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1 66 Journal of Exercise Physiologyonline August 2018 Volume 21 Number 4 Editor-in-Chief Official Research Journal of Tommy the American Boone, PhD, Society MBA of Review Exercise Board Physiologists Todd Astorino, PhD Julien Baker, ISSN PhD Steve Brock, PhD Lance Dalleck, PhD Eric Goulet, PhD Robert Gotshall, PhD Alexander Hutchison, PhD M. Knight-Maloney, PhD Len Kravitz, PhD James Laskin, PhD Yit Aun Lim, PhD Lonnie Lowery, PhD Derek Marks, PhD Cristine Mermier, PhD Robert Robergs, PhD Chantal Vella, PhD Dale Wagner, PhD Frank Wyatt, PhD Ben Zhou, PhD Official Research Journal of the American Society of Exercise Physiologists ISSN JEPonline Impact of Daily Exercise Compared to Exercise on Alternating Days on Excess Post-Exercise Oxygen Consumption Zachary Zeigler, Malachi Votaw, Connor Dreos, Lydia Durnil, Jamie Terran, Ezekiel Palmani, Danielle Akin, Trevor Nordin College of Science, Engineering, and Technology, Grand Canyon University, Pheonix, AZ ABSTRACT Zeigler Z, Votaw M, Dreos C, Durnil L, Terran J, Palmani E, Akin D, Nordin T. Impact of Daily Exercise Compared to Exercise on Alternating Days on Excess Post-Exercise Oxygen Consumption. JEPonline 2018;21(4): The purpose of this study was to compare daily exercise (DE) to exercise performed on alternating days (AE) and a control (CON) on excess post-exercise oxygen consumption (EPOC). Sedentary men aged 18 to 30 yrs were the subjects in this three-arm randomized cross-over trial. The primary comparison was the magnitude of EPOC between the three groups (DE, AE, and CON) and within each group over time (day 1, 2, 3, and 4). Both exercise groups were prescribed the same exercise intensity (70 to 75% HR max), and total duration of exercise (90 min) on a cycle ergometer. The DE group performed exercise on three consecutive days (3 bouts of 30 min). AE performed exercise on two alternating days (2 bouts of 45 min). After exercise the subjects remained in the laboratory for 1 hr while VO 2 was measured continuously. Nine young (21 ± 2) overweight (28 ± 4 kg m -2 ) male subjects completed the study. There was no significant difference on VO 2 or RER over 4 d for CON (P=0.561, P=0.179). EPOC did not accumulate over time during DE (P=0.561). However, DE did produce an overall greater EPOC than CON (P=0.009) while AE did not (P=0.106) (CON: 17 ± 3 L min -1, AE: 19 ± 2 L min -1, DE: 20 ± 3 L min -1 ). RER did not accumulate over time (P>0.05). In conclusion, EPOC was not impacted by back to back exercise sessions. However, when combing all exercise sessions, DE produced more overall EPOC than AE, thus DE could be superior to AE on total energy expenditure. Key Words: Acute Exercise, Daily Exercise, EPOC, Weight Loss

