In te rn a tio n a l J o u rn a l o f S p o rts P h y sio lo g y a n d P erfo rm a nce , 2019, 14, 1140-1146

https://doi.Org/10.1123/ijspp.2018-0854 Human Kinetics £QU

© 2019 Human Kinetics, Inc. ORIGINAL INVESTIGATION

Diurnal Variation of Maximal Fat-Oxidation Rate

in Trained Male Athletes
Francisco J. Amaro-Gahete, Lucas Jurado-Fasoli, Alejandro R. Trivino, Guillermo Sanchez-Delgado,
Alejandro De-la-O, Jorn W. Helge, and Jonatan R. Ruiz

Purpose: To analyze the diurnal variation of maximal fat oxidation (MFO) and the intensity that elicits MFO (Fatmax) in trained
male athletes. Methods: A total of 12 endurance-trained male athletes age 24.7 (4.1) y participated in the study. The authors
measured MFO, Fatmax, maximum oxygen uptake (V 0 2max), and V 0 2 percentage at ventilatory threshold 2 with a graded
exercise protocol performed on 2 days separated by I wk. One test was performed in the morning and the other in the afternoon.
The authors assessed the participants’ chronotype using the HOME questionnaire. Results: MFO and Fatmax were greater in the
afternoon than in the morning (A= 13%, P < .001 and A = 6%, P = .001, respectively), whereas there were similar V 0 2max and
ventilatory threshold 2 in the morning, than in the afternoon test (A = 0.2%, /* = . 158 and A = 7%, P = .650, respectively). There
was a strong positive association between V 0 2max and MFO in both morning and afternoon assessments (R2 = .783, P = .001 and
R~=. 663, P c . 001, respectively). Similarly, there was a positive association between V 0 2max and Fatmax in both morning and
afternoon assessments (P2 = .406, P = .024 and P 2 = .414, P = ,0 2 6 , respectively). Conclusion: MFO and Fatmax may partially
explain some of the observed diurnal variation in the performance of endurance sports.

Keywords: Fatmax, V 0 2max, circadian rhythm, resistance, fuel oxidation

Carbohydrates and fats are the primary substrates oxidized to
fuel energy metabolism during exercise.1 Humans predominantly
store carbohydrates as glycogen in skeletal muscle and the liver, and
- 4 g circulating in plasma as glucose.2 However, these storage
depots are limited, whereas human fat energy storage is effectively
unlimited during prolonged exercise.3 Therefore, the capacity to
adapt fuel oxidation to fuel availability (known as metabolic
flexibility) is a key determinant of endurance sport performance.4
Therefore, the maximal fat oxidation (MFO) capacity during
a graded exercise protocol is considered an important factor in
endurance exercise performance as well and in cardiovascular
health.5 Moreover, another important variable is the exercise inten­
sity at which MFO occurs, so called Fatmax. Both MFO and Fatmax,
together with maximum oxygen uptake (V 0 2max), V 0 2max per­
centage at ventilatory threshold 2 (VT2), and running economy are
considered important outcomes in endurance sports performance.5,6
Endurance sport performance, specifically running and cycling
performance, seems to present diurnal variation, being higher in the
afternoon than in the morning.7 This might be explained by a higher
body temperature, higher neural activation and contractile properties
of the skeletal muscle, or higher plasma catecholamine concentra­
tions immediately after exercise in the afternoon than in the morn­
ing.8,9 It has been observed a higher MFO and Fatmax in the afternoon
than in the morning in nonathlete male students10 and in untrained
normal weight and obese individuals.11 Whether the observed
diurnal variations in MFO and Fatmax also apply to endurance-
trained athletes is unknown. Moreover, despite that it has been
Amaro-Gahete, Jurado-Fasoli. Trivino, and De-la-0 are with the Dept of Medical
Physiology, School of Medicine, and Amaro-Gahete, Sanchez-Delgado, and Ruiz,
the Dept of Physical Education and Sport, Faculty of Sport Sciences, University of
Granada, Granada, Spain. Helge is with the Dept of Biomedical Sciences, Faculty of
Health Sciences, University of Copenhagen, Copenhagen, Denmark. Amaro-Gahete
(amarof@ugr.es) is corresponding author.
1140
reported that individuals with higher V 0 2max present greater muscle
capacity to oxidize fat,12,13 the relation of V 0 2max with MFO and
Fatmax in endurance-trained male athletes remains to be elucidated.
Therefore, the aim of this study was to analyze the diurnal
variations of MFO and Fatmax in endurance-trained male athletes.
We also determined the diurnal variations of V 0 2max and VT2
and examined the association of V 0 2max and VT2 with MFO and
Fatmax. We hypothesized that MFO, Fatmax, V 0 2max, and VT2 are
higher in the afternoon than in the morning, and that V 0 2max and
VT2 are positively associated with MFO and Fatmax in endurance-
trained male athletes.
Methods
Subjects
A total of 14 endurance-trained male athletes age 18 to 32 years
voluntarily participated in the study. Two out of 14 participants did
not meet the predetermined conditions (see below) for MFO and
Fatmax measurements on one of the testing days, and were retro­
spectively excluded from further statistical analyses. All athletes
had extensive experience in endurance events and had a minimum
of 2 years of cycling or running practice as a part of their main
training schedule. They had a body mass index between 18 and
25 k g /n r, were nonsmokers, did not take any medication, and had
no acute or chronic illness. All participants provided written
informed consent to participate in the study, which was performed
in accordance with the Declaration of Helsinki. An ethical approval
was obtained from the University of Granada research ethics
committee (ethical approval code no 507/CEIH/2018).
Design and Methodology
The study was conducted between March and April 2018. MFO
and Fatmax were measured on 2 different days separated by 1 week.

