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Can Neuromuscular Fatigue Explain Running Strategies and Performance in Ultra-Marathons? The Flush Model


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While the industrialized world adopts a largely sedentary lifestyle, ultra-marathon running races have become increasingly popular in the last few years in many countries. The ability to run long distances is also considered to have played a role in human evolution. This makes the issue of ultra-long distance physiology important. In the ability to run multiples of 10 km (up to 1000 km in one stage), fatigue resistance is critical. Fatigue is generally defined as strength loss (i.e. a decrease in maximal voluntary contraction [MVC]), which is known to be dependent on the type of exercise. Critical task variables include the intensity and duration of the activity, both of which are very specific to ultra-endurance sports. They also include the muscle groups involved and the type of muscle contraction, two variables that depend on the sport under consideration. The first part of this article focuses on the central and peripheral causes of the alterations to neuromuscular function that occur in ultra-marathon running. Neuromuscular function evaluation requires measurements of MVCs and maximal electrical/magnetic stimulations; these provide an insight into the factors in the CNS and the muscles implicated in fatigue. However, such measurements do not necessarily predict how muscle function may influence ultra-endurance running and whether this has an effect on speed regulation during a real competition (i.e. when pacing strategies are involved). In other words, the nature of the relationship between fatigue as measured using maximal contractions/stimulation and submaximal performance limitation/regulation is questionable. To investigate this issue, we are suggesting a holistic model in the second part of this article. This model can be applied to all endurance activities, but is specifically adapted to ultra-endurance running: the flush model. This model has the following four components: (i) the ball-cock (or buoy), which can be compared with the rate of perceived exertion, and can increase or decrease based on (ii) the filling rate and (iii) the water evacuated through the waste pipe, and (iv) a security reserve that allows the subject to prevent physiological damage. We are suggesting that central regulation is not only based on afferent signals arising from the muscles and peripheral organs, but is also dependent on peripheral fatigue and spinal/supraspinal inhibition (or disfacilitation) since these alterations imply a higher central drive for a given power output. This holistic model also explains how environmental conditions, sleep deprivation/mental fatigue, pain-killers or psychostimulants, cognitive or nutritional strategies may affect ultra-running performance.
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Can Neuromuscular Fatigue Explain
Running Strategies and Performance
in Ultra-Marathons?
The Flush Model
Guillaume Y. Millet
1 Universite
´de Lyon, F-42023, Saint-Etienne, France
2 Inserm U1042, Grenoble, F-38000, France
Abstract................................................................................. 489
1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490
2. Neuromuscular Alterations in Ultra-Marathon Running . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491
2.1 Central Contribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 492
2.2 Alterations at the Muscle Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493
3. The Flush Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494
3.1 Power Output Change in Ultra-Marathon Runners. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
3.2 Description of the Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
3.3 Feed-Forward and Feedback Mechanisms Influence the Filling Rate. . . . . . . . . . . . . . . . . . . . . . 497
3.4 Apart from the Filling Rate, Which Factors Influence the Quantity of Water?. . . . . . . . . . . . . . . . 499
3.5 The Waste Pipe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500
3.6 The Security Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501
4. Conclusion ........................................................................... 502
Abstract While the industrialized world adopts a largely sedentary lifestyle, ultra-
marathon running races have become increasingly popular in the last few
years in many countries. The ability to run long distances is also considered to
have played a role in human evolution. This makes the issue of ultra-long
distance physiology important. In the ability to run multiples of 10 km (up to
1000 km in one stage), fatigue resistance is critical. Fatigue is generally de-
fined as strength loss (i.e. a decrease in maximal voluntary contraction
[MVC]), which is known to be dependent on the type of exercise. Critical task
variables include the intensity and duration of the activity, both of which are
very specific to ultra-endurance sports. They also include the muscle groups
involved and the type of muscle contraction, two variables that depend on the
