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DOI 10.1007/s00421-009-1008-7
INVITED REVIEW
Introduction
The study of energy consumption in cycling has a long
history. Many factors affecting energy consumption have
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123
work
power
power
:
metabolic rate rest metabolic rate
123
Energy influx
internal
elastic
energy
neg. internal
work
external work
internal
movement
energy
neg. internal
work
pos. internal
work
muscle contraction
muscle contraction
pos internal work
External work
Fig. 1 Conceptual model for energy flow during exercise. The terms
positive (pos) and negative (neg) indicate direction of energy flow
only. One of the main issues is that internal work, as described in the
literature, cannot be assumed to be loss of energy. Further, every
energy conversion (arrow) implies energy loss (as heat)
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as discussed previously, namely that this approach presumes two independent energy flows. Furthermore, by
measuring the energy cost of the unloaded movement, not
only is all internal work dissipated into heat, but it must be
dissipated into heat because the circumstances do not allow
external work production. There is no reason to believe
that in the case where external work can be done, the same
internal work is not (partly) converted to external work.
Moreover, it appears that by moving the lower extremities
passively, i.e. by external forces, metabolic rate increases
significantly (Nobrega et al. 1994). This indicates that
other processes than simply the active limb movements
also affect metabolic rate. Unpublished data from our
laboratory indicate that muscle activity in unloaded cycling
is extremely low and can hardly account for the total
increase of metabolic rate that is usually observed in
unloaded cycling (e.g., about 200450 W energy consumption at 0 external work rate, at 60120 rpm, Hagberg
et al. 1981; Sidossis et al. 1992). Two other processes that
at first sight are obvious candidates are the enhanced
heartand ventilation rate. However, there is ample evidence that these processes require comparatively little
additional energy consumption (e.g., McGregor and
Becklake 1961; Aaron et al. 1992; Kitamura et al. 1972)
and thus can hardly explain this phenomenon. Furthermore, doing more ineffective work due to coordination
challenges (Neptune and Herzog 1999) likely enhances
metabolic rate in passive cycling. This was substantiated
by Bell et al. (2003), who found considerable muscle
activity and a coinciding increase in metabolic rate during
pure passive cycling compared to other modes of passive
leg movements. In that study, subjects reported it was
difficult to relax completely in passive cycling. Thus, it
seems difficult to experimentally determine the energy cost
of true internal work (i.e., work that never appears as
external) in loaded cycling.
Another method to determine the costs of body segments movement energy changes is by extrapolating the
relationship between external work rate and energy cost to
a zero work rate (e.g., Hintzy-Cloutier et al. 2003). Such
an approach requires that the several work rates that are
used entail the same body segments movement energy.
In cycling, this requirement is fulfilled by using the same
cadence. Furthermore, it is expected that the energy
costwork rate relationship is linear, which has been
substantiated empirically in many studies (e.g., Bijker
et al. 2001, 2002; Chavarren and Calbet 1999; HintzyCloutier et al. 2003; Widrick et al. 1992; see also the
literature results gathered in this review, Fig. 2), although
it should be noted that for sustained work rates that
exceed the lactate threshold this relationship may become
nonlinear because of the influence of the so-called slow
component of oxygen uptake (e.g., Poole et al. 1994;
Dpower
;
Dmetabolic rate
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a
Gross efficiency (%)
30
25
**
* *
20
15
*
**
*
*
10
25
20
Luthanen et al., (1987)
15
10
5
30
45
60
75
90
105
120
100
Cadence (rpm)
2500
30
2000
1500
Samozino et al. (2006)
Moseley and Jeukendrup (2001)
Luhtanen et al. (1987)
1000
500
200
300
400
500
400
500
400
500
25
20
15
10
5
0
100
200
300
400
500
100
30
e
2000
200
300
1500
1000
120
500
25
20
15
10
60
5
0
100
200
300
400
500
123
100
200
300
Efficiency in cycling
Here, we are concerned with the literature on efficiency in
cycling, particularly regarding the influence of work rate
and cadence. This paper does not aim to discuss performance enhancement in competitive cycling by optimisation
for energy expenditure in cycling. Nevertheless, it appears
that one tends to freely choose a pedalling rate that is
somewhat above the energetically optimal one (e.g., Foss
and Hallen 2005). Some authors argue that this hampers
performance (e.g., Foss and Hallen 2005; Hansen and
Ohnstad 2008). Others use this finding to argue that the
human body apparently does not care about minimising
energy expenditure (e.g., Redfield and Hull 1986), or
consider other optimisation criteria, such as muscle activation (e.g., Neptune and Hull 1999). While we do not take
a stand on this issue in this paper, the findings summarised
here may be of relevance for its ultimate resolution.