Measurements were perfonned between 8 AM and 11 AM (MFO-
moming, Fatmax-moming, V 0 2max-rnoming, and VT2-moming)
and between 5 PM and 8 PM in the afternoon (MFO-aftemoon,
Fatmax-aftemoon, V 0 2max- afternoon, and VT2-afternoon). The test
order (morning vs afternoon) was randomized using a simple
random function of the software MS Excel for Windows® (Micro­
soft, Redmond, WA). Participants arrived at the laboratory by car or
by bus (avoiding any physical activity) in a fasted state (between
7 and 10 h). They were instructed to avoid moderate or vigorous
physical activity 24 and 48 hours before the testing day. A nutri­
tionist prescribed an individualized pretrial diet (ie, 24 h before
each testing day: 2653 (162) kcal; 50% carbohydrate, 30% fat, and
20% protein), and the participants adhered to it of their own accord.
When the tests were perfonned in the afternoon, we instructed the
participants to consume the same menu (same energy intake and
percentage of macronulrient in each meal) in the same order that was
consumed in the morning test (ie, the breakfast in the morning test
[24 h ago] was the lunch in the afternoon test [24 h ago]). Energy
demand was determined using the Harris-Benedict equation based
on body mass, height, and age. An activity factor of 1.8 was used.14
On day 1, the body weight and height were measured using a
seca scale and stadiometer (model 799, electronic column scale;
Seca, Hamburg, Germany), and the body mass index was calcu­
lated as weight (kg)/height (nr). Participants wore light clothing
and no shoes during the measurements. Fat mass was assessed
by dual energy X-ray absorptiometry (Hologic Discovery Wii;
Hologic, Bedford, MA). The participants also completed the HOME
questionnaire,15 which is a validated questionnaire that determines
the participants’ chronotype (momingness-eveningness). The ques­
tionnaire consists of 19 questions related to sleepAvake behavior
and yields scores ranging from 16 to 86. Based on the HOME score,
the participants were categorized into 1 of 5 chronotype categories:
16 to 30 (definite evening type), 31 to 41 (moderate evening type),
42 to 58 (neither type), 59 to 69 (moderate morning type), and 70 to
86 (definite morning type).
The resting metabolic rate was measured by indirect calo­
rimetry during 15 minutes in peaceful and relaxing room
(temperature: 22.6 [0.7]°C; humidity: 44.5% [6.1%]). After
that, a maximal walking speed protocol on a treadmill (H/P/
cosmos pulsar; H/P/Cosmos Sports & Medical GmbH, NuBdorf,
Germany) was performed on the first day before the graded
exercise protocol to determine MFO, Fatmax, V 0 2max, and
VT2 adapted from a validated protocol.16’17’18 In brief, the
protocol started with a 3 minutes warm-up at 3.5 km/h, and
1 km/h speed increments were programmed every 3 minutes until
the maximal walking speed was reached. Subsequently, the
treadmill speed was constant, and the gradient was increased
by 2% ever)' 3 minutes until the respiratory exchange ratio was
>1.0. Then, after a 5-minute break, a maximal incremental exer­
cise test, using the modified Balke protocol (3-min walking at
5.3 km/h and 1%, followed by increments of 1% every minute)
until voluntary exhaustion, was performed. The final 30 seconds
of the V 0 2 measurement were considered to be maximal (V 02_
max) when the following conditions were met: (1) a plateau (an
increase of <2 mL/kg/min) in V 0 2 with a further increasing
workload, (2) a heart rate at least higher than the age-predicted
maximum minus 10 bpm, and (3) a respiratory exchange ratio
>1.1. If any of these criteria were not met, a V 0 2peak value was
