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Interaction of acute changes in exercise energy expenditure and energy intake Buscar ProQuest ;

on resting metabolic rate

Bullough, Richard C; Gillette, Cynthia A; Harris, Mary A ; Melby, Christopher L . T h e A m e r i c a n J o u r n a l o f C l i n i c a l N u t r i t i o n 61.3
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(Mar 1995): 473.
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The effects on resting metabolic rate of energy intake and exercise energy expenditure were examined in eight trained men under four conditions: high energy Crdoba
flux, low energy flux, negative energy balance and positive energy balance.

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The influence of chronic exercise on resting metabolic rate (RMR) is unclear. An elevated RMR has been observed in endurance-trained compared with untrained
individuals in some cross-sectional studies (1-3). However, other studies have found trained subjects to have RMR values no different from those of sedentary Elementos relacionados
control subjects (4-8). The reasons for the discrepant findings may be related to study differences in the time intervals between the last exercise bout and
measurement of RMR (5, 9), to the energy intake during the days immediately preceding RMR measurement, to the quality of the RMR measurements, or to the
degree of exercise training of the subjects. Several of these factors may influence sympathetic nervous system activity and it is possible that RMR differences Buscar con trminos de indexacin
between groups, as observed in some studies, may at least in part be mediated by catecholamine concentration.

A 1-12-h fast before RMR measurement is the standard followed by investigators to eliminate the thermic effect of food on basal energy expenditure. However, Materia
the time interval required to eliminate any residual effect of exercise on RMR is unclear. Most studies have found that mild to moderate exercise elevates Metabolism
metabolic rate for only a few minutes to a few hours (10, 11). However, we observed elevated RMR in trained men 15 h after 90 min of strenuous resistance Men
exercise (12), suggesting that high-intensity and long-duration exercise may produce prolonged elevations of RMR. If this is true, failure tt allow adequate time to Exercise
elapse between exercise and RMR measurement could influence the RMR values. Clinical trials

Several early studies that identified higher RMR values in trained athletes allowed 24 h between the last exercise bout and RMR measurement. Because subjects Buscar
abstained from exercise for this relatively long period (vs the 10-12-h fast), an acute effect of exercise on RMR was discounted and investigators suggested that
the higher RMR in athletes might be due to adaptations to chronic exercise (1, 2). This hypothesis was supported by studies that allowed more time (36-48 h)
between exercise and RMR measurement and still reported elevated RMR in trained individuals (13-15). However, several studies that used an even longer
interval (48-56 h) between exercise and RMR measurement failed to find elevated RMR in trained subjects (5, 7). Together, these studies suggest that any ebrary e-books
elevation of RMR in athletes may result from acute perturbations of strenuous exercise, possibly lasting as long as 2 d, rather than from long-term adaptations to
chronic exercise training. 1. Exercise and Cognitive
Function
Acute changes in energy intake and energy balance can also influence RMR. Ballor and Poehlman (16) observed elevated RMR in trained women who had
abstained from exercise for >/=36 h but in this study (as in some others) energy consumption was not controlled during the period of exercise abstention. If
subjects did not lower their energy intake during this period, they may have been in an overfed state, which is associated with an increased metabolic rate. In
another study (4), RMR did not increase after 9 wk of exercise training, but subjects were underfed based on doubly labeled water measures. Still other studies
report:increased RMR in overweight subjects placed on an exercise program despite body weight loss suggestive of negative energy balance (17, 18). We 2. Monitoring Metabolic Status:
previously observed that RMR decreases in exercising athletes when they move into an energy deficit state (19). Earlier studies also suggest that the Predicting Decrements in
combination of high energy expenditure and energy intake (high energy flux or turnover) could elevate RMR in athletes (2, 13, 15), even when they are in energy Physiological and C...
balance, and recent data from our laboratory supports this hypothesis (20).

