all
of 0<t<, and endurance at CP is innitely long.
8. The eciency of transformation of metabolic energy
to mechanical energy is constant across the whole
power (and time) domain(s).
9. CP and AWC are constants, independent of P (and/
or of t).
All models of real processes are necessarily simpli-
cations and the above assumptions are indicative of
them. There are of course unrealities to varying degrees
in all of them, and these will be discussed in Extending
the critical power model when extensions to the CP
model and the reasons behind them will be considered.
Parameter interpretation and estimation
The assumptions above suggest clearly enough that CP
is aerobic in nature and that AWC is anaerobic, but
empirical verications are vital. It was Moritani et al.
(1981) who rst provided evidence for these interpreta-
tions. They found CP to be highly correlated with the
ventilatory anaerobic threshold and AWC to be unaf-
fected by hypoxia. Several other early studies (Green
and Dawson 1993; Green et al. 1994; Green 1994, 1995;
Housh et al. 1991; Jenkins and Quigley 1990, 1991;
Talbert et al. 1991) have provided similar evidences,
though the evidence is not all conclusive (Housh et al.
1992). Perhaps the most convincing evidence of the
aerobic and anaerobic natures of these parameters can
be found amongst the collection of studies reporting
interventions and their impact on the CP model, which
will be discussed in Interventions and their impact on
the model.
In order to estimate the parameters CP and AWC for
any subject, it is necessary to complete at least two
separate trials, cycling to exhaustion at two dierent but
constant power outputs. In practice four or more are
common, spread across a spectrum, though six or more
are uncommon. On each occasion P and t (and hence
W = Pt also) are recorded. Any one of the six equation
forms can be tted using any standard statistical curve-
tting software. It is important to choose these experi-
mental design aspects carefully, for the CP and AWC
estimates may dier according to how many trials are
performed (Housh et al. 1990; Taylor and Batterham
2002), what durations (and therefore also what power
selections) are planned (Bishop et al. 1998; Jenkins et al.
1998; Poole et al. 1986) and which of the mathematical
forms of the model may be used (Bull et al. 2000;
Gaesser et al. 1990, 1995; Hill et al. 1993; Hill 2004;
Housh et al. 2001). Furthermore, the rest interval be-
tween trials may aect the estimates also (Bishop and
Jenkins 1995; Hill 1993) varying between 30 min and
over 24 h in dierent studies.
It has been observed too that variations in repeated
performances are larger when durations are longer
(McLellan et al. 1995; Poole et al. 1988). For this reason
the exact nature of the curve-tting routine employed
may also aect the estimates. It may be statistically
advisable to use weighted regression, giving less weight
to long-duration observations and vice versa (Morton
and Hodgson 1996).
Performance implications
Application of the CP model using data from any sub-
ject clearly provides an assessment of two dimensions of
the bioenergetic characteristics of that individual. Then,
340
employing Eqs. 1b, 1f for example and substituting any
given value of P provides predictions of that individuals
performance (i.e. endurance) time and amount of work
that could be accomplished, respectively. Such perfor-
mance aspects are obvious and have been employed in
determining the ability of humans generally as sources of
mechanical power (Wilkie 1960) or specically in activ-
ities such as human-powered ight (Nadel 1988, 1996).
Of far more relevance in todays world of competitive
sports performance is the question of optimal strategy.
For example, can performance time (and the amount of
work accomplished) be improved by working at other
than the constant power throughout determined by the
solution to the CP model equations for that individual?
Fukuba and Whipp (1996, 1999) have solved this
problem (albeit for the running analogue of the CP
model). The answer, or at least part of it, is No. They
also demonstrate that any one of an innite number of
strategies involving stagewise steps of periods of con-
stant power produces an identical endurance time, pro-
vided P never drops below CP. This can also be shown
to be true when P changes (always above CP) in any
smooth and continuous fashion (Morton 2004).
Conceptualisation as a hydraulic model
This conceptualisation is originally due to Margaria
(1976) and can be described as follows. If we envisage
uid contained in a vessel or vessels to represent meta-
bolic energy and the ow of uid therefore to represent
power, the CP model can be diagrammatically repre-
sented as in Fig. 1.
The system is comprised of two vessels representing
the aerobic (A
e
) and anaerobic (A
n
) energy supply
components (assumption 1). A
e
is of innite capacity
(assumption 2). R
1
is a connecting tube of xed diame-
ter, which determines the limit to the uid ow from
A
e
to A
n
. This limit is denoted by CP (assumption 2).
Vessel A
n
is of limited capacity, denoted by AWC
(assumption 3). T is a tap which can be opened to allow
an unlimited positive ow P (assumption 6). This in turn
(since CP is nite) implies that the rate of decrease of the
uid level in A
n
is also unlimited (assumption 3). Pro-
vided P>CP (assumption 6), the uid level in A
n
must
drop to make up the dierence PCP, and P can no
longer be maintained when A
n
becomes empty
(assumption 4). This event occurs at time t>0
(assumption 7). The ow through R
1
is always equal to
exactly CP, determined only by the unchanging depth of
vessel A
e
(assumption 5). When P=CP or less, uid
from A
n
is not required and A
n
will never be emptied,
i.e. t is innite (assumption 7). Since the whole hydraulic
model is a physical construct with xed dimensions and
containing an incompressible uid, the volumetric and
rate (and therefore also eciency) parameters are xed
and independent of P (assumptions 9 and 8).
All the above lead directly to the derivation of Eq. 1b
from rst principles. Knowing also that W = Pt enables
all other mathematical equivalences to Eq. 1b to be
derived algebraically.
Interventions and their impact on the model
The CP model has been the object of a large number of
intervention studies designed to discover what happens
when it is exposed to deliberate changes in conditions.
These studies have assisted signicantly in our under-
standing of the model. In general, their results conform
to what physiological logic might lead us to expect and
arms the overall nature of the concept and its com-
ponent structure.
An early study of Jenkins and Quigley (1990) showed
that a group of trained subjects had a signicantly
higher average CP and AWC than another group of
untrained subjects. Since that time, more specically
designed training studies have examined the direct ef-
fects of various training strategies on changes in CP and/
or AWC in the same subjects. Given the aerobic nature
of CP, it would be expected that endurance or pre-
dominantly aerobic training would increase the CP of
those undertaking such training. A number of studies
have conrmed that this indeed occurs (Gaesser and
Wilson 1988; Jenkins and Quigley 1992; Jones and
Carter 2000; Poole et al. 1986). Likewise it would be
expected that strength or predominantly anaerobic
training would increase AWC and, from the outset,
studies do conrm this (Jenkins and Quigley 1992; Poole
et al. 1990; Stokes et al. 1993). Interval training, in which
periods of low-intensity training alternate with periods
of high intensity, is generally regarded as a particularly
eective overall training strategy (Billat 2001; Laursen
and Jenkins 2002) because it taxes energy supply
mechanisms across the spectrum. Inferring from the
above, it would be expected and has been shown that
interval training improves both CP and AWC (Gaesser
and Wilson 1988; Poole et al. 1990).