Cadence and work rate
The two most obvious variables that may affect efficiency
are cadence and work rate. Cadence is often thought of a
rather simple and straightforward gear between force
(torque) and velocity (angular velocity) of muscle contraction. Thus, the energetically optimal cadence is likely
to be found at a muscle contraction speed that is close to
maximal power and efficiency in isolated muscle (i.e.,
around 0.3 of maximal force and contraction velocity; e.g.,
see Barclay et al. 1993). Kohler and Boutellier (2005)
argued that the freely chosen cadence may not follow the
principle of minimising energy cost because of the discrepancy between velocities giving maximal power and
efficiency. However, their analysis does not account for a
number of processes that are affected by cadence as well.
For example, because of activation-relaxation dynamics,
relatively more time is consumed at higher cadences by the
activation and relaxation process. Furthermore, inertial
effects by rotating lower limb masses lead to a change from
muscle performance to performance on the pedals (e.g.,
Ettema et al. 2009; Kautz and Hull 1993; Kautz and
Neptune 2002; Loras et al. 2009). Ettema et al. (2009)
demonstrated that details in cycling technique change with
cadence. In other words, the concept of treating choice of
cadence as a mere gearing between force and velocity of
muscle contraction may be attractive, but it probably does
not fully hold.
To summarize the extensive literature on efficiency in
cycling and the effect of work rate and cadence, we plotted
the results of a large (but certainly not complete) set of
studies that report gross efficiency in cycling. The studies
include untrained up to elite and professional cyclists
(including world top), different exercise protocols, and
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8
500
400
300
200
100
0
20
40
60
80
Cadence (rpm)
2200
2000
1800
1600
1400
1200
1000
800
600
400
200
0
0
25
Ca
50
den
ce
75
(rpm 100
)
125 0
100
200
300
400
500
wer (W)
External po
Fig. 3 a Cadence plotted against work rate for all data considered in
this overview (see Fig. 2). b Energy expenditure plotted against work
rate and cadence (same data as in Fig. 3a and Fig. 2e and f. Coloured
mesh is the two-dimensional linear regression (red marker indicates
extrapolated intercept at zero load and zero cadence). Markers
distinguish data below (filled) and above the mesh (open). The
regression is described by: Metabolic rate = 39.7 (29.3) ? 2.84
(0.34) 9 rpm ? 3.73 (0.08) 9 External Power
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Table 1 Average values of the reciprocal slope of work ratemetabolic rate relationship (RSep-mr) calculated from (and reported by) a
number of studies on cycling
Source
RSep-mr
22.2
26.2
24.4
17.8
25.5
21.5
25.0
21.6
22.1
25.4
27.6
23.4
Unpublishedelitea
22.7
Unpublishedtrainedb
27.4
Mean
23.8
Standard deviation
2.6
Cyclists from the national Norwegian team (time trial). Measurements done in same lab as (b)
b
Data are from the same study as Ettema et al. (2009) and Loras
et al. (2009), but not reported in these publications
Interestingly, the intercept of the two-dimensional regression at zero work rate and zero cadence (Fig. 3b), which
would be the theoretical value for energy expenditure while
sitting still on a bicycle, reaches a value of 40 W (not
statistically significant from zero). This value is too low,
but still physically possible, despite the rather large
extrapolation range from the experimental data. Overall, it
seems that the very original findings by Fenn (1924) on
isolated muscle also apply to the entire human body in
cycling in a very consistent manner.
