Respiratory quotient during exercise

B. TSSEKUTZ, JR., AND K. RODAHL

Division Research, Lankenau Hospital, Philadelphia, Pennsylvaniu

ISSEKUTZ, B*, JR., AND K. RODAHL. Respirafory quotient during Several observations seem to support this assumption.
exercise. J. Appl. Physiol. IS(~): 606-610, I 961 .-O2 uptake Robinson (I), investigating the physical work capacity
and CO2 output were determined during exercise on the of different age groups, pointed out the parellelism
bicycle ergometer. During moderate and heavy work three between RQ and blood lactic acid level found during
phases could be distinguished in the time course of RQ: a) maximal work in adult men. In a classification of physi-
initial increase, B) secondary drop followed by c) a continuous
cal work, WelIs et al. (3) include not only pulse rate but
rise to a steady state which was reached after 335-4 min work.
also accumulation of lactic acid and RQ. At exhausting

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The rise of work RQ (ARQ) above an assumed metabolic RQ
of 0.75 (or 0.83) showed an approximately logarithmic in- work, blood lactic acid increases six times above the
crease as work load increased. In the same subject there was a resting level, with a concomitant increase of RQ above
straight-line correlation between %onmetabolic excess CO* 1.0. The rise of blood lactate level and the decrease of
( = total CO2 minus 0.75 X 0,) and the increase of blood lac- blood CO2 concentration observed during the first 5
tate level (P < 0.001). In pooled calculations, a correlation min of exercise lead Christensen and Hansen (2) to the
coefficient r = 0.92 was found. Whereas the respiratory minute conclusion that the work RQ could be used in metabolic
volume plotted against 02 uptake or CO2 output showed a studies only if the blood CO2 and lactic acid concentra-
relative hyperventilation as the subject approached maximal
tions reached a steady state after 10-30 min of exercise,
aerobic capacity, excess CQ:! increased with the ventilation in a
depending on the work performed. We felt that a
straight-line fashion. It was concluded that the ARQ represents
the percentual participation of anaerobic glycolysis in the total continuous recording of the RQ during the first 4-5 min
energy expenditure rather than the fuel used during exercise. of work, an estimation of the excess (nonmetabolic) CO2
and its comparison with the blood lactate level might
supply further information in regard to the interpreta-
tion of the work RQ. The results reported here appear to

I T IS WELL KNOWN
exercise of short duration
that the work RQ measured during
(4-5 min) is higher than the
show that RQ during
function
the early phases of work is the
of the 02 supply (or rather deficiency) of the
muscle rather than the function of the fuel utilized in
resting RQ. This rise of RQ seems to depend on the the muscle work.
intensity of the work; at heavy work loads it is always
above LO (1-3); RQ around I .2 is quite common, and
even values as high as 2. I have been reported (4). Ob- METHODS
viously, the usual interpretation of resting RQ cannot
be applied to the work RQ, at least not in the early The experiments were carried out on 19 untrained
phase of exercise (2). The relative increase of CO2 output men 23-65 years of age and 8 women aged 50-63. The
subjects worked f *r m;n nn the bicycle ergometer
compared to the 02 uptake likewise cannot be ascribed
to a relative hyperventilation, because overbreathing described by von Dijbeln (6). This ergometer permits an
would cause a drop in the alveolar as well as the arterial accurate setting of the work load and the work performed
is practically independent of the body weight. The pedal
pCO2, which appears not to be the case. The alveolar
pCO2 was found to be practically the same during frequency was 50 rpm. Each subject started off at light
moderate and maximal work as at rest (I). The recent work (300 or 450 kpm/min) and after about 10-15 min
rest a heavier load was applied. On the same day two
investigations of Mitchell et al. (5) showed that during
periods of exercise were carried out and the experiments
maximal work the pH and CO2 content of the arterial
were continued until the maximal 02 uptake was
blood decreased in such a way that the arterial pC0;
reached. Blood samples were taken from the fingertip
remained fairly constant. Therefore it seemed likely that
before and 1-2 min after work. The lactic acid level of
the increase of respiratory quotient during exercise was
the blood was measured according to S trplms modifica-
partly or entirely due to an imbalance between the
tion (7) of the Barker and Summerson method.
formation and elimination of acids (particularly lactic
Two methods were used to measure the 02 uptake and
acid) and to their reaction with the bicarbonate pool of
CO? output : r) after 3% min work the expired air was
the body (extra- and intracellular HCO,).
collected in two Douglas bags, each for about 45 sec.
Received for publication 26 January 1961. Analyses of room air and expired air were carried out
606
RESPIRATORY QUOTIENT DURING EXERCISE 607
TABLE I. cumparative gas analysis by Nuyods diaferometer TABLE 2. Comparative measurements during work