From the hydraulic representation of Fig. 1, aerobic
training appears to increase the diameter of pipe R
1
. It
would not likely increase the depth of vessel A
e
, as this Fig. 1 The CP concept as a hydraulic model
341
depth could be viewed as the analogue of the O
2
partial
pressure in the inspired air. Strength training on the
contrary appears to increase the volume of vessel A
n
,
probably via a cross-sectional increase rather than a
depth change.
Again, given the aerobic nature of CP, it would be
expected that subjects tested under hypoxic conditions
would evidence reduced CP, while comparative tests
under hyperoxic conditions would evidence an elevated
CP. This appears to be the case (Moritani et al. 1981;
Whipp et al. 1982), whereas AWC is unchanged. From
the hydraulic model perspective similarly therefore, hy-
poxia probably represents reduced depth of vessel A
e
rather than a constriction of R
1
, whereas hyperoxia
represents the converse.
Since AWC is regarded as an anaerobic store repre-
senting energy derived from high-energy phosphates and
glycolysis leading to net lactate formation, it may be
expected that creatine and/or carbohydrate supplemen-
tation would increase AWC. Studies show that this is the
case (Miura et al. 1999; Smith et al. 1998; Stout et al.
1999). Creatine supplementation has no eect on CP,
but its eects on AWC are augmented by carbohydrate
supplementation. No studies have reported the eects of
carbohydrate supplementation alone. Like strength
training, these types of supplementation appear to in-
crease the volume of vessel A
n
.
Likewise, if prior exercise is undertaken with little or
no recovery or if the timing is only started once VO
2
max is reached, a diminution of AWC would be ex-
pected. Once again, empirical evidence conrms these
expectations (Billat et al. 1999a, b; Heubert et al. 2003,
2004; Swanson et al. 1992).
Diering pedal cadences have been considered as a
confounding factor with the potential to cause changes
in the CP and/or AWC estimates. The available evidence
is to some extent contradictory (Carnevale and Gaesser
1991; Green et al. 1995) but it is believed that it is best to
allow subjects to select a cadence at which they are
comfortable, at least between the limits of 60100 rpm
(Hill et al. 1994; Hill 2004).
Another confounding factor which does appear to
aect CP and/or AWC estimates is a learning eect,
manifesting under repeat performance or test/retest
conditions (Scarborough et al. 1991; Smith et al. 1991).
As would be expected, performances are better and
estimates higher on subsequent testing occasions.
In summary, this array of interventions serves to
strongly arm the robust nature of the concept in
assigning an aerobic character to CP and an anaerobic
character to AWC. The interpretation of these param-
eters is now rmly established.
Applications to other constant power exercise modalities
Although Lloyd (1966) rst suggested application of
these ideas to running, it was Hughson et al. (1984) who
rst published the details in which critical velocity (CV,
measured in metre/second) and anaerobic distance
capacity (ADC, measured in metres) were considered
and evaluated as analogues of CP and AWC, respec-
tively. While not subject to quite the same degree of
scrutiny as the original form of the CP concept, studies
do in general conrm those results reported previously.
For example, CV has been found no dierent from the
maximal lactate steady-state running velocity or from
the lactate turn point velocity (Smith and Jones 2001),
and to be a threshold velocity above which exercise of
sucient duration will lead to attainment of VO
2max
(Hill and Ferguson 1999). The success with which the
running version of the concept is able to predict times at
running various distances does, however, seem in some
doubt (Kolbe et al. 1995; McDermott et al. 1993) and
the choice of which equation of the model to use also
seems unclear (Housh et al. 2001).
In the same way as for running, CV and ADC can be
evaluated for swimming (Biggersta et al. 1992; Steward
et al. 1994; Wakayoshi et al. 1992a, b). Although subject
to even less scrutiny than running, the critical swimming
velocity corresponds to the maximal lactate steady-state
swimming velocity (Wakayoshi et al. 1993) and corre-
lates signicantly with swimming VO
2
at the anaerobic
threshold and the onset of blood lactate accumulation
(Wakayoshi et al. 1992a, b) and with the lactate
threshold (Martin and Whyte 2000). Critical swim
velocity has been found to correspond to a maximal
stroke rate which can theoretically be maintained con-
tinuously without exhaustion (Dekerle et al. 2002). The
choice of equation for swimming, however, seems to be
of little consequence (Lane et al. 1994), but there is
conicting evidence on whether the anaerobic swimming
distance capacity provides a reliable estimate of anaer-
obic capacity or not (Biggersta et al. 1992; Dekerle
et al. 2002; Lane et al. 1994).
In theory of course, any exercise modality in which
physical work can be accomplished, or which involves
some form of locomotion, oers scope for application of
the CP concept. Several of these have been briey re-
ported, such as kayaking (Clingeleer et al. 1994),
wheelchair propulsion (Arabi et al. 1999), weightlifting
(Morton et al. 2005) and rowing (Hill et al. 2003; Ken-
nedy and Bell 2000). Cycling when regarded as a means
of locomotion appears to have not been considered.
In all these cases, whatever the form of locomotion,
the relationship between energy cost of transport and
speed over the applicable range must be known and
explicitly introduced into the calculations (di Prampero
1999).
Applications to variable power output exercise
There is nothing particularly special about continuous
exercise at constant power, except perhaps that the
mathematical solutions (and the associated curve tting
to data) to the CP model under such conditions are
extremely simple. One exercise protocol common in
342
laboratory testing settings is ramp exercise, where power
output starts at zero (though in general this need not
necessarily be the case) and increases linearly at some
xed rate, referred to as the ramp slope, s. The CP model
has been adapted for ramp exercise (Morton 1994), ex-
cept that endurance time becomes a function of s, albeit
mathematically slightly more complex than linear or
hyperbolic. Like the original CP model, it too is avail-
able in several dierent mathematically equivalent forms
(Morton 1997). Its validity has been questioned (Van-
dewalle 1995) despite the fact that its assumptions are
precisely the same as for the constant power CP model
(Morton 1995). Indeed, investigation has shown that
estimates of both CP and AWC from ramp tests are not
signicantly dierent from those derived from constant
power tests in the same subjects (Morton et al. 1997).
Similarly, there is nothing special about continuous
exercise per se. Many sports (particularly team games)
involve exercise of an intermittent nature, where periods
of relative rest and intense exercise are interspersed. In-
deed intermittent exercise (Christensen 1960) has for long
been regarded as a particularly benecial formof exercise
(Billat 2001; Laursen and Jenkins 2002). Can the CP
concept therefore be applied to intermittent exercise?