In the literature data in Fig. 2c, some deviations appear: in
two cases the offset in metabolic rate is somewhat higher
(Chavarren and Calbet 1999; Samozino et al. 2006), and in a
few others the metabolic rate increases exponentially at high
work rates (Luhtanen et al. 1987; Moseley and Jeukendrup
2001; Moseley et al. 2004). The higher offset cannot be
explained, but the exponential increase may be because the
subjects exercised above lactate threshold and approached
their maximal work capacity. In such an instance, one may
expect that an increase in work rate requires a disproportional amount of metabolic input (e.g., because of
deterioration of coordination). In the case of Luhtanen et al.
(1987), where the highest three work rates were at and above
anaerobic threshold, this leads to a negative relationship
between gross efficiency and work rate (Fig. 2b). Note that
for all data presented in Fig. 2, the metabolic rate was based
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10
RSep-mr values of a number of studies that reported metabolic rate at various work rates. Most of these values are
not taken from the original papers, but calculated from the
data collected for this review. Thus, for all studies the same
algorithm was used. Some studies show substantially lower
reciprocal slopes than the average, but none much higher.
This is explained by the different weighting in the calculations of these studies: the studies with the higher slopes
have more data points. Thus, it seems more appropriate to
conclude that the RSep-mr as found in the literature averages
around 2324% rather than 26%.
Is there an energetically optimal cadence?
When considering overall effectiveness for the entire
human body (i.e. total energy cost in relation to external
power), most studies that looked at a rather wide range of
cadences and are represented in the analysis in this review
report that the lowest pedalling rate is most effective
(Chavarren and Calbet 1999; Gaesser and Brooks 1975;
Lucia et al. 2004; Nickleberry and Brooks,1996; Samozino
et al. 2006; Sidossis et al. 1992; Widrick et al. 1992). Two
studies claim the highest rate to be most effective, and 2
others an optimal cadence (Foss and Hallen 2005; own
unpublished data). Other studies (e.g., Coast et al. 1986)
also find an optimal cadence with regard to efficiency. In
most of these studies, the optimal cadence lies between 60
and 80 rpm. These contradictory results may, in part, be
explained by the interaction between work rate and
cadence. In short, when an optimal cadence is found, it
increases with work rate (Boning et al. 1984; Coast and
Welch 1985; Francescato et al. 1995; Foss and Hallen
2004; Gaesser and Brooks 1975; Seabury et al. 1977), and
the impact of cadence on efficiency seems most remarked
at lower work rates (Chavarren and Calbet 1999; Samozino
et al. 2006; Widrick et al. 1992). Also di Prampero (2000),
in his review, concluded that efficiency in cycling is
affected by cadence and the optimum by work rate.
Extrapolations from cadence studies to the forcevelocity relationship of muscle should, however, be made
with caution. Indeed, Hill (1934) warned against this in his
critical comments on Garry and Wishart (1931, 1934). Not
only muscle shortening velocity, but also activationrelaxation dynamics are strongly affected by cadence.
McDaniel et al. (2002) attempted to distinguish between
these two factors by manipulating crank length. They found
that pedal speed (marker for muscle shortening speed) and
power (work rate) were the main determinants for metabolic rate, not cadence (marker for activation-relaxation
dynamics). Marsh et al. (2000) found no effect of cadence
on delta efficiency, where values were around 2326%.