taken, defined as the highest V 0 2 measured over a 30-second
period. VT2 was estimated from gas exchange data by 2 inde­
pendent researchers following a validated standard methodology,
as previously described.19
IJSPP Vol. 14, No. 8, 2019
Diurnal Variation of MFO Athletes 1141
An oronasal mask (model 7400; Hans Rudolph Inc, Kansas
City, MO) equipped with a preVent metabolic flow sensor (Medical
Graphics Corp, St Paul. MN) was fitted, and breath-by-breath
respiratory measurements were recorded throughout the test with
the use of an automated gas analysis system (CPX Ultima Cardi02;
Medical Graphics Corp, St Paul, MN). Gas analyzers were calibrated
immediately before each graded exercise protocol according to the
manufacturer’s recommendations. Heart rate was recorded using a
heart-rate monitor (Polar RS800; Polar Electro Inc, Woodbury, NY).
V 0 2, V C 02, and ventilation data were averaged over the
most stable 5-consecutive-minute periods (after discarding the
first 5 min) for analysis of the resting metabolic rate applying
the Weir abbreviated equation (assuming negligible protein oxi­
dation) and expressed as kcal/day: resting metabolic rate =
(3.9 [V 02]+1.1 [VC02])x 1.44. V 0 2, V C 02, and ventilation
data were averaged over and the last 60 seconds of each graded
exercise protocol stage. Stoichiometric equations described
by Frayn20 were used to calculate fat oxidation rates with the
assumption that urinary nitrogen excretion was negligible in all
cases. Fat oxidation rates were plotted against the relative exercise
intensity (% V02max), and a third-degree polynomial regression
was used to determine MFO and Fatmax for each individual
participant.
Statistical Analysis
The determination of the sample size and power of the study are
made based on the data of a pilot study. We considered MFO
differences between the morning and afternoon tests to assess
the sample size requirements for the 1-way analysis of the variance.
As a result, we expected to detect an effect size of 0.05 g/min
considering a type I error of 0.05, with a statistical power of 0.85
with a minimum of 10 participants. Assuming a maximum loss of
20%, we decided to recruit a total of 12 participants.
Results are reported as the mean (SD), otherwise stated. We
used the Shapiro-Wiik test, a visual check of histograms, and
Q-Q plots to verify the normal distribution of all variables. A
repeated-measures analysis of the variance was applied to deter­
mine differences between MFO-morning versus MFO-aftemoon,
Fatmax-morning versus Fatmax-alternoon, V 0 2max-moming ver­
sus V 0 2max-aftemoon, and VT2-morning versus VT2-aftemoon.
A 1-way repeated-measures analysis of covariance was conducted
to study morning versus afternoon differences, including fat mass
percentage and chronotype as covariates.
To analyze the association of V 0 2max and VT2 with MFO and
Fatmax, we conducted a simple linear regression analysis as follows:
(1) V 0 2max-moming with MFO-morning, (2) V 0 2max-moming
with Fatmax-moming, (3) VT2-morning with MFO-moming,
(4) VT2-moming with Fat^-m om ing, (5) V 02max-aftemoon with
MFO-aftemoon, (6) V 02max-altemoon with Fatmax-afternoon,
(7) VT2-aftemoon with MFO-aftemoon, and (8) VT2-aftemoon
with Fatmax-altemoon. We also included fat mass percentage and
chronotype as covariates.
The analyses were conducted using the Statistical Package for
Social Sciences (v 22.0, IBM SPSS Statistics; IBM Corp, Armonk,
NY). For all statistical procedures, the significance level was set
at P < .05.
Results
Descriptive parameters of the study participants are listed in
Table 1. Most of the participants did not lit in definite morning
1142 Amaro-Gahete et al