Examination of the hydraulic representation of the
CP model depicted in Fig. 1 suggests that if tap T were
to be closed, corresponding to the cessation of work (or
a period of relative rest), then vessel A
n
would rell by
virtue of the ow through R
1
. Only when A
n
had been
completely relled would the ow through R
1
cease.
Thus for intermittent exercise one can regard the intense
exercise periods as drains on A
n
and the relative rest
periods as relling A
n
. This is exactly the approach ta-
ken by Morton and Billat (2004), who nd good ts of
the derived model equation to data, but the CV esti-
mated from continuous running was signicantly higher
than the CV estimated from intermittent running in the
same subjects. This discrepancy may be due to increased
sensitivity of the model to the implicit assumption of the
innitely fast step increases and decreases in VO
2
occurring many times in this mode or exercise. In
addition, it should be noted that this particular appli-
cation of the CV concept is quite dierent to that of
Kachouri et al. (1996) and others, who simply apply the
standard model approach to aggregated intermittent
running times. Rather than being an example of the
limits of the CP concept as claimed, Kachouris appli-
cation is perhaps more an example of an inappropriate
or inaccurate procedure, not based on proper consider-
ation of the CP concept assumptions detailed above.
Extending the critical power model
Questioning the assumptions
All models purporting to represent aspects of the human
system are necessarily simplications, and the CP model
is no exception. All nine of the assumptions of Origins,
assumptions and development can be questioned. In
brief, the following critiques can be made of them.
1. There are only two components to the energy supply
system for human exercise, termed aerobic and
anaerobic. While it may be true that there are only
two biochemical pathways, aerobic and anaerobic, to
human energy metabolism, there are certainly more
than two components to the energy source or supply.
The aerobic pathway taps into both glycogen and fat
as fuel sources. The anaerobic pathway comprises a
lactic component in which glycogen metabolises
resulting in net lactic acid production and an alactic
component (essentially the high energy phosphate
compounds or phosphagens) whose metabolism does
not result in lactic acid production. Incorporating all
these aspects realistically will of necessity complicate
the modelling. It has been attempted in a simple
fashion, described in Applications and developments
of the 3-component hydraulic model.
2. The aerobic supply is unlimited in capacity, but is rate
limited. This limiting parameter is the CP. Since the
aerobic pathway requires an energy substrate to be
oxidised, the amount of which is of course limited in
any human, it is clearly untrue to assert that the
aerobic capacity is unlimited. It may well be relatively
large (in energy units) in comparison to other avail-
able energy substrates by an order of magnitude or
more, and this may well explain the robust character
of the CP model at least for exercise of moderate
duration. However, it is well established that the
aerobic supply is rate limited. Whether this rate is
called the critical power, maximum aerobic power
(MAP) or VO
2max
, or indeed some other convenient
term, is not the issue. Some upper limit clearly exists.
3. The anaerobic supply conversely is not rate limited, but
is capacity limited by the parameter AWC. The
anaerobic supply component likewise requires energy
substrates, and so it is quite reasonable to assume a
limited capacity. While explosive muscular power
may well be quite high, it is clearly not unlimited.
Humans cannot run with instantaneous velocity of
more than about 13 m/s, and if the resistance on the
pedal cranks of a cycle ergometer is too high, the
pedals simply cannot be turned. No physical work is
possible and no power is generated. The notion of an
endurance time at such intensities is clearly vacuous.
4. Exhaustion, and by implication termination of exercise,
occurs when all of AWC have been utilised. While it
may be reasonable for a mechanical engine to cease
when its usable fuel is fully depleted (provided there is
no other malfunction), the human engine appears to
be dierent. It has for long been known that at
exhaustion, signicant amounts of unused glycogen
remain, and that the higher is the power requirement
at exhaustion, the more it is that remains (Saltin and
Karlsson 1971). Furthermore, it is well established
that if the power requirement at the point of
exhaustion were to be signicantly reduced, exercise
343
could continue. It is clear therefore that some linkage
or dependency exists between the power demanded at
the point of exhaustion and the residual unused fuel
supply available at that time. Thus while it may be
reasonable to assert that the anaerobic capacity is of
a xed and limited value, the usable amount clearly is
not. Consequently any estimates of this value based
on the CP model assumptions will obviously be
underestimates.
5. Aerobic power is available at its limiting rate CP the
moment exercise begins and remains so right up until
the end of exercise. The kinetics of oxygen uptake,
whether regarded as mono- or bi-exponential and
with or without any delay(s) (Barstow and Mole
1991), clearly dictate that the establishment of the
aerobic supply at any given rate (not just at CP) is not
instantaneous. Three minutes or so may be required.
What this means is that the capacity-limited anaero-
bic supply as reasoned in the CP model must be in
reality larger than the model prescribes. Again, as
above, AWC must be underestimated by the CP
model.
6. The power domain over which the model applies is all of
CP<P<. If P CP, the anaerobic supply is never
required, since under assumption 5, aerobic power at
CP (or less) is immediately available. The left-hand
inequality therefore is obviously reasonable. At the
other end of the scale, human muscle is not all-
powerful and, as indicated above, a nite upper limit
to the power that can be developed must exist and to
the velocity of running, etc. This upper limit could be
denoted P
max
and regarded as a maximum instan-
taneous power.
7. The time domain over which the model applies is all of
0<t<, and endurance at CP is innitely long. The
notion of innite endurance is a somewhat nebu-
lous one. It is not uncommon that long-duration
continuous exercises of moderate power output are
terminated for a variety of non-energy-substrate re-
lated reasons, such as boredom, psychological issues,
the necessity to eat, drink or toilet. This therefore
places an imprecise upper limit to the time domain
over which the CP model applies. In similar vein very
short time intervals at very high power (even if less
than P
max
) are not realistic either. Factors like over-
coming inertia and acceleration come into consider-
ation. Strictly speaking of course, if exercise at P
max
or more was to be attempted, endurance time should
be zero.
8. The eciency of transformation of metabolic energy to
mechanical energy is constant across the whole power
(and time) domain(s). This assumption, implying
that the energy cost of locomotion per unit distance is
constant, seems to depend critically on the form of
locomotion (di Prampero 1999). It seems a reason-
able assumption for running and cycle ergometry (at
least within the sorts of bounds discussed at various
places above), but it changes substantially with speed
in swimming and cycling on the road or track. If the
form of such dependency was known, it would need
to be explicitly introduced into the model. While this
could be done, the result would be cumbersome.