This does not contradict findings about optimal cadence
with regard to energetic cost. For the sake of argument, if
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one assumes that resting metabolic rate and the cost of limb
movements is (physically) independent of work rate, delta
efficiency is a measure of the increasing cost directly
linked to increasing muscle work. Marsh et al. (2000)
basically substantiated that the impact of increasing work
rate is independent of movement speed. Chavarren and
Calbet (1999), however, report a significant positive effect
of cadence on delta efficiency. Their work ratemetabolic
rate data are redrawn in Fig. 2e, inset. Although the trend
of a changing slope is evident, it is small considering the
range of cadence (eD = 21.5% at 60 rpm to near 24% at
120 rpm). On the other hand, Chavarren and Calbet (1999)
report a stronger negative effect of cadence on gross efficiency, indicating that we are not only dealing with
processes associated with work rate, but also other processes (e.g., force-velocity properties, activation-relaxation
dynamics, energy flow associated with internal work,
ventilation and circulation).
In early work, Dickinson (1929), but also Garry and
Wishart (1931, 1934), studied the relationship between
cadence and efficiency, basing their analysis on Hills
force-velocity equation. (Note that they expressed speed in
terms of time of contraction.) Dickinson (1929) established
efficiency for maximal efforts (i.e., well over the lactate
threshold, and with no steady state) by including oxygen
uptake during recovery. She subtracted resting metabolism
(i.e., calculated net efficiency). We calculated gross efficiencies from these original data, leading to values around
49%. These extremely low values occur because of the
long period of oxygen uptake measurement (30 min) in
relation to the exercise period (110 min), once more
clearly addressing the practical and theoretical challenges
around true efficiency of work production. Some of
Dickinsons original data are re-plotted in Fig. 2a against
cadence. The optimal frequency lies around 35 rpm (all
data to the left that are omitted were lower). This is rather
striking as these measurements were made at a high work
rate. On the other hand, the data are not out of the ordinary
compared to the more recent studies accounted for in Fig. 2.
Furthermore, Dickinson calculated net efficiency, assuming
a constant and work independent of resting metabolism.
An important consequence of the analysis of cadence and
work rate effects on efficiency is that power differences
may be a confounder in experimental studies on cadence. In
experimental testing, relatively small differences in external
power that are related to cadence may occur. These differences can have a relatively large impact on the cadenceefficiency relationship. Furthermore, relatively small errors
in power measurements affect the efficiency value considerably (see below, Fig. 2d). An error of 5% at 285 W
(14.25 W) gives an efficiency error of 1% (Fig. 2d). In other
words, it is important that possible systematic errors in
power linked to cadence are eliminated.
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Conclusion
We conclude this review by putting what was discussed
earlier into a simple theoretical framework. In the tradition
of Fenn (1924), one can factorise the metabolic costs as
found in cycling (Fig. 2): E = I ? kW, where E is metabolic costs, I is the constant intercept (maintenance), k a
constant (reciprocal delta efficiency), and W the external
work done. When using delta efficiency as a measure for
muscular efficiency, one assumes that the intercept of the
work rate-metabolic rate relationship is not associated with
muscular contraction and all energy increase is linked to
the work accomplished. This is, of course, a tempting
thought, but the physiological basis for it can be challenged; the only matter that is clearly established is a very
consistent linear work rate-metabolic rate relationship. The
equation is likely better rewritten as E = Ic ? k(Iw ? W),
with Iw = qW. In other words, total metabolic rate is built
up from a constant rate (Ic), a work related rate (kW), and a
component of energy consumption that is not directly
associated with the work conversion process (muscle
contraction) but changes linearly with it (Iw). These processes may be, for example, ventilation and circulation, but
also digestive processes. Energy loss associated with relative movements of segments (note once more, not the
entire internal work) would logically be accounted for by
the Ic component: losses associated with internal work do
not depend on external work rate, but more likely on
cadence (i.e., the amount of kinetic energy changes of the
moving limbs). Indeed, both Sidossis et al. (1992) and
Chavarren and Calbet (1999) clearly show that with
increasing cadence only Ic increases in a more or less linear
fashion (see Fig. 2e, inset). It is tempting, but likely
incorrect, to conclude that this increase is solely due to
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12
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