Given the possible influence of the length of abstention from exercise, the interaction of exercise and energy intake, and the acute state of energy balance on
RMR, it is important to examine these issues further. Therefore, the purpose of this study was to investigate the influence of different levels of exercise energy 3. Metabolic Syndrome :
expenditure and energy intake, and their interaction, on RMR in trained male subjects. The mean RMR values under each condition in the trained subjects were Underlying Mechanisms and
compared with the mean RMR of untrained subjects who abstained from exercise. Additionally, the influence of these different conditions on norepinephrine and Drug Therapies
epinephrine concentrations was investigated and the relationship of changes in RMR to changes in catecholamine concentration was examined.

Subjects and methods

Subjects

Sixteen (eight trained and eight untrained) young (aged 18-35 y), healthy male subjects were recruited for this study. Trained subjects were cyclists or
triathletes/biathletes currently exercising >/=5 d/wk, and had engaged in training for >/= 2 y previously. The untrained subjects had not engaged in any formal
exercise during the previous 2 y and had never engaged in any athletic competition involving endurance activities. Subjects were nonsmokers, were weight stable
(+/= 2.5 kg) during the year before the study, were not taking any medication known to affect metabolic rate, and were free of any injury or illness that could
influence metabolic rate during the study. Before the study, approval was obtained from the Colorado State University Human Research Committee and each
subject signed an informed consent document.

Experimental design

According to a 2 X 2 factorial design (exercise or no exercise by high or low energy intake), RMR, respiratory exchange ratio (RER), and plasma concentrations of
norepinephrine, epinephrine, triiodothyronine, insulin, glucose, and creatine phosphokinase (CPK) were measured in each trained subject under four different
controlled conditions: 1) a condition in which subjects were in a state of high energy flux (HF), exercising at 75% of maximal oxygen capacity (VO sub 2 max) for
90 min on each of 3 d while consuming a controlled diet adequate to maintain energy balance; 2) a condition in which subjects were in a state of low energy flux
(LF), abstaining from exercise for 3 d while consuming a controlled diet adequate to maintain energy balance; 3) a condition in which subjects were in negative
energy balance (NEB), exercising at 75% VO sub 2 max for 90 min on each of 3 d while consuming the controlled diet designed to maintain energy balance during
LF; and 4) a condition in which subjects were in positive energy balance (PEB), abstaining from exercise for 2 d while consuming the controlled diet designed to
maintain energy balance during HF. The untrained subjects were tested for 2 d only in their usual state of LF. Subjects were asked to abstain from exercise for 2 d
immediately before the study. To minimize the possibility of a testing order effect, trained subjects were randomly assigned (by using computer generated
random numbers) to one of two testing order groups. There were four individuals per group. In group 1, the order of testing was HF, LF, NEB, and PEB, whereas in
group 2 the order of testing was NEB, PEB, HF, and LF. Subjects went immediately from one condition to the next. The rationale behind these groups was to allow
the measurement of RMR over time when subjects moved from a period of exercise to a period of inactivity (eg, HF to LF and NEB to PEB). Note that in this
design PEB always followed NEB. Therefore, it may be appropriate to interpret PEB as being a "refeeding" condition. A sample testing schedule for trained
subjects is presented in Figure 1. (Figure 1 omitted.) As trained subjects moved from an exercise to a nonexercise protocol, RMR was measured ==21, 45, 69, and
93 h after the last exercise bout.

Preexperimental procedures

Approximately 2 wk before the experiment began, a series of tests were performed on each subject. Body composition was estimated from body density by
underwater weighing and the Siri equation (21). Residual lung volume was determined by the oxygen-dilution technique of Wilmore (22).

Maximal exercise tests were performed on all subjects by using a Monark bicycle ergometer (Stockholm) and a progressive maximal workload protocol. Oxygen
consumption was assessed by using indirect calorimetry (Horizon MMC; SensorMedics, Anaheim, CA) and a mouthpiece and Hans-Rudolph three-way valve.
Heart rate (HR) was measured and recorded throughout the exercise test by using CM-5 electrode placement.