or definite evening chronotypes (-92% ). The test order was Fatmax -morning and Fatmax-afternoon (59.0% [8.1%] vs 62.6%
morning-afternoon in 7 participants and afternoon-morning in 5 [7.0%] V 0 2max, respectively; P = .0 0 1 , Figure 1C and ID) that
participants. Fasting time was similar in the morning and in the remained once fat mass percentage and chronotype were included
afternoon (8.4 [1.2] vs 8.2 [1.0] h, respectively, P = .554). in the model (P = .018 and P c . 001, respectively).
We observed significant differences between MFO-moming There were no significant differences between V 0 2max-moming
and MFO-aftemoon (0.55 [0.12] vs 0.63 [0.15] g/min. respectively; and VCHmax-aftemoon (63.7 [9.5] vs 63.9 [9.7] mL/kg/min, respec­
P < . 001, Figure 1A and IB), which persisted after controlling for tively: P = .158, Figure 2A and 2B). which persisted after controlling
for fat mass percentage and chronotype (P = .288 and P = .5 6 l,
fat mass percentage and chronotype (P = .023 and P < .001, respec­
respectively). Similarly, there were no significant differences between
tively). Similarly, there were significant differences between
VT2-moming and VT2-aftemoon (78.2% [4.9%] vs 78.8% [6.9%]
V 0 2max, respectively; P = .650, Figure 2C and 2D) that remained
T a b le 1 D e s c rip tiv e P a ra m e te rs of S tu d y P a rtic ip a n ts once fat mass percentage and chronotype were included in the model
(N = 12) (P = .309 and P = .784, respectively).
V 0 2max was positively associated with MFO in both morning
Age, y 24.7(4.1) ( P c . 001, Figure 3A) and afternoon assessments (P = .001,
Weight, kg 69.5 (9.2) Figure 3B). A positive association was observed between V 0 2max
Height, m 1.75 (0.04) and Fatmax in both morning (P = ,024, Figure 3C) and afternoon
assessments ( P - . 026, Figure 3D). VT2 was positively associated
Body mass index, kg/m2 22.7 (2.3)
with MFO in both morning (P = .005, Figure 3E) and afternoon
Fat mass, % 16.7 (3.7)
assessments ( P = .034, Figure 3F). A positive association was
Resting metabolic rate, kcal/d 2096.8(212.8) observed between VT2 and Fatm;ix in the afternoon assessment
Resting fat oxidation, g/min 0.068 (0.014) (P = .024, Figure 3H), whereas a tendency toward significance
Maximal oxygen uptake, mL/kg/min 63.8 (9.6) was noted in the morning assessment (P = . 105, Figure 3G). We
HOME questionnaire score 47.1 (13.7) repeated all the regression analysis after controlling for either fat
Definitive evening type 1 (8.3%) mass or chronotype, and the results did not change (data not shown).
Moderate evening type 2 (16.7%)
Neither type 6 (50.0%) D is c u s s io n
Moderate morning type 3 (25.0%)
The main finding of this study shows that MFO and Fatmax are higher
Definite morning type 0 (0 %) in the afternoon than in the morning in endurance-trained male
Note: Values are presented as mean (SD) or n (%). athletes. Moreover, we observed no differences in V 0 2max and VT2