Noyons Diaferometer SchoIander-Noyons

Room Air Minus
DiluteiExpired Scholander, % Difference, %
/ N1 Range
1d%Ekce
] SD / SE
AQ2 73 0.670 0.68 -0.01
108 1.068 I. IO -0.032
95 I .882 1.88 +0.002
AGO* 78 O-559 0.54 +o.o1g
157 1.152 I.14 +0.012
252 I ,808 I .81 -0.002
flow rate of about 4 liters/min. This flow was con-
tinuously sampled by the diaferometer at a rate of 0.7
liters/min. The diaferometer readings were compared
according to the micro method of Scholander (8). with the difference between room air and bag air as
Usually the average of the two bags was used for the measured by the Scholander method. As Table I shows,
estimation of 02 uptake and CO2 output. If, however, the differences between the two methods do not exceed
at heavy work load the second bag gave a higher 02 ko.03 %. Table 2 records comparisons between the

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uptake, this value was used. two methods on subjects working on the bicycle ergom-
2) The 02 uptake and CO2 output were continuously eter at 300 kpm/min for 30 min. In these cases, im-
recorded by means of a Noyons diaferometer (manu- mediately after taking the diaferometer readings, the
factured by Kipp et Zonen, Delft, Holland). This instru- expired air was collected in a bag. The volume of the
ment, working on the principle of heat conductivity bag was measured by a dry gasometer and the 02
of gases, showed the differences between room air and uptake and GOa output were computed (STPD). The
expired air. The expired air, flowing through a Krogh mean difference in the RQ was less than 0.01.
valve and a hose, entered a plastic hood, the volume of Finally, the correct time course of the RQ in the first
which was about 50 liters. This reduced the high flow 2 min of exeycise, calculated according to the equation
rate of the expired air and served as a mixing drum. An given above, was compared with the Douglas bag
air-flow (at light work I 70 liters/min, at heavy work 240 method, repeating the experiment on the same subject
liters/min) maintained by a pump, diluted and carried at the same work load at a different time (Fig. I).
the expired air into the diaferometer. The equipment
worked with a constant lag of 42 set; this lag was paid RESULTS
off at the end of the measurement and therefore caused
a simple linear shifting of the time curve. In non-steady Fig. I shows the changes of RQ in the same subject
state, for instance during rising metabolism, there was at six different work loads, from the lightest-150 kpm/
an additional delay caused by the mixing drum. How- min-to the over-maximal, 1500 kpm/min, which could
ever, the actual, a, 02 uptake (or CO2 output) can be be continued only for 2% min. Three distinct phases
calculated for any given time, t, according to the were found in the time course of the work RQ: a) an
equation : initial increase, 6) a characteristic drop to a low level,
in this subject to about 0.70, followed by c) a gradual
increase which reached a steady state after 346-4 min.
At very light work (150 and 300 kpm/min) no initial
increase is found and the 2nd phase lasts longer,
whereas above 450 kpm/min the initial rise becomes
where m is the measured 02 uptake at time t, r = flow more pronounced and the following drop is less marked.
rate (I 70 or 240 liters/min) and v is the volume of the Finally, at 1500 kpm/min work the RQ rises in the I st
mixing drum (50 liters). This equation was used to phase to a peak value of I. I g and falls only to about 0.82.
follow the quick changes of RQ and to estimate the A similar finding was described by Robinson (I) for
total amount of bicarbonate CO2 expired during the maximal work on the treadmill. In the 3rd phase the
first 4 min of work. Comparisons between the excess CO2 RQ rises gradually, the rate of increase depending
and the increase of blood lactate were based on the entirely on the work load applied. While it was about
Douglas bag method. o.oq/min at 150 kpm/min, it reached the maximal rate
The diaferometer is supplied with a factory-made of increase of about o.35/min at the highest load studied.
calibration table which was checked from time to time In order to obtain an approximate estimation of the
according to the following procedure: 10-20 liters bicarbonate, nonmetabolic, excess COZ, it was necessary
expired air were collected in a Douglas bag and sufficient to assume a reference RQ, the value of which was
room air was added to it that the CO2 concentration expected to be reasonably close to the real metabolic RQ
would be approximately 0.5; x and 2 %. After thorough during the 4- or 5- min exercise. During the calculation
mixing, a small pump pulled the air from the bag at a of excess COZ, two reference metabolic RQs were used,
608 B. ISSEKUTZ, JR., AND K. KODAHL