Diet and assessment of energy requirements

All meals were controlled during the 13-d testing period for each subject. The diet contained (by energy) 64% carbohydrate, 23% fat, and 13% protein and met or
exceeded the recommended dietary allowance (RDA; 23) for all nutrients according to analyses with the Nutritionist III computer software program (N sup 2
Computing, Salem, OR). Although the energy content of the meals varied between conditions, the relative proportion of macronutrients remained constant for all
individuals on all days. All meals during the entire study period were prepared in a research kitchen and given to each subject. Subjects were instructed to
consume all of the food provided them (but no more), to abstain from alcohol, and were encouraged to report any deviations from protocol. Breakfast was
consumed from 0745 to 0800, lunch and snacks from 1100 to 1500, and dinner from 1830 to 1900, after which time subjects fasted and abstained from caffeine-
containing drinks. The energy content and macronutrient composition of all breakfasts and dinners within subjects remained the same throughout the study (ie,

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only the lunch and snacks changed between treatments). This was done to minimize the possibility of a confounding effect of meal size and composition
differences on exercise metabolism in trained subjects, and to standardize the effect of the evening meal on the next morning's RMR in all subjects.

The daily energy content of the diet used during LF and NEB conditions in trained subjects was calculated by multiplying the individual's RMR (measured that
morning) by 1.5. This value approximates the energy required for zero energy balance when people perform their obligate daily activities while abstaining from
additional physical activity (24). The energy content of the diet used in HF was calculated for each individual on each day according to the following formula:

(Energy cost of exercise X 1.15) + (RMR X 1.5)

The value equal to 15% of the energy cost of exercise was added to the total in an effort to compensate for elevated postexercise oxygen consumption (EPOC).
This adjustment was based on data presented in studies by Bahr (25) and Maehlum et al (26) in which they studied EPOC in subjects cycling at an intensity and
duration similar to that used in this study. HF exercise data were used to calculate the energy content of PEB meals. Therefore, any variation in the daily energy
intake when HF and PEB data were compared was due to variations in that day's RMR. The energy content of meals for untrained subjects, who were in their
usual state of LF, was calculated by multiplying individual RMR values by 1.5, as it was in trained subjects.

In an effort to verify that trained subjects were in energy balance during HF and LF, in energy surplus during PEB, and in energy deficit during NEB, and that
untrained subjects were in energy balance in their LF state, plasma triiodothyronine was measured under each flux condition (triiodothyronine concentrations are
sensitive to under- and overfeeding; the specific procedures regarding this hormone are described later). Additionally, although not highly sensitive to short-term
changes in energy balance, body weight was measured to the nearest 100 g each morning and skinfold-thickness measurements were performed on day 1 of the
study period and on the final day of each energy flux condition for each subject. Skinfold-thickness measures were taken at the chest, abdomen, and thigh by
using a Lange caliper (Cambridge Scientific, Cambridge, MD). All skinfold-thickness measurements were performed by the same investigator (RB) on the right
side of the body to the nearest 0.5 mm.

Measurement of RMR

RMR was measured by indirect calorimetry using a ventilated-canopy system (Horizon MMC). Energy expenditure was calculated by using the Weir formula (27).
Metabolic testing was performed under outpatient conditions, with subjects being driven by automobile to the laboratory by an investigator before testing. The
RMR measures in our laboratory were shown previously to be highly reproducible (CV for repeated measures, 3.0 +/= 1.3%), with outpatient measures similar to
inpatient measures when subjects are transported to the laboratory as described above (28). On each of the 11 consecutive days that RMR was measured,
subjects were awakened at home at ==0630 and transported to the laboratory for metabolic testing at 0700. Upon awakening, subjects were asked to rate the
quality of that night's sleep on a subjective 1-10 scale, with 10 being the highest-quality sleep. All RMR tests were conducted with subjects in a 12-h fasted state.
Metabolic tests lasted 45 min and were conducted in a thermoneutral (23 degC), semidarkened environment.