B 0 .9 P < .0 0 1

M orning Afternoon

M orning A fternoon

Figure 1 — MFO (A and B) and Fatmax (C and D) in the morning and in the afternoon. Results are shown as the individual observations for each
participant (gray lines) and as the mean for all participants (black line). P value obtained by repeated-measures analysis of variance. Fatmax indicates the
intensity at which MFO occurs: MFO, maximal fat oxidation; V 0 2max, maximum oxygen uptake.

IJSPP Vol. 14, No. 8, 2019

 Diurnal Variation of MFO Athletes 1143

A 90 B

80-
E E
o> ra
_J
E 70-
x x
to
E E
O
CN
.......... . * "7 O
> 60- >

50
M o rn in g A fte rn o o n M orning A fterno on

M orning A fterno on

Figure 2 — V 02max (A and B) and VT2 (C and D) in the morning and in the afternoon. Results are shown as the individual observations for each
participant (gray lines) and as the mean for all participants (black line). P value obtained by repeated-measures analysis of variance. V 02max indicates
maximum oxygen uptake; VT2, V 02max percentage in ventilatory threshold 2.

in the morning versus the afternoon. We also observed a significant
positive association of V 0 2max and VT2 with both MFO and
Fatmax. These findings support the idea that the MFO and Fatmax
diurnal variation should be considered for repeat laboratory testing in
research, clinical, and athlete monitoring settings, as maintaining the
same fasting time does not seem to nullify these effects. In addition,
these findings may partially explain the observed increased endur­
ance sport performance in the afternoon, specifically in events
limited by endogenous carbohydrate availability.7
Our results extend those reported by others in untrained
individuals.10,11Mohebbi and Azizi11reported that MFO and Fatmax
were higher in the afternoon than in the morning in untrained
normal weight and obese individuals aged 19-25 years. Similarly,
Darvakh et a l10 observed significantly greater MFO and Fatmax in
the afternoon than in the morning in nonathlete male students. The
MFO differences observed in the present study were larger than
those observed by Mohebbi and Azizi11 (14.5% vs 8.9%) and
Darvakh et al10 (14.5% vs 6.7%), whereas Fatmax differences
were smaller than those obtained by Mohebbi and Azizi11 and
Darvakh et al10 (6.1% vs 12.2% and 10.7%, respectively).10’11
It is well known that the catecholamine peak induced by
exercise is higher in the afternoon than in the morning.9 Consider­
ing that the catecholamine release activates the lipolysis in skeletal
muscle and in adipose tissue,21 it seems reasonable that this will
lead to increased plasma fatty acid content, which could explain the
elevated fat oxidation rates observed in the afternoon. However, a
higher catecholamine release in the afternoon may also increase the
glycogenolysis during exercise,22 producing a potential decrement
of fat oxidation during exercise. Future studies are needed to
investigate whether a higher plasma catecholamine concentration
can induce higher MFO and Fatmax levels in the afternoon, as
we have no data on exercise-induced catecholamine release.
In addition, a number of studies suggest that body temperature,
time to the exhaustion, and V 0 2max are higher in the afternoon
than in the morning in active and untrained individuals.11-23
Moreover, our data indicate that both V 0 2max and VT2 are similar
in the morning and in the afternoon, which does not agree with
others.11>23 Other studies are warranted to determine diurnal differ­
ences in V 0 2max and VT2 in trained athletes.
The association of V 0 2max with MFO and Fatmax remains
unclear, as controversial results have been reported. Several studies
showed positive associations of V 0 2max with MFO in moderately
trained men (V 0 2max ranged from 50 to 55 mL/kg/min),24 in trained
male endurance athletes (VO2m ax> 70 mL/kg/min),25 in healthy
young adults (V 0 2max = 43.9 [7.2] mL/kg/min),13 and in a very
heterogeneous group of 300 men and women (V 0 2max = 46.3
[0.7] mL/kg/min),12 which concurs with the results obtained in our
study. However, these findings differ from those obtained by
others,26-27 who did not find significant associations between V 0 2_
max and MFO in healthy trained individuals (VO2max = 58.0
[1.6] mL/kg/min)26 and in male ironman athletes (V 0 2max ranged
from 43.9 to 72.5 mL/kg/min).27 It has been suggested that this
association is only present when heterogeneous groups are com­
pared. 12’27-2S These results concurred with our findings that showed a
strong positive association between V 0 2max and MFO in a hetero­
geneous cohort of endurance-trained male athletes (V 0 2max ranged
from 52.9 to 83.7 mL/kg/min). The fact that individuals with higher
V 0 2max normally had greater capacity of the muscle to oxidize fat12
could partially explain the observed association. In addition, it has
previously reported that trained individuals use more fat at the same
relative exercise intensity than untrained individuals in both longi­
tudinal29 and cross-sectional28 training studies.
Previous studies suggested that greater endurance performance
is most frequently observed in the afternoon.7 Atkinson et al30