2 67 INTRODUCTION It is well established that following exercise, oxygen consumption is increased beyond what it was prior to the exercise session. This has been termed by Gaesser and Brooks as Excess Post-Exercise Oxygen Consumption (EPOC) (8). Many have now suggested that EPOC may play a greater role in weight loss than once was thought, thus highlighting the need to evaluate how to increase it (4,13). Exercise duration and intensity have been identified as key factors in determining the magnitude of EPOC. It is well known that exercise intensity is curvilinearly related to EPOC with stark rises when intensity is beyond 75% of VO 2 max (5). The results of the impact of exercise duration on EPOC are equivocal. Nevertheless, much data suggests a linear relationship is seen with increased duration of exercise provoking the increase in EPOC (3). Many studies and reviews have assessed the impact of exercise intensity and duration on EPOC, yet there is a paucity of research on the impact of exercise frequency on EPOC (5). Split exercise sessions have shown greater EPOC values compared to one exercise session (1,11). For example, when comparing 50 min of continuous running to two 25-min sessions, a greater total EPOC was found for the split group (11). However, completing two exercise sessions in one day may not be feasible for many. EPOC has been shown to be elevated for upwards of 36 to 48 hrs (23). This suggests that if daily exercise were undertaken, subsequent exercise sessions could be conducted when oxygen consumption is elevated and may drive oxygen consumption higher, somewhat of an accumulating effect. Thus, because physical activity and exercise are a fundamental component of a weight loss program, it is of interest to assess the impact of exercise frequency, independent of exercise energy expenditure, on EPOC. Whether or not exercise performed on consecutive days would lead to an accumulating EPOC effect has yet to be determined. If an accumulation effect is seen, it is unknown if daily exercise (DE) is superior to exercise performed on alternating days (AE) when compared to a non-exercise control (CON). The purpose of this study was to compare the impact of 90 min of moderate intensity exercise (70 to 75% heart rate (HR) max) split up into either two nonconsecutive 45-min exercise sessions or three consecutive 30-min exercise sessions on EPOC and substrate utilization. It is hypothesized that EPOC will accumulate such that EPOC will be the highest during the last training session of DE. We also hypothesize that fat usage will be increased during the EPOC phase to a greater degree in the DE condition. Our hypothesis is formed by examining prior research showing an additive effect of consecutive exercise on glycemic control (15) triglycerides (9), and blood pressure (24). METHODS The present study was a three-arm randomized experimental design. The Grand Canyon University Institutional Review Board approved this study (IRB # ). The primary comparison was the magnitude of EPOC and change in substrate utilization between the 3 groups and within each group over time. Figure 1 gives graphical depiction of the research design. This cross-over experimental trial consisted of subjects randomized into 3 groups (i.e., conditions): (a) DE; (b) AE; and (c) a non-exercise control (CON). After completion of the first condition, subjects were randomly assigned to the last two conditions. All subjects completed each condition. A period of 1 wk was used as the wash-out time, thus each

3 condition was performed 1 wk apart. Exercise duration between groups was matched at 90 min split into either three 30-min sessions (DE) or two 45-min sessions (AE). Exercise was performed on a cycle ergometer at an intensity of 70 to 75% HR max. Gas analysis was measured continuously for 1 hr following each exercise session. 68 Subjects Inclusion Criteria Nine sedentary male subjects (aged 18 to 30 yrs) who did not meet the physical activity guidelines of 150 min wk -1 as determined by the International Physical Activity Questionnaire (18) were recruited to participate in this experimental trial. Exclusion Criteria The following exclusion criteria were adopted: (a) having known cardiovascular, pulmonary, renal or metabolic disease; (b) having symptoms suggestive of these diseases; (c) current smoker; and (c) having orthopedic limitations for performing physical activity. Consent Process The informed consent was given to potential participants and adequate time was allowed for them to read over it. Once the potential participant finished reading the informed consent, the investigator gave the individual an opportunity to ask any questions. Procedures The three conditions were assigned randomly and performed on the same days of the week and time of day to minimize differences in subject s daily routines and account for diurnal variation. During all three visits the subjects were asked to refrain from physical activity for 24 hrs prior, and to come to the lab in a fasted state ( 6 hrs without food). Food and beverage consumption was not altered but the subjects were given a food recall form and asked to eat