Upon arriving at the laboratory, subjects were allowed to void and height and weight were then measured according to standard procedures. While metabolic
equipment was being calibrated before testing, subjects were familiarized with the canopy system. Subjects were then placed in a supine position on a
comfortable bed, chest electrodes were attached, the respiratory canopy was placed over the subject's head, the room was darkened, and, after instrument
wash-in, data collection was begun. At 12 min, the subjects were inspected to ensure that they were comfortable and awake. The first 15 min of the 45-min
metabolic test were used as a habituation period as described previously (28), with only the final 30 min of data being used to calculate the subject's RMR and
RER.

Exercise protocol

After measurement of RMR and consumption of a small meal at 0800, trained subjects began exercise at 0900 on each of the 3 exercise days during both HF and
NEB. Using a bicycle ergometer, subjects exercised for 90 min at a workload approximating 75% of their predetermined Ozmax. HR was continuously monitored
by using a portable chest strap monitor (Polar CIC Inc, Port Washington, NY). Expired gases were measured for 5 min at the end of each 15-min period to
determine exercise VO sub 2 and to make workload adjustments to maintain exercise intensity at 75% of VO sub 2 max. Additionally, during each 5-min
measurement period, workload, HR, and perceived exertion (29) were recorded. Subjects were allowed to consume water ad libitum during exercise.

Plasma assays

Immediately after the end of the final RMR test in each condition, a venous blood sample was drawn from the antecubital space of the arm for measurement of
resting epinephrine, norepinephrine, triiodothyronine, CPK, glucose, and insulin concentrations. Samples were collected in evacuated tubes containing EDTA,
centrifuged at 22 degC (2500 x g for 20 min), and then separated. Hematocrit was measured in duplicate to facilitate adjustment for changes in plasma volume
(30). Plasma was frozen immediately and stored at -70 degC until analyzed. Catecholamine plasma samples were stored with 5 mmol dl-dithiothreitol. Plasma
was assayed for norepinephrine and epinephrine concentrations by using HPLC as modified by Hallman et al(31) and described by Mazzeo and Marshall(32).
Plasma triiodothyronine and insulin were analyzed by radio-immunoassay. Plasma glucose and CPK were measured by using colorimetric assays (Sigma, St
Louis).

Data analyses

Statistical analyses were performed by using SAS (Statistical Analysis System, Cary, NC) and Minitab statistical software (Minitab, State College, PA). Differences
were considered significant when P < 0.05. Descriptive statistics were computed to describe the groups and to characterize the distributions of variables under
each of the four testing conditions. The primary dependent variables were RMR (kJ/min), RMR statistically adjusted for fat-free mass (FFM), and plasma
catecholamine concentrations. Within the trained subjects, a 2 x 2 factorial analysis of variance (ANOVA) (exercise and no exercise by HF diet and LF diet) was
used to examine the main effects of exercise, the main effects of diet, and any exercise by diet interactions. Appropriate preplanned pair-wise comparisons were
made by using the least-squares difference posthoc test when F ratios were significant. To examine whether RMR changed as the time interval increased from
the last exercise bout in the trained subjects, these data were analyzed by using a within-subjects repeated-measures ANOVA (subject by condition by time).
Comparisons were made between untrained and trained subjects by using independent t tests and analysis of covariance (ANCOVA), with FFM serving as the
covariate. Additionally, Pearson product-moment correlations were performed by using the entire sample of trained and untrained subjects to investigate the
relationship of VO sub 2 max (L/min) and RMR (kJ/min) adjusted for FFM.

Results

The physical characteristics of the trained and untrained subjects are presented in Table 1. There were no differences in age, height, FFM, or maximal HR
between these groups. Resting HR, total body weight, and percent body fat were lower in trained subjects than in untrained subjects. VO sub 2 max was greater
in trained subjects, expressed either in L/min or when adjusted for total body weight and for FFM. Additionally, there was no difference in the quality of sleep
between the conditions within trained subjects or between groups.