8 = Scho/onder vohes
reached a value of at least 65 mg/roo ml (A lactate was
Anw fm//
more than 55 mg/~oo ml). In -these untrained subjects
2269 with a wide age range, the maximal 02 uptake varied
from I. 7I to 3.03 liters/min. It is well known that the
02 uptake at a given submaximal load is about the same
1.0 whether the subject is trained or untrained, while
0.9
200 0.9 increase of blood lactate depends on the aerobic work
0.8 100
0 0.8 capacity of the individual. The closer the subject works
0.7 -100
0.7 to his maximal 02 uptake, the more pronounced is the
rise of lactic acid in the blood as a result of increasing
insufficiency of the 02 supply. This is reflected in Table
3 by the wide range of coefficient of variability in the A
lactate (26.9-66.7 %) compared to the narrower range
in the 02 uptake (4.7-14 %). Also the excess COP,
calculated on the basis of a metabolic RQ 0.75, showed a
-2-101234 I234
high coefficient of variability ranging from 10.6 to 41 %*
More important is the fact that the correlation coefficient
FIG. I. Time course of RQ and excess CO2 ( = total CO2 minus between A lactate and excess COZ, based on I 02 meas-

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.075 02 uptake) at different work loads on same subject. urements, was found to be r = 0.92; calculated with a
metabolic RQ of 0.83~ r = 0.91 was obtained. Con-
0.75 and 0.83, respectively. Thus: excess CO2 = total sidering the wide range of body weights, from 62 to
CO2 minus 0.75 X 02 (or 0.83 0,). We preferred the 94 kg and hence the individual differences in blood volume
value of 0.75, however, for reasons discussed below. and size of the lactate space (factors which affect the
In the 1st min of work (above 450 kpm/min), some values of A lactate but not the excess CO& it is reason-
excess CO* was expired and in the 2nd phase some CO2 able to say that the correlation is very good.
was retained; the amounts are negligible, however, When the respiratory minute volume was plotted
compared to the changes observed in the 3rd phase. Fig. against 02 uptake, CO2 output and excess COZ, respec-
I shows the estimated output of excess CO* (assumed tively, burves were obtained as shown in Fig. 4 (one
metabolic RQ: 0.75) in this 3rd stage until the end of male and one female subject). At the beginning the
the 4th min. While the 02 uptake plotted against work ventilation rose proportionally with the 02 uptake; but
load increases in the usual straight-line fashion, the as soon as one approached the maximal 02 uptake of
excess CO2 rises rapidly, reaching a value at I 350 kpm/ the subject, the minute volume increased sharply, that is,
min (the maximal aerobic capacity of the subject) the oxygen extraction dropped. At heavy work there was
which is almost go times higher than that measured at a relative hyperventilation as far as the 02 uptake
x50 kpm/min (Fig. II)* The changes in the amount of was concerned, a finding which has been described by
excess CO2 as the work load changed resembied closely many i;lvestigators (I, I I, I 2). A similar although less
the changes in blood lactate levels found by Astrand et pronounced hyperventilation was found also in regard
al. (g, IO) under experimental conditions similar to those to the total CO2 output, whereas the excess CO2 showed
used here. a straight-line increase proportional to the ventilation.
A calculation based on RQmet = 0.83 yielded no Our interpretation is that the so-called relative hyper-
excess CO2 at 150 kpm/min and only g ml during the 4 ventilation is brought about by accumulation of
min at 300 kpm/min, while the maximal work load metabolic acids and consequently by formation of
brought an almost 2oo-fold increase. The ARQ (= work excess COZ. This, of course, does not exclude other
RQ minus 0.75) at the 4th min of exercise showed an causes, such as reflexes arising from the working legs (I 3)
approximately logarithmic increase between 300 kpm/ or the effect of rising body temperature. It should be
min and the maximal work load. At the maximal 02 pointed out, however, that these factors are expected to
uptake of the subject, ARQ reached a value of 0.40. decrease the arterial pCOz, while Mitchell et al. (5) by
In a series of experiments the expired air was col- direct sampling of the brachial artery did not find any
lected in two Douglas bags at 3% and 4% min of cycling significant change of pCOz in maximal exercise.
at different work loads. The 02 uptake and CO2 output
DISCUSSION
were measured, and the excess CO2 was calculated (for
1~0th RQmet = 0.75 and 0.83, respectively) and plotted In this study the calculation of excess CO2 was based
against the increase of blood lactic acid level (A lactate on two hypothetical metabolic RQs: 0.75 and 0.83,
mg/ I oo ml). The regression coefficient was significant respectively. It was felt, however, that the value of 0.75
at the P < 0.00~ level (Fig. 3). was a closer approximation of the real metabolic RQ
Table 3 summarizes the results obtained on 19 male during the first 5 min of exercise. This assumption was
subjects at work loads from 300 kpm/min up to the supported by three observations: a) Huckabee found
point where the subject reached his maximum aerobic (14) that very mild exercise which increased the 02
work capacity; that is, higher load did not cause any uptake only to about 0.58 liter/min induced a 0.6 mM/
further increase of 02 uptake and blood lactate level liter (5.4 mg/Ioo ml) increase of blood lactate. By using
 RESPIRATORY QUOTIENT DURING EXERCISE