1 J S P P V o l. 1 4 , N o . 8 , 2 0 1 9
1144 Amaro-Gahete et al

A Morning B Afternoon

VT2 (% V 0 2max) VT2 (% V 0 2max)

Figure 3 — Association between (A, B) V 0 2max and MFO. (C, D) V 0 2max and Fatmax, (E, F) VT2 and MFO, and (G, H) VT2 and Fatmax in the
morning and in the afternoon. (3 indicates unstandardized regression coefficient: Fatmax, the intensity at which MFO occurs; MFO. maximal fat oxidation;
R2, coefficient of determination; V 0 2max. maximum oxygen uptake. VT2. V 0 2max percentage in ventilatory threshold 2. P value obtained from a simple
linear-regression analysis.

showed that aerobic cycling performance (measured by peak power)
is greater in the afternoon than in the morning in trained cyclists.
Moreover, Souissi et al23 also found greater peak power and V 0 2max
in the afternoon than in the morning, yet no differences were observed
from morning to afternoon in V 0 2max when corrected for total work
done. Taken together, it is plausible that the diurnal variation of MFO
and Fatmax might be the key factor in endurance performance diurnal
variation, rather than of V 0 2max and VT2, specifically in events
limited by endogenous carbohydrate availability.
The results of this study should be considered with caution.
The lack of body temperature data and blood parameters assess­
ments during the graded protocol lest did not allow us to confirm
whether metabolic and hormonal variables play a role in MFO and
Fatmax diurnal variation. It should also be acknowledged that the

