J. Phyeiol. (1972), 223, pp.

355-363 355
With 2 text-figure8
Printed in Great Britain

THE MECHANICAL
EFFICIENCY OF TREADMILL RUNNING AGAINST A
HORIZONTAL IMPEDING FORCE
BY B. B. LLOYD* AND R. M. ZACKSt
From the University Laboratory ofPhysiology, Parks Road,
Oxford, OX1 3PT
(Received 6 December 1971)
SUMMARY
1. A method for determining the apparent mechanical efficiency of
running against a horizontal impeding force is described. The results of
studies on three well trained athletes have been summarized.
2. A linear relation was found between metabolic rate and external
work rate. The apparent mechanical efficiency was calculated as the inverse
of the slope of the linear regression line of metabolic rate on external work
rate x 102 (percentage).
3. The over-all mean for apparent load-running efficiency (LRE) was
36x1 %. The differences in LRE between subjects were not statistically
significant. No measurable change in LRE occurred with habituation.
4. Values are given for E4, the net energy cost of unloaded horizontal
treadmill running per kilogram of bodyweight and per kilometre of dis-
tance covered. The over-all mean for Ek was 0-83 kcal. kg-'. km-'.
5. The efficiency values obtained are discussed and compared with other
values of running efficiency obtained by different methods, and with studies
on isolated muscle.

INTRODUCTION
A problem in the evaluation of the mechanical efficiency of most human
activities lies in obtaining an accurate estimate of the external work done
or in setting up a situation in which accurately measurable work is done.
In running over horizontal ground energy is used in producing changes in
the potential and kinetic energies of the body as a whole and of the indi-
vidual limbs, and in overcoming the resistance of the air. The energy used
for these processes is difficult to measure, and the assumptions involved
make the accuracy of the measurements questionable. In running on a
* Present address: Oxford Polytechnic, Headington, Oxford.
t Rhodes Scholar.
14 P HY 223
356 B. B. LLOYD AND R. M. ZACKS
horizontal treadmill the body as a whole is effectively stationary relative
to the surrounding air, so that no energy is needed for overcoming air
resistance. The need for evaluation of the potential and kinetic energy
changes in the body may be obviated by the imposition of an extra work
load by inclining the treadmill. The external work done in lifting the body
weight can be simply calculated, and measurement of the increase in
metabolic rate above that for horizontal running required for doing this
lifting work enables the mechanical efficiency of grade running to be
estimated-.
Confusion has arisen over the measurement of mechanical efficiency,
some of this difficulty being semantic. In this paper
net efficiency = external work rate
corresponding increase in metabolic rate
above resting metabolic rate
and
apparent efficiency =external work rate
corresponding increase in metabolic rate
above metabolic rate at zero load
Furusawa, Hill & Parkinson (1927) first attempted to measure the
energy used in sprint running by collecting the expired air during recovery
for up to 50 min after runs of up to 150 yd performed during breath-
holding. The mechanical work was estimated by evaluation of the pro-
pelling force from the distance-time record. The mechanical efficiency of
sprint running calculated in this way averaged 38 % for ten subjects of
widely varying ability, age and state of training.
Fenn (1930) estimated the total work done in sprint running by calcu-
lating the energy changes involved in acceleration of the limbs and changes
in potential energy of the body. Using the data of Sargent (1926) on the
total oxygen cost of running, Fenn calculated sprint running efficiency as
being 22-7 %, neglecting lateral body movements, work against internal
friction and possible energy storage in muscles and tendons.
Cavagna, Saibene & Margaria (1964) measured the mechanical work in
running at 11-20 km/hr by making accelerometric recordings in the direc-
tion of motion and perpendicular to it. The potential and kinetic energy
changes were found to have the same sign throughout the step cycle,
which precluded the possibility that a substantial part of one component
was transformed into the other. On the assumption that no transfer of
energy occurred within the body, the method indicated that the net
efficiency of running was between 40 and 50 %, and independent of speed.
The authors postulated that elastic energy stored when contracted muscles
are stretched (as in the negative work phase of the step cycle) can be used
 LOAD-RUNNING EFFICIENCY 357
for work during the next shortening of the muscles. Cavagna, Dusman &
Margaria (1968) showed in isolated frog muscle and in muscle in working
man that the work done by muscle shortening at a given velocity was
greater if the shortening was preceded by a stretch during stimulation, and
that this effect was partly due to an increase in the force of contraction of
the contractile component.
Pugh (1971) reported studies on a well trained athlete running on a
treadmill in a wind tunnel. The apparent efficiency of running was estimated
by measuring the increased metabolic rate required to overcome increased
wind force calculated from the aerodynamic characteristics of the human
body. The results for apparent efficiency were substantially higher than
any values previously quoted, the mean for four experiments being 69 %.
Lloyd (1966) derived a time, distance equation for world running records
by considering the energy available to a runner and the energy required
for running. An analogue computer solution of the equation (Lloyd &
Moran, 1966) suggested that a good fit to world records was obtained with
a value of about 50 % for the apparent efficiency with which the runner
converted metabolic energy into external work against air resistance and
into the kinetic energy of the body. Subsequent work on this equation has
confirmed this indication of high mechanical efficiency, and the present
paper describes a direct experimental investigation of the efficiency of
horizontal running in man. We have imposed an easily measurable external
load on subjects running on a treadmill, and calculated the apparent
efficiency of running by measuring the metabolic rates at various imposed
external work rates.