Exercise metabolism

All trained subjects completed the entire 90 min of exercise on all exercise days. The average total energy cost of the exercise bouts was 6297 kT/d for the
90-min exercise bout. The mean VO sub 2 max, percent VO sub 2 max, and mean workload at which subjects cycled on exercise days (HF and NEB) were 3.3 +/=
0.02 L/min, 74.2 +/= 0.6%, and 1380 +/= 13.2 kg m/min, respectively. There were no differences in any of these exercise variables for any single days of exercise
or between the two exercise conditions of HF and NEB. Additionally, there was no difference in mean exercise HR (HF = 160.2 +/= 0.4, NEB = 160.9 +/= 1.2
beats/min) or mean perceived exertion according to the Borg scale (29) (HF = 14.1 +/= 0.1, NEB = 14.2 +/= 0.4) between conditions. Mean RER during exercise in
the NEB condition was lower when compared with exercise RER during the HF condition (0.913 vs 0.933, respectively).

Energy intake and body weight

Energy intake in trained subjects was greater during HF and PEB (18 384 +/= 862 and 17 769 +/= 849 kJ/d) compared with LF and NEB (10 711 +/= 519 and 10
745 +/= 485 kJ/d). Intake in untrained subjects (10 941 +/= 339 kJ/d) was less than in trained subjects in IIF and PEB, but not different from LF and NEB. There
was no change in body weight when subjects went from HF to LF, but as expected, subjects lost weight during NEB and gained the majority of this weight back
during PEB. When the final day of each condition were compared, mean body weight was lower during NEB than during HF, LF, and PEB (-0.90, -0.75, and -0.80 kg,
respectively). There was no difference between mean body weight when the first and last days of the entire study period were compared. There were no
differences in skinfold-thickness measurements between conditions.

RMR

Comparisons within trained subjects. When mean RMRs between conditions were compared, data from the first day of each condition were not included in data
analyses, which allowed for a washout period between conditions. A 2 X 2 ANOVA revealed that there were significant main effects of exercise and diet on RMR
and an exercise by diet interaction (P < 0.05). Figure 2 shows that the mean RMR in trained subjects was significantly greater during HF than during the other
three conditions--LF, NEB, and PEB. (Figure 2 omitted.)

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Mean RMRs for each day within the entire study period for trained subjects are shown in Figure 3. (Figure 3 omitted.) RMR on HF day 3 (measured 21 h
postexercise after 3 consecutive days of exercise) was elevated when compared with all other days within each condition with the exception of HF day 2. RMR on
LF day 3 (measured 93 h postexercise) was lower than the RMR on each day of HF.

Comparisons benyeen trained and untrained subjects. Comparisons of average RMRs for each of the four conditions in the trained subjects to the LF RMR of the
untrained subjects were made by using ANCOVA, with FFM serving as the covariate. The adjusted mean RMR during HF in trained subjects was significantly
greater (+0.604 kJ/min, P = 0.01) than was the mean RMR in untrained subjects. The RMR values for the trained subjects under the other three conditions were
not different from the RMR values of the untrained subjects.

Comparisons of daily RMR between trained and untrained subjects were again made by covarying with FFM. Because RMRs on days 1 and 2 in untrained
subjects were not different, all comparisons were made by using the mean of these 2 d. The mean CV (SEM) for RMR measured on these 2 d in untrained
subjects was 3.99 +/= 0.74%. The adjusted mean RMR was greater (p </= 0.05) on HF days 1 (+0.401 kJ/min), 2 (+0.520 kJ/min), and 3 (+0.685 kJ/min) and on
LF day 1 (45 h postexercise, +0.337 kJ/min) than was the mean RMR for untrained subjects. There were no other RMR differences.