b= 0.017

- Men
x 402
--I- Women *A )Excess CO2

Wome
b=0.014

40

20

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300 600 900 0 20 40 60 0 20 40 60 0 I 2 3
kpm /min A Luctafe (mg //UU m/J O2 I CO2 md excess CO2

2. Relationship
FIG. of 02 uptake, excess CO2 (shaded area on FIG. 4. Relationship of respiratory minute volume to 02 uptake,
Fig. and ARQ ( = RQ - 0.75) to work load.
I) CO2 output and excess CO2, respectively.
FIG. 3. Correlation between excess CO2 (after 3%-5 min work)
and increase of blood lactic acid level (A lactate mg/r oo ml).

TABLE 3
Work, kpm/min 300 450 750 Max.
02 liters/min. o*g7 * 0.02 1.28 A 0.01 1.53 32 0.03 r-77
zt 0.08 2.07 h 0.08 2.39 * 0.06
SD ho.07 0.06 0.15 0~24 0.29 0.34
cv % 7.2 (10) 4*7 (21) 9.8 (22) 13.6 (9) 14 (13) 14 (28)
RQ 0.87 =t 0,006 0.96 =t 0.012 0.98 rt 0.012 1.08 h 0.023 1.06 dz 0.007 I.19 * 0.013
SD 0*04 0.06 0,05 0,07 0.03 0.06*
cv % 4.6 (10) 6-3 (21) 5-I (21) 6.5 (9) 2.8 (13) 5 (23)
A Lactate mg/roo mlt 5.4 * I.2 13.5 It I.2 19.1 4~ 2.4 34.7 A= 3-2 42.6 j, 4.5 64.7 & 4.6
SD 3.6 56 II .4 9*5 16.9 17.4
cv % 66.7 (IO) 41.5 (21) 59.7 (22) 27*4 (9) 39-7 (14) 26.9 (14)
Excess CO 2 liters/min $ 0.15 It 0.02 0.27 =t 0.02 0.35 * 0.02 o-57 xt 0.06 0.66 * 0.04 o*g7 * 0.02
SD 0.061 0.086 0.100 0.184 0.136 0.103*
cv % 41 (10) y*g (21) 28.6 (21) 32.3 (9) 20.6 (13) 10.6 (28)

Values are means & standard error; SD = standard deviation; CV = coefficient of variability. Correlation coefficient
rALactate--exc.COZ = 0.918 (102). * Maximal RQ, ext. CO2, and lactate depend on motivation and do not show normal distribution.
t Work lactate less rest lactate. $ Total CO2 less 0.75 OZ.

RQ met = 0.83 in our estimation of excess COZ, the c) Our unpublished experiments have shown that
lightest work, 150 kpm/min (= 0.651 liter/mm O,), trained persons working at 300 kpm/min for 30 rnin had
would not produce any excess COZ, and 300 kpm/min a work RQ of o-74, and 0.73, although the resting RQ
( = 0.97 liter/min 0,) would cause only about g ml measured under basal conditions was found to be 0.80.
excess COZ. Even if we assume that only the lactic acid Therefore, it seems to be a reasonable assumption that
produced entered a blood volume of about 6 liters the metabolic RQ during work, at least in its early
and no lactate left the circulation during the 4 min work, phase, is somewhere between 0.70 and 0.80.
the increase of blood lactate would not be measurable (g During moderate and heavy work three phases were
ml CO2 = 36 mg lactate = 0.6 mg%). In our subjects, observed in the time course of RQ. Whereas the 3rd
however, the average increase of blood lactate found at phase correlated with the changes of lactic acid level
300 kpm was 5.4 mg/roo ml. This suggests that the in the blood, the explanation of the first two phases is not
metabolic RQ was less than 0.83. yet clear. One possibility is that the initial increase of
6) Recently, Christensen (15) has shown that with RQ is brought about about by a cerebra-cortically
intermittent work on the treadmill (20 km/hr run, 5 set induced hyperventilation, followed by a compensatory
work and 5 set rest) it was possible to increase the 02 hypoventilation (I). This assumption is based on the
uptake for 30 min to as much as 4*4 liters/min without fact that, at rest, arbitrary hyperventilation increases, hy-
any appreciable rise in blood lactate. The average RQ poventilation decreases the RQ; and on the observation
measured at a fairly steady state of blood lactic acid of Krogh and Lindhard (16) that in the 1st min of
level (23-20 mg%) was 0.78. work there is a transitory drop of the alveolar pCOz be-
6IQ B. ISSEKUTZ, JR., AND K. RUDAHL