METHODS
The experiments were performed on a motor-driven treadmill, with a maximum
speed of 13 km/hr. A low-resistance, low-dead-space valve, similar to that of Cun-
ningham, Johnson & Lloyd (1956), was connected by 25 mm internal diameter
tubing to a 1501. polyvinylchloride Douglas bag. The bags were emptied through a
recently calibrated dry gas meter (Parkinson Cowan DI), with a standard procedure.
Gas samples were collected in glass sampling tubes and stored under pressure over
mercury for subsequent analysis for CO2 and 02 with a Lloyd-Haldane gas analyzer
(Lloyd, 1958).
External work was imposed on the subjects by means of a horizontal impeding
force. A wide canvas belt round the subject's waist was attached to a scale-pan with
a cord over a pulley supported on a firm gantry behind the treadmill. The heights of
the belt and pulley were adjusted to ensure that the pull was horizontal and the
restriction to breathing minimal. The pulley ran freely on ball bearings and no
change in metabolic rate was measured when it was replaced by a very free-running
wheel. The apparatus is illustrated in Fig. 1.
Usually eight bag collections were made, unloaded running first. A minimum of
4 min before the first collection and 1 min between later collections was allowed
for the attainment of a steady-state oxygen uptake. Bag collection times were usually
14-2
358 B. B. LLOYD AND R. M. ZACKS
2-2& min so that the duration of an experiment was seldom longer than 35 min. The
order of loading was varied throughout the experimental series, and no differences
attributable to its variation were found.
The subjects used were well trained cross-country and track athletes whose parti-
culars are given in Table 1. J. V. was an experienced treadmill runner and physiologist,
but neither H. S. nor T. M. had participated in exercise experiments previously. No
special instructions were given to the subjects about diet or activity preceding any

Fig. 1. Illustration of the apparatus.

TABLE 1. Particulars of the subjects

Best track times
Age Height Weight
(yr) (cm) (kg) 1500 m 3000 m 5000 m
J.V. 24 173 58-1 3 min 53 see 8 min 12 see 14 min 09 sec
H.S. 23 183 57-2 3 min 53 see 8 min 09 see 14 min 10 sec
T.M. 21 171 56-3 3 min 57 see 8 min 15 see 14 min 22 see

experiment. Ir, values were not measured directly but lower limits for rA
were gauged on the basis of other experimental evidence. The maximum load imposed
on any of the subjects required less than 85 % of this estimated lower limit so that
the danger of underestimating the true oxygen cost of the exercise as a result of
appreciable anaerobic metabolism was not encountered.

RESULTS
Metabolic rate was calculated as in Douglas & Priestley (1948) with
calorie values from the table of Zuntz & Schumburg cited therein (p. 64).
The lowest external load imposed was that of the empty scale-pan ( 1-24 kg),
which corresponded to a work rate of 40-5 W at a treadmill speed of
12-0km/hr. The maximum work rate imposed was 190 W (weight 5-5 kg
at speed 12-6 km/hr), and at this work rate subject J.V. had an oxygen
 LOAD-RUNNING EFFICIENCY 359
consumption of 3-96 1./min s.t.p.d. The maximum work rates were usually
lower than this, demanding oxygen uptakes of about 3-20 1./min s.t.p.d.
Table 2 gives the values obtained in all experiments for the apparent
efficiency of running against a horizontal impeding force (load-running
efficiency, LRE). Each value is the inverse of the slope of the linear regres-
sion line of metabolic rate on external work rate for the particular experi-