9. CP and AWC are constants, independent of P (and/or
of t). The constancy (or otherwise) of these two
parameters of the model introduces issues of inter-
pretation. For example both CP and AWC are not
constant over time with respect to changes in exer-
cising habits (training in particular is discussed in
Interventions and their impact on the model above)
and, in any event, are nevertheless probably subject
to the usual intra-subject variations (other factors
being equal) typically inherent in any biological
organism. In another sense, Bishop et al. (1998) have
shown that the estimates of CP and AWC dier when
diering sets of P are used in the testing sessions.
While in some sense this non-constancy might seem
disturbing, relative stability of CP and AWC does
seem reasonable over a short term and indeed with-
out such an assumption not much progress can be
made.
Towards a more realistic critical power model
Bearing in mind the many deciencies in the assump-
tions as described above, the CP model can nevertheless
be improved without a great deal of added complication.
In several instances this has been done.
For example, Wilkie (1980) focussed on assumption
5, introducing a correction factor for oxygen uptake
kinetics based on a single exponential with time constant
s and without delay. Wilkies version of Eq. 1a now
becomes:
W a bt bs(1 e
t=s
), 2a
where a and b correspond to AWC and CP, respectively,
and the third term represents the amount of energy re-
leased from anaerobic sources before the attainment of
an aerobic steady state at CP. This factor is the degree
by which AWC underestimates the true anaerobic
capacity and makes up for the dierence in energy terms
between the assumed innitely fast (square wave) VO
2
kinetics of the CP model and real VO
2
kinetics with a
time constant equal to s. Wilkie also presents a version
of Eq. 1c with a corresponding correction term, but
derivations of corresponding equations 2a to 1b, 1d, 1e
and 1f are mathematically not straightforward.
The bioenergetics of Wilkies correction can be
demonstrated by Fig. 2. Here the real anaerobic
capacity is not a as it would have been under the CP
model, but a plus the approximately triangular area
below the dotted line and above the curve (the correction
factor in Eq. 2a).
Wilkie showed his correction to be appropriate for
times to exhaustion between 50 s and 10 min, giving s a
value of about 10 s. Although 10 s is rather too fast
(30 s might be more reasonable), this term does adjust
344
appropriately for oxygen kinetics in exercise of fairly
short duration. For exercise of longer duration, its eect
is relatively lessened.
Wilkies correction can be modelled by a simple
change to the hydraulic model presented in Fig. 1. All
that is required is to lower the vessel A
e
and its con-
necting tube R
1
such that when A
n
is full, the uid levels
in A
n
and A
e
are in alignment (Fig. 3)
Following Conceptualisation as a hydraulic model,
this model operates as follows. When T is opened to
some value P>0 the level in A
n
immediately begins to
drop, creating a dierential in levels between A
n
and A
e
.
This dierential induces a ow through R
1
directly
proportional to h, corresponding to a rise in VO
2
above
rest. This ow in turn slows the rate at which the level in
A
n
drops, slowing the rate at which VO
2
began to rise. If
P CP, then h and VO
2
will reach a steady state with
the level in A
n
no lower than the connecting tube R
1
.
That is, the height h is exactly sucient to induce a ow
through R
1
exactly equalling the ow P through T.
Mathematically, the ow through R
1
, and hence VO
2
,
can be shown to be mono-exponential, whether or not a
steady state is reached.
Of course if P>CP, h will increase beyond the depth
of vessel A
e
, the ow through R
1
will have reached its
limit at CP, and A
n
will continue to empty in order to
sustain the ow P>CP. In the model context, exhaus-
tion occurs when A
n
is fully depleted as P can no longer
be sustained. Mathematically, this can be shown to be
precisely the Wilkie formulation of the CP model.
While other corrections may be feasible focussing on
any one or more of the problems associated with the
original CP model, probably the most important from a
physiological point of view is the way in which AWC
appears to not necessarily all be usable.
A feedback control system and the 3-parameter model
It is well recognised that the last 100 m of a longer race
(say 5,000 m) cannot be run at the same speed as that
which could be achieved had the runner been sprinting
100 m afresh. Taking this in conjunction with the work
of Saltin and Karlsson (1971), there appears to be some
form of control or feedback system at work, which
operates to limit the maximum power output achievable
inversely according to how fresh the runner is or more
precisely according to how much energy store remains.
Therefore we can retain assumption 2 and modify
assumption 6 such that CP<P<P
max
and introduce a
simple linear feedback control system in accordance
with the amount of the A
n
store remaining.
This means that the maximum achievable power at
any time becomes a function of the amount of A
n
, spe-
cically:
CP + A
n
(P
max
CP)=AWC. 3
If the individual is fully rested and replete,
A
n
= AWC and P
max
can be achieved. On the contrary
if A
n
is empty then only CP can be achieved, and as
before the notion of exhaustion in vacuous unless
P>CP. This assumption of a linear control feedback
forms the basis of the 3-parameter CP model and is fully
described by Morton (1996).
It is immediately apparent that this model avoids, or
at least mitigates, the problems associated with
assumptions 3, 4 and 6. Research at the time (Gaesser
et al. 1995; Morton et al. 1997) and subsequently for
example Bull et al. (2000) and Hill et al. (2003) suggest
that AWC estimates are higher and CP estimates lower
than those obtained from the 2-parameter model, and
that endurance at CP (while not innite) is signicantly
longer than previously (assumption 7).
Diagrammatically, the 3-parameter model can be
represented by the hydraulic model of Fig. 1, but with
the addition of an appropriate linkage between the uid
level in A
n
and the tap T in accordance with Eq. 3
above.
Further developments
The most obvious development of the above discussion
is to combine Wilkies correction with the feedback
Fig. 3 Wilkies correction to the CP hydraulic model
Fig. 2 Wilkies correction to the CP model
345
assumption of the 3-parameter CP model. The details of
this combination have not previously been published, so
it can be achieved quite simply as follows.
di Prampero (1999) gives a version of Wilkies cor-
rection as his Eq. 3 on p. 163, which since W = Pt can
be adapted to
A
n
P CPt CPs1 e
t=s

and when combined with Eq. 3 above produces

CP P
max
CPAWC P CP
t CPs1 e
t=s
=AWC P
in the manner described by Morton (1996), where t is the
time to exhaustion.
This equation is transcendental (t occurs both linearly
and in an exponent), so cannot be solved explicitly for t.
Numerical methods need to be employed. However,
since Wilkie asserts s is about 10 s, and even if it is a
more realistic 30 s, then provided t is much greater than
s (by a factor of say 4, which it usually is), the exponent
can be neglected to a good order of agreement.
Neglecting the exponential term and solving for t
produces the following equation for time to exhaustion:
t AWC=(P CP) AWC=(CPP
max
) CPs(P CP).