low the resting level. It should be mentioned, however, exercise. Since the lactate concentration is higher in
that Christensen and Hansen (2) found about a I mm Hg the working muscle than in the blood, and since the
increase of alveolar pCOz in the I st min of exercise. diffusion of bicarbonate COe is presumably more
Another explanation would be that the observed rapid than that of lactate it appears likely that the
changes of RQ in the 1st and 2nd min of work are the excess CO2 follows the anaerobic metabolism more
result of a delayed circulatory response and consequently closely than does the blood lactate level.
of an initial discrepancy between 02 demand and 02 An accelerated fat and protein metabolism appears
supply of the working muscle. The percentual participa- to play a cardinal role from the start of the work. The
tion of anaerobic glycolysis in the total energy expendi- site of this may be partly the working muscle itself
ture is considerably greater in the first few seconds of (2~22), partly the liver in the form of an increase&
exercise than in the steady state. This was shown by glyconeogenesis, probably hormonally induced. Wright-
Mtiller and Hettinger (I 7), who measured the 02 debt ington (23) reported that the work RQ (I o min work on
after IO set work on the bicycle. Therefore the initial the bicycle) tended toward a central value of o-88-0~89
rise of RQ may be caused by an increased formation of although the resting RQ varied from 0.69 to I .oo as a
lactate. In the 2nd min, the anaerobic glycolysis is result of different diets. This is to be expected if the
slowed down by the rapidly increasing 02 supply metabolic work RQ, regardless of the rest RQ, was
(Pasteur effect) and temporarily more lactic acid will between 0.70 and 0.8o and if the accumulation of acids
be eliminated than formed, causing a secondary drop of caused the rise from this low value to 0.88. Special
the work RQ. The observation that the initial increase dietary conditions such as extremely high or low carbo-

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is more pronounced and the secondary drop is less hydrate (24) or a sugar load before the work (23) may
marked during heavy than during light work, supports influence the work RQ, perhaps because they induce a
this explanation. A cortically induced hyperventilation shifting in the insulin glucocorticoids-hypophyseal
is expected to be more effective under a light load, since growth hormone balance (25) which may inhibit (insu-
here much less excess CO% could increase the respiratory lin) or promote (corticosteroids) the glyconeogenesis
quotient than during heavy work with higher 02 during exercise.
uptake. Finally, it should be pointed out that great caution
The 3rd phase rise of RQ above 0.75 as reference should be used in the interpretation of work RQ even in
value, called ARQ, is the ratio: excess C02/02 uptake, prolonged exercise. In this case we must assume a lactate
and seems to reflect the percentual participation of space of 600 ml/kg body wt; definitely measurable
changes in the work RQ during exercise may therefore
anaerobic glycolysis in the total energy expenditure
simply be caused by changes in the size of the pool of
rather than the fuel used for the work, The formation
lactate (and other acids) without appreciable changes in
uf lactate and other acids may entirely mask the metabo-
the blood lactate level. This is especially the case at
lic RQ and a stoichiometric estimation of excess CO2 moderate work loads with low 02 uptake when very
seems to be impossible because of the rapid diffusion small amounts of excess CO2 can cause a considerable
of lactic acids through the cell membranes (13, 18, 19). shift in the RQ. The extremely low recovery RQs ob-
The space of distribution may shift from the volume of served after work, repeatedly reported (4, 26), may be
blood to the volume of body water (about 60 % of caused partly by a gradual decrease of the lactate pool,
body wt) depending on the intensity and duration of with a concomitant increase of the bicarbonate pool.

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