TABLE 2. Values for apparent efficiency (LRE), Ek and mean running speed for male
runners. Correlation coefficients and probability values for significance refer to the
linear regressions of metabolic rate on external work rate
Mean Ek
No. of speed LRE Correlation (kcal.
points (km/hr) (%h) coefficient P < kg-'. km-')
J.V. 4 11-4 29-6 0 9975 0.01 0-84
5 11-8 33-6 0-9988 0*001 0-89
6 11-7 38-3 0 9994 0.001 0 90
3 11*3 31-4 0-9998 0*01 0-84
3 12-3 32-2 1*0000 0*001 0-85
6 12-4 30-2 0 9993 0*001 0-82
8 11.9 37-9 0-9862 0.001 0-81
8 11-7 38-5 0-9986 0*001 0-81
8 12*6 40-2 0-9963 0.001 0-83
8 13-0 37.7 0 9955 0*001 0-80
8 13-0 38-9 0-9790 0.001 0-81
Mean 35.3 0-83
+ 1 S.E. + 1.19 + 0*009
T. M. 8 11-3 32-4 0-9917 0-001 0-85
8 11-5 33-6 0 9940 0-001 0-82
4 11.0 32*2 0-9971 0.01 0-80
4 12-5 34*1 0*9988 0.01 0-81
4 10-4 32-4 0.9909 0.01 0-83
4 12*3 35 0 0 9994 0'001 0-85
7 11.9 36-3 0 9905 0.001 0-84
8 12-0 38-7 0-9843 0*001 0*87
8 11.9 36-7 0-9897 0.001 0-86
Mean 34-6 0-84
+ 1 S.E. 0.75 + 0X008
H. S. 8 11-6 44-1 0 9947 0.001 0-87
8 11-6 36-0 0-9966 0.001 0X76
8 11-8 36-2 0-9910 0.001 0-82
4 11.4 34-3 0-9989 0.01 0X80
4 12.6 36-2 0-9955 0-01 0-83
4 11-5 -38-7 009990 0-001 0-78
4 13-0 38-6 0 9947 0*01 0-81
8 12-2 39-4 0-9962 0*001 081
8 12-2 45-1 0-9922 0-001 0-86
Mean 38-7 0-82
+ 1 S.E. + 1-23 0-012
360 B. B. LLOYD AND R. M. ZACKS
ment multiplied by 102 to give a percentage. The correlation coefficients
are given in the Table, all being significant at the 1 %/ level and the majority
at the 0-1 % level, and justify regarding the relation between the work
and metabolic rates as being linear. The plot of a typical experiment is

1000

4-

._

-o0
4)

800

External work rate (W)

Fig. 2. The relation between metabolic rate against external work rate.
Line fitted is the linear regression. Note the linear relation over the entire
range of external work rate.

shown in Fig. 2. The Table also gives values for speed and for the net
energy cost Ek (kcal.kg-'. km-i) of horizontal treadmill running per unit
distance and per unit bodyweight.