4 and drink the same items during the day prior to the lab visits. Following subject consent and prior to the first condition, body composition, height, weight, and VO 2 peak were assessed. Baseline Measurements Once the subjects met the inclusion criteria and signed the consent, baseline measurements were determined. Body composition was measured via whole body air displacement plethysmography (Bod Pod). The subjects were asked to wear a bathing suit or tight-fitting shorts for testing and remove all jewelry. They were first weighed on a scale in bare feet. Then, the subjects were asked to place a cap on their heads to cover their hair. The subjects sat in the bod pod for 2 to 3 measurements of 50 sec each. Following the Bod Pod, a VO 2 peak test was completed. The subjects VO 2 peak was assessed with the Vmax metabolic cart (CareFusion, Franklin Lakes, NJ). Gas analyzers were calibrated with a certified mixture of oxygen at 17.01% and carbon dioxide at 5.00% balanced with nitrogen. Flow and volume were calibrated with a 3-L syringe (Hans Rudolph TM, Kansas, MO, USA). The test was a ramp cycle ergometer protocol. Subjects began on the cycle ergometer at 25 watts for 5 min for a warm-up. Every minute the wattage increased by 20 watts until the subject reached volitional failure or requested to stop. The subjects were given a 5-min cool down period on the ergometer at 25 watts. The two consecutive highest 15-sec VO 2 values were taken as VO 2 peak. Experimental Approach Subjects completed each of the three conditions: (a) non-exercise control (CON); (b) daily exercise (DE); and (c) exercise on alternating days (AE). All subjects were prescribed the same exercise intensity (70 to 75% of HR max achieved from the VO 2 peak assessment), and total duration of exercise (90 min). Exercise was performed on a cycle ergometer. The only difference between the two exercise conditions was that the DE condition performed exercise on 3 consecutive days (3 bouts of 30 min) compared to AE that performed exercise on 2 alternating days (2 bouts of 45 min). The subjects were required to perform exercise at the laboratory under supervision of one of the researchers. To ensure adherence to the exercise intensity, HR monitors were worn by the subjects and a researcher monitored HR every 5 min during the exercise session making changes to alter wattage if needed. Subjects remained in the lab for 1 hr following exercise to collect oxygen consumption data. Upon completion of exercise, the subjects were asked to sit quietly of which they were free to surf the internet, watch a movie, or read. During the non-exercise day of AE, subjects sat quietly for 1.5 hrs while gas analysis was being completed. For the control visit, subjects were asked to sit quietly at the lab for 1.5 hrs. During this time, they were free to surf the internet, watch a movie, or read. However, the subjects were not allowed to walk or stand during this time. Gas analysis was measured continuously. All subjects, and for all conditions, were required to again return to the lab a 4th day to assess resting oxygen consumption one last time. Upon arrival to the laboratory, the subjects were asked to sit quietly in a calm environment for 15 min. Then, gas analysis was done for an additional 20 min with the first 5 min being discarded. The last 15 min was averaged and used as the final resting VO 2. 69

5 70 Measurement of EPOC Subjects were asked to come to the laboratory at the same time during each of the three conditions. They were told to fast for >6 hrs, and to have abstained from any physical activity for 24 hrs. Upon arrival to the laboratory, the subjects were asked to sit quietly in a calm environment for 15 min. Then, the subjects were fitted for a two-way non-rebreathing mask that they were required to wear at baseline, during exercise, and for 1 hr following. Subjects continued to sit quietly for 20 min of which this 20-min average was considered baseline. The first 5 min of baseline data were discarded to allow subjects to relax and ventilation data to normalize. Upon completion of exercise, the subjects were asked to immediately sit down. Gas analysis continued to be collected for 1 hr following exercise. The gas while sitting was averaged into 5-min averages. Total EPOC was calculated as the difference between oxygen consumption during the 1 hr post-exercise period after the exercise conditions minus total oxygen consumption during the corresponding time control trial. Statistical Analyses All statistical analyses were done using SPSS software version 23 (SPSS 23.0 IBM Corporation, Armonk, New York, USA). Data are expressed as means ± standard deviation (SD) unless otherwise specified. Data were analyzed for normality and values with skewed distribution were transformed to achieve normality. Descriptive statistics were used for the demographics of the subjects. All P values were calculated assuming two-tailed hypothesis; P<0.05 was considered statistically significant. Statistical analysis included gas analysis from data collected during baseline and the 1 hr following the exercise bout. Linear mixed models were used to detect group (CON, AE, and DE) and time (between day 1 through day 4) differences in oxygen consumption over the 1 hr measurement period. Both within condition over time effects, and between conditions over time effects were analyzed. The analysis was conducted in a hierarchical fashion using Restricted Maximum Likelihood model and variance components covariance error structure. Both fixed and random effects were explored in the model. Treatment condition, age, and BMI were used as fixed effects. Unless specified, post-hoc analysis was assessed using Bonferoni to examine group and time differences in oxygen consumption data. One-way ANOVA was used to test for baseline VO 2 differences (Homogeneity of Variance was analyzed and correction made if applicable). RESULTS Nine subjects completed the study. Table 1 presents demographic characteristics of the subjects. Briefly, the subjects were relatively young, overweight, and on the lower end of the fitness spectrum for their age (21).