RMR correlation analyses. Pearson product-moment correlations were examined by using the entire sample of trained and untrained subjects to investigate the
relationship of VO sub 2 max (L/min) and RMR (kJ/min) under two sepate scenarios: 1) when the RMR data for trained subjects were those measured on the final
day of HF and the data for untrained subjects were those measured on the final day of LF (trained and untrained were in different metabolic states) and 2) when
all subjects, trained and untrained, were measured on the final day of the LF condition (trained and untrained were in the same metabolic state). Because both VO
sub 2 max and RMR are influenced by FFM, FFM was partialed out and correlation analyses were performed by using the residuals. Figure 4a demonstrates that
there was a significant correlation between RMR and VO sub 2 max when trained subjects were in HF (r = 0.82, P < 0.001) whereas Figure 4b shows that there
was no correlation between RMR and VO sub 2 max when trained subjects were in LF (r = 0.39, P = 0.13). The change in this relationship between HF and LF can
be explained by the response of the trained subjects to these conditions. When the RMR data from trained subjects only were correlated with VO sub 2 max,
there was a significant correlation when subjects were in HF (r = 0.84, P < 0.01) but no correlation (r = 0.39, P = 0.34) when subjects were in LF. Further correlation
analyses using work (the average exercise work performed during the 3 d of HF) and RMR, both statistically adjusted for FFM, demonstrate a positive
relationship between the amount of work performed and RMR measured 1 d postexercise (r = 0.70, P < 0.05). There was no correlation between RMR and VO sub
2 max within the untrained subjects when these data were analyzed separately.

Plasma measures

The mean values of measured plasma components for trained subjects under each of the four conditions, and untrained subjects in their usual state of LF, are
presented in Table 2. (Table 2 omitted.) Resting plasma catecholamine concentrations were measured for trained subjects only. Additionally, there was one
untrained subject from whom a blood sample could not be obtained. There were no differences in hematocrit between conditions and, therefore, no adjustments
for plasma volume were made.

There was a main effect of exercise on norepinephrine (P < 0.05) and an exercise by diet interaction (P < 0.01). Norepinephrine concentration was greater during
HF (measured 21 h after completion of the last exercise bout) than during NEB and PEB. Also, there was a trend for norepinephrine to be higher during HF than
during LF (P < 0.06). Seven of eight subjects had increased norepinephrine concentrations during HF when compared with their LF concentrations. The remaining
subject displayed a very high norepinephrine concentration during LF (the highest concentration of any condition for that individual). When norepinephrine data
were analyzed excluding this outlier, the difference between concentrations during HF and LF was highly significant (P = 0.007). There were no differences in
epinephrine concentration between conditions and no effects of exercise, diet, or their interaction on epinephrine concentration.

Correlation analyses were performed to investigate the relationship between the change from the mean norepinephrine concentration and the change from the
mean RMR under each condition (eight subjects, 16 data points). The analysis of data from the HF and LF conditions (the zero energy balance conditions that
most closely represent the habitual conditions of trained and untrained subjects, respectively) revealed a positive correlation between the change in
norepinephrine and the change in RMR (r = 0.82, P < 0.01, Figure 5). (Figure 5 omitted.) This positive relationship persisted when the analyses were performed by
using the simple difference in these variables between HF and LF conditions (eight data points, r = 0.72, P < 0.05). Additionally, significant correlations were
observed when analyses were performed by using the data from HF and NEB (r = 0.56, P < 0.05) and LF and PEB (r = -0.54, P < 0.05).

The plasma concentrations of triiodothyronine, insulin, and glucose are also presented in Table 2. (Table 2 omitted.) There were main effects of diet and exercise
on triiodothyronine but no exercise by diet interaction. Plasma triiodothyronine concentrations were elevated during PEB when compared with NEB in trained
subjects. There were no differences in triiodothyronine between untrained and trained subjects. Plasma insulin concentration was decreased during NEB when
compared with LF and PEB and there was a nonsignificant trend for it to be lower than during HF (P = 0.08). The insulin concentration was higher in untrained
subjects than in trained subjects during HF, NEB, and PEB, and nearly greater than during LF (P = 0.06). There were no differences in CPK or glucose
concentrations between conditions in trained subjects or between the trained and untrained groups.

Discussion

The discordant findings among previous studies that have sought to determine whether or not exercise training is associated with an increased RMR, led us to
explore several factors (ie, the magnitude of energy intake and the time interval between RMR measurement and previous exercise), which we believed could help
clarify the nature of this relationship. Specifically, we hypothesized that among trained endurance athletes, acute HF (high energy turnover characterized by high
intake and exercise energy expenditure) would elevate RMR compared with other LF conditions. We also hypothesized that only when trained subjects were in an
acute state of HF would they exhibit higher RMR values than untrained subjects. Our findings in this study support these hypotheses.