Here we note that if VO
2
kinetics is assumed innitely
fast (a square wave), then s=0 and Eq. 4 reverts exactly
to the 3-parameter model equation. In addition, we note
that Wilkies correction depends, as indeed it should, on
the feedback property of the extent by which P exceeds
CP, i.e. more so if P greatly exceeds CP, precisely when t
is short.
A dierent development of the original CP model,
adapted for running, is due to Keller (1973), who poses
the question of an optimal running velocity in order to
cover any given distance (from 50 to 10,000 m and
corresponding time 6 s to 30 min). This treatise is quite
mathematical, requiring the calculus of variations, but
its results are straightforward and logical. Keller makes
no adjustments to the original CP model, but does allow
for the work required to accelerate the body in the early
stages of a race (a point mentioned earlier). Interest-
ingly, the optimal solution is in two parts. For races less
than about 291 m, an all-out sprint is optimal. For
longer races an all-out sprint is required only up to a
certain velocity (lower for longer distances), followed by
constant maintenance of this velocity almost to the very
end of the race (eectively to when A
n
is emptied), n-
ishing with the briefest collapse over the nish line
where CP and residual momentum are the only driving
forces.
The general notion that constant pace throughout a
race represents the duration-minimising strategy (as
distinct from a winning one) is widely accepted. While
this appears to be but one among countless such optimal
strategies for the 2-parameter CP model (Performance
implications), it appears to be sub-optimal for the 3-
parameter model. It has been shown (Morton 2004) that
under the linear feedback system assumed for the 3-
parameter model, an all-out eort is the duration-
minimising strategy. Consequently it can be conjectured
that for any linear feedback of the type in which the
maximum power attainable is linearly dependant on the
amount of fuel source remaining, an all-out strategy
would be optimal. Observers of human performance and
athletes themselves will surely be sceptical. Thus al-
though the 3-parameter CP model has certain advanta-
ges over the original CP model, it still has properties
and/or assumptions open to further questioning.
More detailed hydraulic models
Assumption 1, as indicated in Towards a more realistic
critical power model, is one which though appropriate
from a broad biochemical pathway point of view, is
metabolically inadequate. There should be a method of
incorporating the aerobic, alactic anaerobic and lactic
anaerobic pathways into a model scheme. This section is
devoted to showing how this has been done, elucidating
some of the properties of these models and considering
their further development.
Margarias seminal ideas
It was Margaria (1976) who rst proposed a 3-compo-
nent hydraulic model of human bioenergetics. It took
the following form (Fig. 4).
The uid in vessel P (representing the alactic anaer-
obic phosphagens) is directly connected with the outside
through the tap T, which regulates the ow (net energy
expenditure above rest). At rest, with T closed, the upper
level of uid in P is the same as in the communicating
vessel O (representing the oxidative or aerobic source).
The vessel O is of innite capacity and is connected
through tube R
1
. The second communicating vessel L
(representing the lactic or glycolytic anaerobic source) is
of nite capacity, with upper level the same as the bot-
tom level of vessel O, apart from a very narrow exten-
sion tube, B. The uid in B, corresponding to the resting
blood lactic acid, is of very small volume relative to L
and does not contribute to the ows to any signicant
degree. It is, however, measurable and corresponds to
the so-called early lactate which is produced at the
onset of even mild aerobic exercise. L is connected to P
through a wider, but one-way tube R
2
, and P is con-
nected back to L by another, but very much smaller one-
way tube, R
3
.
If T is partly opened, corresponding to a workload
W, the level in P falls, inducing a ow through R
1
(oxygen consumption, VO
2
) in accordance with the
dierence in levels, h, between the two vessels. This
induced ow slows the rate at which the level in P falls
346
and, provided W is not too large, equilibrium will be
reached at a level above the outlet R
1
. The level in P is
below the resting level, and uid ows continuously
from O to P and out through T. If the equilibrium
level is exactly at the level of R
1
, then the oxidative
mechanism is at its maximum, denoted VO
2max
. Once
the equilibrium is established, the only energy mecha-
nism contributing is the oxidative; the exercise is
purely aerobic and, in theory, could continue inde-
nitely. Prior to equilibrium of course, P has contrib-
uted some of its supply, and the empty volume in P
above the equilibrium level is known as the alactic
oxygen debt.
If T is now closed, i.e. exercise ceases, P will begin to
rell through R
1
, but at a slower and slower rate as the
level in P returns to normal. When it does so, the ow in
R
1
ceases and the subject is said to have repaid his
oxygen debt during this recovery period.
If T had been widely opened (severe exercise), the
initial situation would be as described above, but the
level in P would fall below R
1
. This happens after about
50% of the uid in P has been utilised, and the subject is
said to have crossed his anaerobic threshold. As soon as
this happens, two things occur: the ow in R
1
has
reached and continues at its maximum determined only
by the height of the vessel O and a ow through R
2
is
induced. This ow is also in accordance with the dif-
ference in levels between vessels L and P (the level in L
lagging behind the level in P). The ow through R
2
will
slow the fall of level in P, but since the ow through R
1
is insucient and the capacity in L is limited, the levels
in both L and P will continue to fall. If exercise is pro-
longed, L and P will be emptied and the subject will be
exhausted.
If T is closed at or before exhaustion, P will again be
relled. Initially it will be lled both through R
1
at the
maximal rate and through R
2
until the lag in levels
between L and P has been eliminated. This latter ow
through R
2
is a delayed lactic acid formation, which has
been observed to occur for a short period after cessation
of exercise (di Prampero et al. 1973). Once the levels
have been equated, P will ll through R
1
, initially at the
maximal rate and thereafter at a progressively slower
rate as described previously. L will be relled from P
through R
3
at a rate in accordance with the dierence in
levels between the two. As R
3
is so small, the level in L
will lag behind the level in P and the repayment of this,
the lactic oxygen debt, is very slow. Finally both P and L
are relled and the subject has fully recovered.
Although Margarias model was not quantied, suf-
cient information was available to do so. No mathe-
matical solution was oered therefore, though a
purported graphical solution (Margarias Figs. 1.27 and
1.28) was presented. A detailed examination of, and
mathematical solution to, Margarias model has been
undertaken from several perspectives (Morton 1984,
1985a, b, 1986a, b). This reveals that the actual solution
diers from both Margarias sketch solution and from
what actually happens under experimental conditions.
Thus although Margarias model clearly represents a
signicant step forward, the observed discrepancies are
such that it must be rejected in its original form and
reformulated in an attempt to alleviate the discrepancies.
This has been done by Morton (1986a, b) and has led to
development of the 3-component MM model.