6 71 Table 1. Descriptive Data of the Subjects (N=9). Mean ± SD Height (cm) Weight (kg) BMI (kg m -2 ) Body Fat (%) Age (yrs) VO 2 Peak (ml kg -1 min -1 ) ± ± ± ± ± ± 7.3 EPOC Data Table 2 indicates that during the CON condition there were no statistical differences for VO 2 (P=0.561) between days 1 through 4. Table 2 also illustrates that there was not a significant within condition over time effect for CON (P=0.561) or DE (P=0.236). There was a significant within condition over time effect for alternating exercise (P=0.005) such that EPOC on day one averaged the highest. Table 2. Mean VO 2 Within Each Condition Over the Four Measurement Days. Condition Day 1 Day 2 Day 3 Day 4 P-value CON 3.4 ± ± ± ± AE 3.8 ± 1 a 3.3 ±.6 b 3.5 ± 2 a 3.5 ± 1 ab DE 3.9 ± ± ± ± VO 2 expressed as ml kg -1 min -1, Different letters signify statistical difference from each other at a P<0.05 Figure 2 (total EPOC per day) illustrates that overall, there was a significant condition (P<0.001) effect for VO 2 such that VO 2 averaged the highest during the DE condition. Post hoc analysis revealed that differences were found between the CON and the DE (3.4 ±.89 ml kg -1 min -1 vs. 3.9 ± 1.4 ml kg -1 min -1-1 ; P<0.001), not CON and AE (3.4 ±.89 ml kg -1 min vs. 3.5 ± 1.2 ml kg -1 min -1 ; P=0.194). The DE EPOC was 2.4 L min -1 per day higher than the CON (P=0.009). The AE condition was not higher than the CON condition when averaged over the period of the intervention (P=0.317). The DE and the AE were no different from each other (P=0.306).

7 72 Table 3 shows there was a significant condition effect for day 1 (P<0.001) such that both the exercise conditions produced a higher VO 2 when compared to the CON condition. Also, there was a significant condition effect for day 2 (P<0.001) such that the DE had the highest average VO 2. There was a trend during day 3 and for day 4 for the DE to be higher than the CON (P=0.070 and P=0.079, respectively). Table 3. Mean VO 2 Within Each Day and Each Condition. Day CON AE DE P-value ± 1 a 3.8 ± 1 b 3.9 ± 1 b < ±.8 a 3.3 ±.6 a 3.8 ± 1 b < ± ± ± ± ± ± VO 2 expressed as ml kg -1 min -1, Different letters signify statistical difference from each other at a P<0.05

8 73 Substrate Utilization During the CON condition there were no statistical differences for RER (P=0.179) between days 1 through 4. Figure 3 illustrates that for the overall average RER for all days, there was a significant condition effect such that RER averaged lower on both exercise conditions when compared to the CON (P<0.001). The AE condition produced the lowest RER value. Post hoc analysis revealed that differences were seen between CON and AE (.86 ±.1 vs..82 ±.1, P<0.001), and a non-significant trend between CON and DE (.86 ±.1 vs..86 ±.1, P=0.066). Table 4 indicates there was not a significant within condition over time effect for CON (P=0.179). There was a significant within condition over time effect for AE (P<0.001) such that day 4 had the lowest RER. There was also a significant within condition over time effect for the DE condition such that all days had a lower RER compared to day 4 (P=0.002).