Comparisons within trained subjects

Mean RMR was greater during the period when subjects were exercising daily while consuming a controlled diet adequate to maintain energy balance than
during the 3 d of no exercise and lowered energy intake necessary to achieve energy balance. Mean RMR measured on the final day of HF (21 h after 3
consecutive days of exercise) was ==11% (0.558 kJ/min) higher than when measured on the final day of LF (96 h postexercise), with every subject having a
higher RMR during HF. The average RMR during HF was also higher than during the conditions of exercise with reduced energy intake (NEB), and no exercise with
excess energy intake (PEB). Thus, it appears that the combination of energy expended and consumed (total energy flux) is an important contributor to RMR. It
was shown previously that acute PEB and NEB can influence RMR (33-35). Data from the present study now indicate that the magnitude of energy flux (turnover)
may influence RMR even when subjects are kept in energy balance.

The mechanisms responsible for the observed elevation of RMR during HF remain speculative and might include postexercise changes in muscle cell structure
(36), immune system response (37, 38), neuroendocrine function (5, 39), and substrate cycling (25, 40). In the present study, no attempt was made to directly
assess changes in muscle cell structure (eg, cellular damage) or changes in immune function after exercise. The factors responsible for the higher RMR during
HF would likely also be present during the NEB condition. However, any tendency for an elevated RMR under the latter condition might be negated by the energy
deficit condition, which is associated with a reduction in RMR. The fact that CPK concentrations were not different between conditions suggests that severe
tissue damage was not responsible for the differences in RMR.

The thermogenic hormones triiodothyronine, norepinephrine, and epinephrine were measured under each condition to assess their possible roles in explaining
observed differences in RMR among conditions. Because triiodothyronine is acutely influenced by over- or underfeeding, it is important to focus on the plasma
triiodothyronine concentrations in the two conditions in which attempts were made to maintain energy balance (HF and LF). Although RMR was different
between these conditions, triiodothyronine concentrations were not. Therefore, it appears that in this study the observed differences in RMR among conditions
was disassociated from plasma triiodothyronine concentrations. Changes in norepinephrine concentrations were observed between conditions, with
norepinephrine being greater during HF than during NEB and PEB and nearly greater than during LF (P = 0.06). As Figure 5 illustrates, there was a correlation
between the change from the mean in NE concentration and the change from the mean in RMR between the energy balance conditions of HF and LF (r = 0.82, P <
0.001). Although a cause-and-effect relationship cannot be established, this finding suggests that even 21 h postexercise norepinephrine may contribute to
observed elevations in RMR during HF conditions. Similar findings have been reported previously (13).

Comparisons benyeen trained and untrained subjects

RMR, statistically adjusted for FFM, was greater on every day of HF and on day 1 of LF (measured 45 h postexercise) in trained subjects than was RMR in
untrained subjects. There were no other differences between RMR in untrained subjects and RMR measured on any other day under any other condition in
trained subjects. Additionally, a change in the relationship of RMR (kJ/min) and VO sub 2 max (L/min) when subjects went from HF to LF conditions was
observed in the present study. The residuals of RMR and VO sub 2 max (adjusted for FFM) were correlated (r = 0.82, P < 0.001) when all subjects (trained and
untrained) were evaluated and when the RMR data of the trained subjects were those measured the day after the last HF exercise bout (21 h postexercise)
(Figure 4a). Poehlman et al (2) found a similar correlation (r = 0.77, P < 0.01) between RMR (kcal X kg body wt sup -1 X h sup -1 ) and VO sub 2 max (mL X kg body
wt sup -1 X min sup -1 ) in males of varying fitness levels when RMR was measured 24 h postexercise. On the basis of these data, they suggested that chronic
exercise training results in an adaptive, long-term elevation in RMR. In the present study, however, RMR and VO sub 2 max were no longer correlated (r = 0.39, P =
0.13) when the RMR data of trained subjects were those measured on the final day of LF (93 h postexercise) (Figure 4b). These data suggest that observed
differences in RMR between trained and untrained individuals in several previous studies (1-3) may have resulted from residual perturbations of acute exercise

