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Respiratory Quotient During Exercise
Respiratory Quotient During Exercise
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
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
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-
b= 0.017
- Men
x 402
--I- Women *A )Excess CO2
Wome
b=0.014
40
20
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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