IJSPP Vol. 14, No. 8, 2019


present study was performed in endurance-trained male athletes;
thus, these results cannot be extended to women or a sedentary
population. Despite we established a fasted state (between 7 and
10 h) as a pretesting previous condition, a stricter control of fasting
conditions should be considered in future studies (ie, 8 h), although
our results remained after controlling by fasting time (data not
shown). In addition, we do not know whether the differences found
in MFO and Fatmax are determined by the individual chronotype, as
the small sample size made it difficult to study. Finally, Croci et al24
reported a coefficient of variation ranging from 16% to 21% for
MFO estimation determined from 2 progressive exercise protocols
completed 3 to 7 days apart.24 This variability may bias the results
obtained by the current study.
Practical Applications
• MFO and Fatmax are greater in the afternoon than in the
morning in endurance-trained male athletes.
• No diurnal variation was observed in V 0 2max and VT2.
• There is a strong positive association of V 0 2max and VT2
with MFO and Fatmax.
Conclusion
In summary, our results indicate that MFO and Fatmax are greater in
the afternoon than in the morning in endurance-trained male
athletes, whereas there is no diurnal variation in V 0 2max and
VT2. Moreover, we observed a positive strong association of
V 0 2max and VT2 with MFO and Fatmax. These data are relevant
when scheduling training times, and specifically for coaches, who
usually engage in athletic testing and monitor training session that
can occur during different hours of the day, whenever the training
intensity will be the Fatmax. Further studies are needed to investi­
gate whether these results remain when a running or cycling
protocol are used to estimate MFO and Fatmax.
Acknowledgments
The study is supported by the Spanish Ministry of Education (FPU
13/04365, FPU 14/04172, and FPU 15/03960). The study was partially
supported by the University of Granada, Plan Propio de Investigacion
2016, Excellence actions: Units of Excellence; Unit of Excellence on
Exercise and Health (UCEES), and by the Junta de Andalucla, Consejerfa
de Conocimiento, Investigacion y Universidades and European Regional
Development Fund (FEDER), ref. SOMM17/6107/UGR. The authors
would like to thank all the athletes who participated in this study for their
time and effort. They are grateful to Ms Carmen Sainz Quinn for assistance
with the English language. The authors declare that the research was
conducted in the absence of any commercial or financial relationships that
could be construed as a potential conflict of interest. This study is part of a
PhD thesis conducted in biomedicine doctoral studies of the University of
Granada, Spain.
References
1. Jeukendrup AE. Regulation of fat metabolism in skeletal muscle. Ann
N Y Acad Sci. 2002;967:217-235. PubMed ID: 12079850 doi: 10.
1111/j. 1749-6632.2002.tb04278.x
2. Jensen J, Rustad PI, Koines AJ, Lai YC. The role of skeletal muscle
glycogen breakdown for regulation of insulin sensitivity by exercise.
IJSPP Vol. 14, No. 8, 2019
Diurnal Variation of MFO Athletes 1145
Front Physiol. 2011:2:112. PubMed ID: 22232606 doi:10.3389/
fphys.2011.00112
3. Gonzalez JT, Fuchs CJ, Betts JA, van Loon LJ. Liver glycogen
metabolism during and after prolonged endurance-type exercise. Am J
Physiol Endocrinol Metab. 2016;31 LE543-E553.
4. Fernandez-Verdejo R, Bajpeyi S, Ravussin E, Galgani JE. Metabolic
flexibility to lipid availability during exercise is enhanced in indivi­
duals with high insulin sensitivity. Am J Physiol Endocrinol Metab.
2018;315(4):E715-E722. PubMed ID: 29870678 doi:10.1152/
ajpendo.00126.2018
5. Maunder E, Plews DJ. Kilding AE. Contextualising maximal fat
oxidation during exercise: determinants and normative values. Front
Physiol. 2018;9:599. doi: 10.3389/fphys.2018.00599
6. Barnes KR, Kilding AE. Strategies to improve running economy.
Sports Med. 2015;45:37-56. doi:10.1007/s40279-014-0246-y
7. Drust B, Waterhouse J, Atkinson G, Edwards B, Reilly T. Circadian
rhythms in sports performance—an update. Chronobiol Int. 2005;
22:21 —44. PubMed ID: 15865319 doi:10.1081/CBI-200041039
8. Teo W, Newton MJ, McGuigan MR. Circadian rhythms in exercise
performance: implications for hormonal and muscular adaptation.
J Sports Sci Med. 2011;10:600-606.
9. Kim HK, Konishi M, Takahashi M, et al. Effects of acute endurance
exercise performed in the morning and evening on inflammatory
cytokine and metabolic hormone responses. PLoS One. 2015; 10:
eO 137567.
10. Darvakh H, Nikbakht M, Shakerian S, Mousavian AS. Effect of
circadian rhythm on peak of maximal fat oxidation on non-athletic
men. Zahedan J Res Med Sci. 2014; 16(6):8—11.
11. Mohebbi H, Azizi M. Maximal fat oxidation at the different