DISCUSSION
Previous workers have reported values for the apparent efficiency of
grade walking. Smith (1922) found a mean value of 31-3 % for four subjects
with a minimum of 25-2 % and a maximum of 48-7 %. Erickson, Simonson,
Taylor, Alexander & Keys (1946) found a variation of similar magnitude,
 LOAD-RUNNING EFFICIENCY 361
the range being from 25-9 to 40-3 % for one subject and from 24-8 to 35-2 %
in another.
Our three subjects gave ranges of 29-6-40*2, 32-4-38*7, and 34-3-45.1 %
the mean values being 353, 34-6 and 38-7 %. The differences between the
subjects were not statistically significant. The means for Ek were 0-83,
0 84 and 0 82 kcal. kg-1. km-1 the differences also not being significant.
No influence of habituation on apparent efficiency could be detected.
Erickson et al. (1946) reported an increase in grade walking efficiency with
habituation, but this was smaller than the replicate variability.
As our subjects were well trained athletes little if any improvement in
running skill was to be expected, and this expectation was borne out by
the Ek values for T. M. and H. S. For J. V., however, Ek seemed to decrease
through the series and was consistently greater for the first five experiments
than the mean for all eleven. This may indicate some habituation in J. V.
who, though experienced on the treadmill, had not previously run with
belt-loading.
Linear regressions of Ek on speed show no significant correlation for
T.M. or H.S. There was, however, a significant (P < 0 02) decrease in
Ek with increase in speed for J.V. B0je (1944) reported that Ek appeared
to be independent of running speed, and this was supported by Margaria,
Cerretelli, Aghemo & Sassi (1963), but Knuttgen (1961) and Pugh (1970)
found that the relation between metabolic rate and running speed was
not linear for all subjects over the speed ranges of 8-0-21-5 km/hr.
The magnitude of Ek obtained by Margaria et al. (1963) was 1 kcal.
kg-'. km-', this being the mean for two athletic subjects. The grand mean
in our experiments was 0 83 kcal. kg-. km-', somewhat lower than the
value quoted above. Pugh (1971) obtained values of 0-82 and 0-85 kcal.
kg-'.km-' for this energy cost at running speeds of 13-5 and 16-1 km/hr
respectively for one well trained athletic subject.
Our criterion for acceptance of the results of an experiment was that the
relation between metabolic and external watts was significant at least at
the 1 % level. This was found to be so in all experiments, including the
initial ones, so that no results were discarded.
The important point that emerges from our study is that the mechanical
efficiency of load-running is higher than that accepted for bicycling and
grade walking, and than the 25 % often assumed to apply to running. The
over-all mean from all the experiments was 36-1 %.
The net efficiency for grade running seems to be higher than for walking.
Margaria (1968) states that for grade walking and grade running the net
efficiency tends to a constant value of 25 % as the grade increases. This
constant value is attained when no 'negative work' is performed, i.e.
when no decreases in potential energy occur during a step cycle, and for
362 B. B. LLOYD AND B. M. ZACKS
walking occurs at grades greater than 22 %. This implies that calculations
of net grade walking or running efficiencies are incorrect in so far as they
assume that the relation between the lifting work rate and the metabolic
rate is linear. Pugh (1971) has found that the relation between oxygen
uptake and lifting work rate is linear for grade running, and linear except
at low grades for walking, so that, apart from this small non-linear region,
apparent efficiencies for grade walking and grade running are independent
of the grade. In this respect apparent efficiency may be a more useful
measure than net efficiency for comparison of results. Whipp & Wasserman
(1969) have suggested that net efficiency is inaccurate and misleading, and
that apparent efficiency is a more valid measure of the efficiency of
muscular work.
In biochemical terms such high efficiency values as those obtained by
Pugh (1971) and even those in this study appear to be inexplicable. The
standard free energy of oxidation of glucose is 686 kcal/mole (Lehninger,
1965), and 1 mole releases 38 moles ATP, each mole of which liberates
11 kcal (Woledge, 1971), so that a maximum of less than 70 % of the free
energy of the oxidation of glucose is available for muscle contraction. Not
all the energy of hydrolysis of ATP is, however, transformed into con-
tractile energy by muscle, some being lost as heat. Davies (1965) has
reported that the efficiency of external work performance in isolated muscle
calculated from the ATP energy used varies from approximately 50 % in
slow muscle to 35 % in fast muscle. It thus appears that at least half of the
ATP energy is lost as heat, implying that values for whole-body efficiency
-above 35 % are unlikely. For isolated muscle, it has been estimated on the
basis of thermodynamic and mechanical measurements that the efficiency
of the entire chain of reactions between the oxidation of glycogen and the
performance of mechanical work is less than 20 % (Wilkie, 1960). Factors
such as increased circulation, respiration, posture and conditions of move-
ment combine to decrease the measured efficiency of exercising muscle in
man. There is then a considerable discrepancy between the apparent
efficiency of the exercising runner and the efficiency of isolated muscle.
The reasons for this are unknown, but it is possible that there are major
differences in the performance of isolated muscle (usually frog muscle at
0 C) and mammalian muscle at 38 C.
All methods of determining running efficiency have, however, given
higher values than those for bicycling and grade walking. This may be
further evidence in support of the hypothesis of Cavagna et al. (1964) that
substantial energy storage occurs in muscle stretched during contraction.
 LOAD-RUNNING EFFICIENCY 363

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