The 3-component hydraulic model
It is clear from Fig. 4 that many dierent congurations
of the hydraulic model are possible depending on the
various heights of the tops and/or bottoms of the vessels,
relative locations of the connecting tubes, etc. Morton
(1986a, b) argues that only 4 of 16 are consistent with
Fig. 4 Margarias hydraulic
bioenergetic model
347
known physiological facts and later (Morton 1990) re-
duces the number to one unique conguration shown
below in Fig. 5.
The operation of this conguration can be deduced
from Fig. 4 and the description in Margarias seminal
ideas, with appropriate amendments, and is described
fully by Morton (1986a, b, 1990). Nevertheless it is
worth noting some of its salient features.
The top of vessel A
n
L, apart from the very narrow
tube B, is below the level of uid in vessel O, but above
the inlet tube R
1
(as is the bottom of vessel A
n
L). This
means that the anaerobic threshold (as signalled by the
commencement of lactic acid production represented by
a ow through R
2
) does not commence until VO
2
reaches some value above rest and clearly below VO
2max
,
say at 40% of VO
2max
.
This commencement of ow through R
2
changes the
dynamics of the ow through R
1
, i.e. changes VO
2
dynamics at this point, as described in some detail by
Cerretelli and di Prampero (1987). This elbow can be
deduced from a breath-by-breath VO
2
plot and is the
commencement of the VO
2
slow component (Gaesser
and Poole 1996). In other words, the crossing of the
anaerobic threshold can be modelled using the hydraulic
model (Morton and Gass 1987). What happens is that at
this delay point VO
2
dynamics change from mono-
exponential to bi-exponential as has since been empiri-
cally veried by Barstow and Mole (1991). The
hydraulic model formulation diers from the Barstow
and Mole formulation only in that the former proposes
a smooth transition across the threshold, whereas the
latter does not.
A steady state may or may not be reached thereafter
depending on whether W is so large as to drive the level
in A
n
A below R
2
. If it does then A
n
L will ultimately
empty and exhaustion is sure to occur. The maximal
aerobic steady state is clearly the point where h is exactly
opposite to R
2
and with A
n
L empty, the ow through R
1
exactly equals P. This is the true maximal sustainable
aerobic power and is clearly less than VO
2
max. Con-
sidering Fig. 5, this may be at 85% of VO
2max
, say.
Of course if T is closed, the system rells, initially
through R
1
and R
2
, then through R
1
and R
3
. The
reversal of the ow from R
2
to R
3
suggests that recovery
VO
2
kinetics has two bi-exponential phases, a phenom-
enon which does not seem to have been examined in
detail. Since R
3
is relatively so narrow, A
n
L takes a
considerable time to rell, during the early stages of
which A
n
A is almost completely lled. Thus the model
predicts that for some long time after cessation of
exercise, VO
2
would not in fact return fully to the resting
level, but remain just above it. Indeed this phenomenon
was described by Hill in the 1920s and is further re-
viewed in di Prampero (1981). This phenomenon is akin
to the well-known post-prandial minor elevation in VO
2
.
It is a quite straightforward exercise to programme a
computer with the equations of this hydraulic model.
For any specied time course of W, running the pro-
gramme would simulate VO
2
, phosphagen depletion,
lactic acid production (or glycogen depletion) and the
replenishment thereof if W reduces signicantly or takes
the value zero. Given the right scaling constants and
osets for resting values, quite realistic smooth data
can be generated.
What the model of course does not incorporate is any
criterion for exhaustion, other than A
n
L emptying when
VO
2
exceeds MAP. There is no feedback system of the
type described in A feedback control system and the
3-parameter model.
Feedback control systems, maximal and optimal
performance
Since the 3-component hydraulic model has two anaer-
obic vessels which partially empty during exercise, there
are two fuel starvation feedback systems possible.
Morton (1990) has investigated both. A phosphagen-
based limitation does not lead to the right sort of time
Fig. 5 The generalised MM
hydraulic model
348
course for all-out power decline such as in Wingate test
or prolonged sprint, despite the observation that chan-
ges in these muscle metabolites do tend to correlate with
power decline. Much better correspondence occurs if the
maximum attainable power is assumed directly propor-
tional to the remaining glycogen store. The properties
and predictions of the MM model under this assump-
tion are examined in detail by Morton (1990).
All-out power is constant at its maximum (governed
in the model by the diameter of tap T when open) for
only a few seconds. This is the time taken for the level in
A
n
A to drop to the top level of vessel A
n
L, during which
of course there is no ow through R
2
. Then only does
the level in A
n
L begin to drop and the feedback begins
to operate. Using published data for t males, all-out
power declines after about 6 s at 972 W to very low
levels after about 2 min. Plenty of data are available to
compare this prediction for eorts of 30 s or so, but data
lasting 2 min or more appear to be sparse. Very well
motivated subjects would be needed to provide it.
Endurance at constant power declines from ad in-
nitum at 208 W (about 89% VO
2
max) to 6 s at 972 W
in a hyperbolic shaped curve (actually it is the solu-
tion to a ratio between two bi-exponentials). At VO
2max
an endurance of around 9 min is predicted. The model
also allows investigation of endurance at ramp tests of
various incremental rates. For rates between 30 and
60 W/min, endurance reduces from 14 to 9 min, and the
anaerobic threshold is reached in times between 6 and
3 min. Interestingly, in these ramp tests the model pre-
dicts that power output at exhaustion (peak power) in-
creases with increasing ramp slope, but terminal oxygen
consumption (VO
2
peak) is unaected.
Contrarily, the MM model has not yet been inves-
tigated to determine any optimal strategy for completing
a given amount of work in minimum time (or running a
xed distance in minimum time). It is conjectured (as an
inference from the work by Morton (2004) reported in
Further developments) that this feedback system could
lead to implicating an all-out strategy. However, it
should be noted that the form of the original glycogen-
based limitation [Eq. 7 in Morton (1990)] is slightly
dierent from the form of feedback Eq. 3 above, and so
this implication is by no means assured.
Applications and developments of the 3-component
hydraulic model
One obvious application of the MM model is its use as
a teaching and learning tool to give a macroscopic
overview of the operation of the human bioenergetic
system during exercise and recovery. At the simplest
level the concepts only can be used, since understanding
the ows of a uid between interconnecting vessels is
quite intuitive. Indeed I believe this to be the motivation
behind Margarias original 3-component model. The
analogies between the hydraulic systems attributes
(capacities, ow rates, height dierentials, etc.) and the
corresponding human bioenergetic attributes (energy
stores, power, etc.) are also most intuitive. While not
absolutely realistic in its properties, the MM model
presents a signicant advance on the earlier described
CP model.
At an intermediate level, graphical illustrations of
changes in various components of the system (for
example of the ow through R
1
, analogous to VO
2
) can
provide more insight. These can be enhanced by the
selection of corresponding real data sets to verify the
accuracy (at least to a reasonable degree) of the work-
ings of the model.