9 74 Table 4. Mean RER Within Each Condition Over the Four Measurement Days. Condition Day 1 Day 2 Day 3 Day 4 P-value CON.87 ±.1.85 ±.1.85 ±.1.87 ± AE.82 ±.1 a.84 ±.1 a.82 ± 1 a.76 ±.1 b DE.82 ±.1 a.83 ±.1 a.81 ± 1 a.89 ±.1 b VO 2 expressed as ml kg -1 min -1, Different letters signify statistical difference from each other at a P<0.05 There was a significant difference on day 1 (P<0.001) such that both the exercise conditions produced a lower RER when compared to the CON condition. There was not a significant RER difference on day 2 (P=0.365). During day 3 both exercise conditions produced lower RER values when compared to the CON (P=0.007). Additionally, during day 4 there was a significant RER difference such that the AE condition had the lowest RER value (P<0.001). DISCUSSION The main findings of this study are the following: (a) There was not an accumulating impact of DE on VO 2 or RER; (b) The added exercise duration on the AE condition did not increase EPOC greater than the shorter durations of the DE condition; and (c) DE, when expressed in L min -1 and averaged over the experimental protocol produced a significantly greater amount of EPOC when compared to the CON condition while the AE condition did not. The absence of an accumulating effect of DE does not come as a huge surprise. EPOC increases in a curvilinear fashion for exercise intensity (6,12). The absence of sustained EPOC after low intensity exercise and/or duration is consistent with prior research (6). The current protocol took either 30 min or 45 min to complete and the intensity was approximately 75% of the subjects HR max. Intensities above 50 to 60% of VO 2 max are typically required to induce EPOC that lasts several hours (6). Thus, the elevated oxygen consumption more than likely returned to pre-exercise values prior to the onset of another exercise session. The present protocol was used to emulate the current public guidelines (10). Exercise of greater intensities may produce different outcomes. The mechanisms of EPOC can be broken down into two phases. The Rapid EPOC Component is well understood to be due to replenishment of O 2 stores in blood and muscle, resynthesis of adenosine triphosphate and creatine phosphate, lactate removal, increased body temperature, circulation, and ventilation (2,3,8). These factors are more than likely to be the mechanisms responsible for the increased VO 2 of the DE condition. The fact that AE did not exercise on day 2 combined with an inability for the longer exercise duration of AE to make up for the missed exercise session is more than likely the responsible factor for DE being superior to AE.