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rather than to an adaptive elevation in RMR produced by chronic exercise. This point is further supported by the significant correlation (r = 0.70, P < 0.05)
between the amount of physical work performed during the 3 d of high flux and RMR measured ==1 d after exercise. Additionally, these results suggests that a
period longer than 45 h (possibly 3 d), in which trained subjects abstain from exercise before RMR testing, may be necessary to obtain RMR values that are not
influenced by the last exercise bout.

An important methodological issue regarding this study should be addressed. It is virtually impossible to precisely quantify energy intake and expenditure in
free-living subjects. We wanted the trained subjects to be in energy balance during HF and LF conditions. Our method of calculating energy requirements during
LF conditions (RMR X 1.5, which also served as a baseline for energy intake during HF) provided only a crude estimation. The body weight and triiodothyronine
data suggest, however, that subjects were in approximate energy balance during HF and LF. Body weights and triiodothyronine concentrations did not change
when subjects went from HF to LF (6 d total) despite energy expenditure and intake being ==7500 kJ/d greater during HF. Additionally, triiodothyronine
concentrations were intermediate during HF and LF when compared with NEB and PEB, further suggesting that deviations from energy balance under these two
conditions were small. For the PEB and NEB conditions, body weights and triiodothyronine concentrations varied in a predictable manner. Plasma
triiodothyronine concentrations are responsive to acute changes in energy balance, increasing with overfeeding and decreasing with underfeeding (41, 42). These
data suggest that our subjects were in energy deficit during NEB and in energy surplus during PEB, although the magnitude of each could not be precisely
quantified.

Additionally, three important points need to be made with respect to the PEB and NEB conditions in this study: 1) Subjects remained in positive energy balance
for only 2 d (compared with 3 d for each of the other conditions) and we cannot predict what the response of RMR would have been to 3 d of overfeeding, 2) PEB
always followed NEB. Therefore, PEB was actually a refeeding condition, which may have influenced the RMR response during this condition, and 3) NEB was
achieved through exercise rather than through dietary restriction only. It is possible that the exercise during this condition may have attenuated the decrease in
RMR observed during energy restriction in some earlier studies. Also, note that although this study was designed and analyzed by using a 2 X 2 factorial design
within the trained subjects, it could possibly be viewed as having an incomplete 3 x 2 design. The two missing conditions under this scenario are 1) a condition
as described above in which NEB is attained by reducing energy intake while the subjects abstain from exercise (NEB without exercise), and 2) a PEB condition
attained by overfeeding exercising subjects (PEB with exercise). It is impossible to predict from the present study what the metabolic responses to these
conditions may be.

In conclusion, data from the present study indicate that RMR in exercise-trained individuals is influenced by the total flux of energy through the body (as
indicated by the significant effect of the exercise by diet interaction on RMR), even under conditions of approximate energy balance. The data further indicate
that RMR in trained vs untrained subjects is elevated under acute conditions of high exercise energy expenditure and high energy intake, but that the elevation is
attenuated as the time interval increases from the last exercise bout to the measurement of RMR. These data suggest that RMR may be chronically elevated in
individuals who engage in daily high-intensity, prolonged exercise owing to an effect of acute exercise rather than to an adaptation to chronic exercise.

From the Human Energy Laboratory, Department of Food Science and Human Nutrition, Colorado State University, Fort Collins. Supported by the Colorado
Agricultural Experiment Station, project no. 616, Colorado State University. Reprints not available. Address correspondence to CL Melby, Department of Food
Science and Human Nutrition, Human Energy Laboratory, 226 Gifford Building, Fort Collins, CO 90523.

Received January 13, 1994.

Accepted for publication September 27, 1994.

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