At the highest level, those with appropriate knowl-
edge of ordinary dierential equations can gain an even
deeper comprehension by working through the mathe-
matics to obtain solutions to the system under a variety
of conditions.
Taking this application one step further and into the
research context, the model provides a means of
answering a wide variety of what if questions. The
resulting model predictions can then be checked against
appropriate existing data or data from suitably designed
experiments. Just how many of the MM model pre-
dictions conform to reality or not remains to be seen.
For example, as already observed, the bi-exponential
nature of the model prediction for VO
2
for constant
power exercise above the anaerobic threshold is now
clearly established (Barstow and Mole 1991).
For ramp exercise, the MM model predicts a fast
exponential trend towards linearity of VO
2
, but only up
until the AT, followed by a second exponential trend
towards lesser sloped linearity until exhaustion.
Unequivocal verication of this has not yet been estab-
lished, for available data seems to be contradictory
(Morton and Bocquet 2003).
As a further example, glycogen loading obviously
enlarges the capacity of vessel A
n
L, but whether this is
accomplished by increasing the cross-sectional area and/
or the depth (either by raising the top or lowering the
bottom or both) is unclear. Changes in VO
2
dynamics
forecast by the model as a result of such structural
change may provide clues.
Behncke (1993) has incorporated the MM model as
one submodel in a larger mathematical model for the
force and energetics of competitive running. The other
submodels include the biomechanics of running and an
optimisation determination. Behncke modies the MM
model rst by regarding vessel O as nite, though very
large, because glycogen depletion and fatigue are serious
problems for long-distance events like 10 km or mara-
thon runs. As a second modication, vessel O is con-
sidered as the union of two vessels, the smaller and
uppermost having lesser cross-sectional area, although
both are large. This is done because even though it may
be reasonable to consider separate compartments for
glycogen and lipids, it cannot be done because
these processes are not additive as far as their power is
concerned. They run in parallel using the same substrate
(oxygen). As a third modication Behncke regards
349
maximal available power at any time to be a function of
the level of uid in vessel A
n
A, but only after the
anaerobic threshold is reached. Specically, maximum
available power declines linearly from P
max
to zero as
A
n
A empties beyond the anaerobic threshold.
However the time course for changes in maximum
available power starting from 0.85P
max
(Behnckes
Fig. 2, p. 865) does not entirely conform to the experi-
mental data described in Morton (1990).
The optimisation submodel is mathematically com-
plex, but in essence produces a strategy very similar to
that of Keller (1973) as described in Performance
implications. Constant power throughout most of the
race appears to negate the conjecture of all-out eort
suggested above. Such matters are clearly in need of a
more careful general solution.
Behncke (1997) then applies this model to world
running records of the time in order to determine key
parameters. The values so obtained were in good
agreement with those published in the literature and
those obtained by other means. The model is also ap-
plied to determine the velocity prole for a 100 m race
and used to compare the eects of adverse wind and
altitude. In as much as there are data available, Behncke
evidences good agreement.
Other bioenergetic models
A model based on the rst law of thermodynamics
Ward-Smith (1985a) proposed a theory of running
based on the rst law of thermodynamics. In confor-
mity with this law, the energy released by chemical
conversion, C, must ultimately be accounted for either
in the form of physical work, W, or degraded into
heat, H. The overall energy balance equation is
therefore
C H W
and by dierentiation with respect to time, a power
equation can be derived. Ward-Smith traces the origins
of his modelling back to Lloyd (1967) who proposed C
to consist of two terms: one expressing the energy
available from a store and the other expressing the en-
ergy available at a steady state. The similarity to
assumptions 13 of the CP model are immediately
apparent.
Of interest here are Ward-Smiths assumptions about
the derivative dC/dt, regarding the two terms as anaer-
obic and aerobic, respectively, based on Margarias
(1976) earlier work. Specically
dC=dt P
max
e
kt
R1 e
kt
;
where P
max
is the maximal anaerobic power, k a rate
parameter governing energy release and R the steady-
state maximum aerobic power (the critical power). There
aerobic power, P
aer
, given by the expression
R(1 e
kt
)
is the same as that previously referred to as Wilkies
correction, and the robust nature of this expression has
already been noted. Ward-Smith does argue (p. 341) that
the anaerobic energy store has two components, alactic
and glycolytic, but nevertheless combines them as one. A
separation is investigated in more recent papers (Ward-
Smith and Radford 2000a, b).
Where a departure from most earlier work is evident
is in the expression for anaerobic power, P
an
, given by
P
max
e
kt
rather than by the dierence between dC/dt (which
commonly is constant, but is by no means necessarily
so), as determined by the individual exerting the eort,
and P
aer
. What this means is that under the Ward-Smith
assumption dC/dt always begins at P
max,
declining
exponentially thereafter to an asymptotic value of R and
quite independent of the power requirement of the
exercise, constant or otherwise. This of course is not
necessarily so, for the athlete may choose to start at
some value higher than R, but less than P
max
. Further-
more if the expression for P
aer
is adopted, then P
an
must
be a function of both the power requirement of the
exercise and of t, not just of t alone. Closer investigation,
however, reveals that the departure is only an illusion,
for Ward-Smiths equations are mathematically equiv-
alent to those of Lloyd.
The feedback nature in the Ward-Smith model is the
hypothesis of Margaria that prior to the activation of
the glycolytic mechanism, the rate of energy release due
to oxidation is dependent on the amount of split
phosphagen. Overall, the bioenergetic components of
the Ward-Smith model are therefore a compilation of
prior model themes.
Where there is originality is the manner in which the
biomechanical equations of running are linked to the
bioenergetics, which is the incorporation of running
resistance and the thermodynamic equation for increase
of thermal energy. When all is said and done, the Ward-
Smith model predictions (given appropriate selection of
parameter values) conform very well with actual running
data from real athletes. Indeed the robust nature of
Ward-Smiths model has been further demonstrated in a
number of successful applications (Ward-Smith 1985a,
b, 1999a, b; Ward-Smith and Radford 2000a, b).
Other bioenergetic models
Another detailed and often quoted model is due to
Peronnet and Thibault (1989) which describes the aver-
age power output sustained over time t. It is based on a
2-component (aerobic and anaerobic) system, closely
related to the models described above, and based on the
work of Lloyd (1966) and Ward-Smith (1985a, b). The
aerobic component is again a single exponential analo-
gous to Wilkies correction as described previously.
350
However, Peronnet and Thibault argue that the
maximal aerobic power can only be sustained for a nite
length of time (about 7 min) and that the fraction of
MAP that can be sustained for longer times than this
declines linearly with the natural logarithm of t, ln(t).
The amplitude of the aerobic component is therefore
reduced accordingly for times in excess of 7 min.