10 75 The observation that a longer duration of exercise on the AE condition did not produce greater EPOC is of interest. Chad and Wenger (6) found that when exercise duration was increased from 30 min to 45 min there was a 2.3-fold increase in EPOC. On the other hand, Sedlock (20) conducted a similar study and found EPOC did not change with altered exercise durations. However, in the Sedlock s study the intensity was 40 to 60% of VO 2 max compared to 70% used by Chad and Wegner. The current study used an intensity that is closer to Sedlock s work, and it could be that the intensity was not high enough to maximize the benefits of an increase in exercise duration. The interaction between exercise duration and intensity is not well understood, and it is extremely hard to separate the two out from each other. It is important to note that total exercise duration in the current study was matched. It is already established that a dose response is present as to the amount of exercise undertaken and fat loss (22). For example, those who exercised on a treadmill for longer time increased fat loss to a greater extent than those exercising on a treadmill for shorter periods of time (19). However, it is important to point out that the current study was addressing the role of exercise frequency that was independent of total exercise time. It was hypothesized that fat utilization during the DE recovery period would be greater than the AE and the CON conditions. This hypothesis was put forth because repeated exercise provokes an increase in muscle contraction (16) that is thought to increase the uptake of circulating glucose concentrations (7). Glycogen depletion may take place shifting substrate utilization to favor fat in the exercise recovery period. However, both conditions elicited lower RERs compared to the CON condition, which was probably due to the increase in glycogen depletion above the non-exercise CON. The DE condition was not any better than the AE on fat utilization during the recovery period. Interestingly, on day 4, RER was the lowest during the AE condition. It could be possible that the longer duration of the exercise bout on the day preceding caused the substrate shift. We did not measure glucose concentration. Hence, we can only speculate that both exercise conditions increased glucose uptake, which resulted in an increase in fat utilization following the exercise. The RER values following exercise must be taken in context however. It is known that the first hour following exercise produces potential instability of the plasma bicarbonate pools, making RER values questionable during this time (14). Limitations in this Study First, the age of the subjects in this study was relatively young and healthy. Divergent results may be seen in diseased or older populations. Second, the subjects were free to eat on their own and eat what they liked. The subjects were asked to eat the same thing before coming to the lab and between conditions and a dietary recall was given to aid in this endeavor. Third, EPOC was measured only for 1 hr following exercise. Although unlikely, it could be that additional oxygen consumption may have been missed due to this short measurement time window. Zeigler and Swan (25) measured oxygen consumption for 3 hrs following a moderate bout of exercise and found that virtually all EPOC was accumulated during the first 15 to 30 min post-exercise.

11 76 CONCLUSIONS The DE condition did not cause EPOC to accumulate but it was superior to the AE condition on total daily EPOC (L) compared to the AE condition. In context of the external relevance of our findings, the additional EPOC energy expenditure of the DE contributed to a net increase of 12 kcals d -1 compared to the CON condition. The AE did not reach statistical significance with EPOC when compared to CON. If DE were to be adopted that would be roughly an additional 85 cal wk -1 or 1.2 lbs yr -1. When considering that average weight gain for middle aged adults is a mere 2 lbs yr -1 (17), calories from the DE condition EPOC could account for over half of that. Because EPOC increases linearly with exercise intensities, future research should assess if higher exercise intensity would alter these findings. ACKNOWLEDGMENTS We would like to thank the group of subjects for their participation. Address for correspondence: Zachary S. Zeigler, PhD, College of Science, Engineering, and Technology, Grand Canyon University, Phoenix, AZ. zachary.zeigler@gcu.edu REFERENCES 1. Almuzaini KS, Potteiger JA, Green SB. Effects of split exercise sessions on excess postexercise oxygen consumption and resting metabolic rate. Canadian J Appl Physiol. 1998;23(5): Bahr R, Sejersted OM. Effect of intensity of exercise on excess postexercise O 2 consumption. Metab Clin Exp. 1991;40(8): Bahr R. Excess postexercise oxygen consumption--magnitude, mechanisms and practical implications. Acta Physiol Scand Suppl. 1992;605: Bergman BC, Brooks GA. Respiratory gas-exchange ratios during graded exercise in fed and fasted trained and untrained men. J Appl Physiol. 1999;86(2): Borsheim E, Bahr R. Effect of exercise intensity, duration and mode on post-exercise oxygen consumption. Sports Med. 2003;33(14): Chad KE, Wenger HA. The effect of exercise duration on the exercise and postexercise oxygen consumption. Can J Sport Sci. 1988;13(4): Di Loreto C, Ranchelli A, Lucidi P, et al. Effects of whole-body vibration exercise on the endocrine system of healthy men. J Endocrinol Invest. 2004;27(4): Gaesser GA, Brooks GA. Metabolic bases of excess post-exercise oxygen consumption: A review. Med Sci Sports Exerc. 1984;16(1):29-43.

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