exercise intensity in obese and normal weight men in the morning
and evening. J Hum Sport Exerc. 2011;6:49-58. doi:10.4100/jhse.
2011.61.06
12. Venables MC, Achten J, Jeukendrup AE. Determinants of fat oxida­
tion during exercise in healthy men and women: a cross-sectional
study. J Appl Physiol. 2005;98:160-167. PubMed ID: 15333616
doi: 10.1152/japplphysiol.00662.2003
13. Ponce Gonzalez JG, Guadalupe-Grau A, Rodnguez-Gonzalez FG,
et al. Androgen receptor gene polymorphisms and maximal fat
oxidation in healthy men. A longitudinal study. Nutr Hosp. 2017;
34:1089-1098.
14. Harris J, Benedict F. A biometric study of human basal metabolism.
Proc Natl Acad Sci USA. 1918;4:370-373. PubMed ID: 16576330
doi: 10.1073/pnas.4.12.370
15. Home JA, Ostberg O. A self-assessment questionnaire to determine
momingness-eveningness in human circadian rhythms. Int J Chron­
obiol. 1976;4:97-110. PubMed ID: 1027738
16. Dandanell S, Prasst CB, Spndergard SD, et al. Determination of the
exercise intensity that elicits maximal fat oxidation in individuals with
obesity. Appl Physiol Nutr Metab. 2017;42:405^112. PubMed ID:
28177732 doi: 10.1139/apnm-2016-0518
17. Amaro-Gahete F, De-la-0 A, Jurado-Fasoii L, et al. Exercise
training as S-Klotho protein stimulator in sedentary healthy adults:
Rationale, design, and methodology. Contemp Clin Trials Commun.
2018;11:10-19. PubMed ID: 30023455 doi:10.1016/j.conctc.2018.
05.013
18. Amaro-Gahete F, Sanchez-Delgado G, Ruiz JR. Commentary: con­
textualising maximal fat oxidation during exercise: determinants
and normative values. Front Physiol. 2018;9:1460. PubMed ID:
30405428 doi: 10.3389/fphys.2018.01460
19. Lucia A, Hoyos J, Chicharro JL. The slow component of V 0 2 in
professional cyclists. Br J Sports Med. 2000;34:367-374. doi: 10.
1136/bjsm.34.5.367
1146 Amaro-Gahete et al
20. Frayn KN. Calculation of substrate oxidation rates in vivo from
gaseous exchange. J Appl Physiol. 1983;55:628-634. PubMed ID:
6618956 doi: 10.1152/jappl. 1983.55.2.628
2 1 . Jocken JWE, Blaak EE. Catecholamine-induced Iipolysis in
adipose tissue and skeletal muscle in obesity. Physiol Behav. 2008:
94:219-230. PubMed ID: 18262211 doi: 10.1016/j.physbeh.2008.
01.002
22. Watt MJ, Howlett KF, Febbraio MA, Spriet LL, Hargreaves M.
Adrenaline increases skeletal muscle glycogenolysis, pyruvate dehy­
drogenase activation and carbohydrate oxidation during moderate
exercise in humans. ./ Physiol. 2001;534:269-278. PubMed ID:
11433007 doi: 10.1111/j.l469-7793.200l.t01-l-00269.x
23. Souissi N, Bessot N. Chamari K, Gauthier A, Sesboiie B, Davenne D.
Effect of time of day on aerobic contribution to the 30-s Wingate
test performance. Chronobiol Ini. 2007;24:739-748. PubMed ID:
17701684 doi: 10.1080/07420520701535811
24. Croci I, Borrani F, Byrne NM. et al. Reproducibility of Fatmax and
fat oxidation rates during exercise in recreationally trained males.
PLoS One. 20l4;9:e97930. PubMed ID: 24886715 doi:10.1371/
joumal.pone.0097930
25. van Loon LJ, Greenhaff PL, Constantin-Teodosiu D, Saris WH,
Wagenmakers AJ. The effects of increasing exercise intensity on
IJSPP Vol. 14, No. 8, 2019
muscle fuel utilisation in humans. J Physiol. 2001;536:295-304.
PubMed ID: 11579177 doi:10.1111/j. 1469-7793.2001.00295.x
26. Astorino TA, Edmunds RM, Clark A, et al. Change in maximal fat
oxidation in response to different regimes of periodized high-intensity
interval training (HIIT). Eur J Appl Physiol. 2018;117:745-755.
PubMed ID: 28251399 doi: 10.1007/s00421-017-3535-y
27. Frandsen J, Vest SD, Larsen S, Dela F, Helge JW. Maximal fat
oxidation is related to performance in an ironman triathlon. Int J
Sports Med. 2017;38:975-982. PubMed ID: 29050040 doi: 10.1055/
s-0043-117178
28. Nordby P, Saltin B, Helge JW. Whole-body fat oxidation determined
by graded exercise and indirect calorimetry: a role for muscle
oxidative capacity? Scand J Med Sci Sports. 2006;16:209-214.
PubMed ID: 16643200 doi:10.1111/j. 1600-0838.2005.00480.X
29. Tan S, Wang J, Cao L. Guo Z, Wang Y. Positive effect of exercise
training at maximal fat oxidation intensity on body composition and
lipid metabolism in overweight middle-aged women. Clin Physiol
F und Imaging. 2016;36:225-230. PubMed ID: 27072372 doi: 10.
I l l 1/cpf. 12217
30. Atkinson G, Todd C, Reilly T, Waterhouse J. Diurnal variation in
cycling performance: influence of warm-up. J Sports Sci. 2005;23:
321-329. PubMed ID: 15966350 doi:10.1080/026404!0410001729919
Copyright of International Journal of Sports Physiology & Performance is the property of
Human Kinetics Publishers, Inc. and its content may not be copied or emailed to multiple
sites or posted to a listserv without the copyright holder's express written permission.
However, users may print, download, or email articles for individual use.