In the case of the anaerobic component, Peronnet
and Thibault adopt essentially the same assumption as
Ward-Smith that the available anaerobic component
contribution declines exponentially with t (again with
the same time constant as the aerobic component) for
any power output, provided it is sucient to lead to
exhaustion. Peronnet and Thibault recognise that not all
the anaerobic store is necessarily available for races of
any duration. They argue that all is available for races of
up to 7 min duration, but that for longer races the
availability decreases with ln(t).
This last assumption represents a signicant contrast
to one important property of the 3-parameter CP model,
where availability of AWC increases more and more
with exercise lasting longer and longer. Nevertheless,
Peronnet and Thibault evidence very good ts of their
model to mens world records of the time from 60 m to
the marathon.
di Prampero et al. (1993) published a model to predict
running performances rather similar to that of Peronnet
and Thibault, showing good agreement between actual
and calculated best times over distances from 800 to
5,000 m, not only for their own data, but also for
independent data of Lacour et al. (1990). A summary of
this and related studies can be found in di Prampero
(2003).
Olds et al. (1993) incorporate a bioenergetic compo-
nent into their modelling of road cycling performance.
This component consists of a xed anaerobic contribu-
tion, referred to as the maximal accumulated oxygen
decit (their equivalent to AWC), together with a vari-
able aerobic contribution. In the manner of Peronnet
and Thibault, this aerobic contribution increases mono-
exponentially at exercise onset and reaches a steady state
at some fraction of VO
2max
depending on the duration
of exercise. The only dierence is that whereas Peronnet
and Thibault regard this fraction as decreasing linearly
with ln(t), Olds et al. regard it as unity for times
10 min, equal to the individual anaerobic threshold
for times 30 min, and as declining linearly with t be-
tween the two. Empirically derived eld and laboratory
data were used to assess the accuracy of the model,
yielding mean (observedpredicted) dierences of 1.3%,
very good, but marginally worse that the 0.73% of
Peronnet and Thibault.
Further application of this model investigated the
impact of a number of biophysical, environmental and
physiological variables on cycling performance (Olds
et al. 1995). The most important physiological factors
were found tobe VO
2max
, fractional utilisation of VO
2max
,
mechanical eciency and projected frontal area. A
number of practical issues, such as the eect of drafting
and the normalisation of performances under dierent
conditions, were also considered. Maronski (1994) also
considers an optimising problem in cycling, and a feed-
back mechanism limits force development according to
the cyclists velocity. In a more recent reviewof modelling
human locomotion, Olds (2001) makes a further modi-
cation to the modelling of the anaerobic contribution, by
asserting (in the manner of Ward-Smith) that the oxygen
decit may not all be utilised by the end of exercise. In
other words, it is regarded as declining exponentially with
a xed time constant, once again independent of the
power requirement of the exercise.
In the more complex (at least for physiologists) ap-
plied mathematical literature can also be found some
models appropriate to mention. These can be viewed
alongside the Keller (1973) model, as most seek to ob-
tain optimal strategies in some way. Like Keller, Cooper
(1990) adopts no feedback system to a model for opti-
mising wheelchair propulsion. On the contrary, Mathis
(1989) adapts Kellers model by introducing a feedback
due to a fatigue eect in such a way that the propulsive
force available to the runner declines linearly with the
oxygen decit of the runner. The similarity with the
feedback mechanism of the 3-parameter critical power
model is immediately apparent. Of interest is that this
dependency leads to an optimal solution in which the
constant velocity segment is run at a slightly lower pace
and which includes a period of perceptible acceleration
over the last 1040 s of the race. The slower pace is to
ensure that the oxygen decit does not become too high,
and the stronger the dependency the lower the velocity
and the shorter the duration of the nal sprint. As far as
I can determine, this is the only application where the
mathematically optimal strategy includes a nal sprint.
Readers will no doubt take note that this is an extremely
common strategy in real races. Woodside (1991) adopts
a similar approach, using a feedback fatigue constant
which relates the rate of energy loss due to fatigue to the
energy already spent in running. This analysis does not
result in a nishing sprint.
Both Tibshirani (1997) and Mureika (1997) follow
Mathis example in that the propulsive force during
sprinting declines linearly, but with t rather than with
the oxygen decit. As it turns out, this dierence is of
little consequence, evidencing just as good ts to data as
does Mathis.
Harman (2002) modies the Ward-Smith and Rad-
ford formulation to include three anaerobic compo-
nents: ATP, phosphagen and glycolysis. Each
component is regarded as bi-exponential in nature with
its own rise and fall rate parameters. In this model the
limited ATP supply is largely depleted within the rst
few seconds, albeit that in reality ATP concentration is
essentially constant (though maybe low) throughout the
exercise intensity range because of the equilibrium con-
stant of the creatine kinase reaction. In this model all
three anaerobic mechanisms have peaked within 10 s.
Nevertheless, as with many of the models described
previously, very good ts to real data are evidenced.
351
Conclusions and further considerations
I have traced the development of whole-body bioener-
getic models over the last 50 years, from the early simple
2-component ideas of Monod and Scherrer (1965) and
Lloyd (1966) and others; through Margaria (1976) and
Mortons (1986a, b) 3-component extensions; to the
more complex recent models of Behncke (1993, 1997),
Ward-Smith (1985a, b) and Harman (2002). It is useful
now to view the current situation in summary with ref-
erence back to the ten assumptions of the original CP
model (Origins, assumptions and development and
Questioning the assumptions).
It would seem that signicant progress has been made
on all counts, simply by an evolutionary development of
existing ideas. Indeed such is the nature of science and the
advancement of knowledge. Nevertheless it appears that
there is at present no single published model which
incorporates all the developments. Herein lies a challenge.
One issue of note where it appears there has been no
development is with respect to assumption 6 (and to a
certain extent 7 also). Why should the bioenergetic
models available so far and surveyed herein apply only
to/or have been tested only against) exercises of a
maximal nature or which ultimately lead to exhaustion?
Why should they not equally be applied to and tested
against data from submaximal work which does not
necessarily lead to exhaustion. Herein lies a second
challenge.
A second issue of note is that a fundamental char-
acteristic of the bioenergetic process is that it is demand
driven by a single element, the power output required. It
is supply limited obviously, though this may not neces-
sarily be the only feedback mechanism. However, it is
clear that all amplitude parameters must depend on the
demand (if power is high, VO
2
will be high, etc.) and all
rate parameters depend on the structure (i.e. capacities,
maximal rates, etc.) of the system. No single model so
far presented is of this character, though the work of
Behncke comes closest. Herein lies a third challenge.
Finally, the single most remarkable conclusion is on
the one hand such a plethora of models t the real world
so well, yet there is so much more to discover.
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