sport under consideration. The first part of this article focuses on the central
and peripheral causes of the alterations to neuromuscular function that occur
in ultra-marathon running. Neuromuscular function evaluation requires
measurements of MVCs and maximal electrical/magnetic stimulations; these
provide an insight into the factors in the CNS and the muscles implicated in
fatigue. However, such measurements do not necessarily predict how muscle
REVIEW ARTICLE Sports Med 2011; 41 (6): 489-506
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function may influence ultra-endurance running and whether this has an ef-
fect on speed regulation during a real competition (i.e. when pacing strategies
are involved). In other words, the nature of the relationship between fatigue
as measured using maximal contractions/stimulation and submaximal per-
formance limitation/regulation is questionable. To investigate this issue, we are
suggesting a holistic model in the second part of this article. This model can be
applied to all endurance activities, but is specifically adapted to ultra-endurance
running: the flush model. This model has the following four components: (i) the
ball-cock (or buoy), which can be compared with the rate of perceived exertion,
and can increase or decrease based on (ii) the filling rate and (iii) the water
evacuated through the waste pipe, and (iv) a security reserve that allows the
subject to prevent physiological damage. We are suggesting that central reg-
ulation is not only based on afferent signals arising from the muscles and peri-
pheral organs, but is also dependent on peripheral fatigue and spinal/supraspinal
inhibition (or disfacilitation) since these alterations imply a higher central drive
for a given power output. This holistic model also explains how environmental
conditions, sleep deprivation/mental fatigue, pain-killers or psychostimulants,
cognitive or nutritional strategies may affect ultra-running performance.
1. Introduction
More than 30 000 articles have been publish-
ed about fatigue. Limiting keywords to ‘muscle’
and ‘fatigue’ still gave us more than 12 000 arti-
cles. It is known that the magnitude and aetiology
of fatigue depend on the exercise under con-
Critical task variables include the
muscle activation pattern, the type of muscle
group involved and, the type of muscle con-
traction. However, the intensity and dura-
tion of activity are probably among the most
important factors. This article focuses on ultra-
endurance running exercises, the so-called ultra-
Throughout the world (e.g. in the US, Europe,
Japan, Korea, South Africa), ultra-marathons have
become increasingly popular in the last few years.
For example, Hoffman et al.
recently analysed
the participation in 161 km ultra-marathons in
North America and showed that the number of
finishes increased exponentially over the period
19772008 through a combination of increases
in the number of participants, average annual
number of races completed by each individual,
and number of races organized every year. It is
considered that more than 30 000 runners took
part in at least one ultra-marathon in France in
2009. There is no consensus about the definition
of contemporary ultra-marathons; some authors
consider it to be any distance greater than a
marathon, while for others, it is any event that
exceeds 4 hours
or 6 hours
in duration. Ultra-
marathons can last for up to 40 hours or even
several days (e.g. 6-day races) and are basically of
two types: (i) ultra-marathons performed on a
mostly flat road (24 hours, 100 km); and (ii) those
run on varying terrains (e.g. 100 miles in the US).
Contrary to what is usually claimed, ultra-marathon
running is not new; the Six-Day Professional
Pedestrian Races in London and New York have
existed since the 1880s.
Importantly, the ability
to run, rather than only walk, over long distances
(i.e. without fatigue) may have played a role in
human evolution.
For example, it has been sug-
gested that endurance running may have helped
hominids to exploit protein-rich resources. Thus,
while endurance running is now primarily a form
of recreation, its roots may be as ancient as the
origin of the human genus.
This type of extreme
event can also be seen as a testbed for ideas on
how some people manage to perform physical
feats at which others can only marvel.
Several models of fatigue have been proposed in
the literature. For example, Abbiss and Laursen
reviewed the following eight different models that
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may be applied to prolonged cycling: cardiovascular/
anaerobic, energy supply/energy depletion, neuro-
muscular fatigue, muscle trauma, biomechanical,
thermoregulatory, psychological/motivational and
the central governor model. All of these are
interrelated. For example, changes in biomecha-
nical patterns may be both the cause and con-
sequence of neuromuscular fatigue, which is also
influenced by modifications in cardiovascular/
anaerobic metabolism, muscle trauma and thermal
conditions, and all potentially associated with the
central governor. Similarly, depending on the
authors and the scientific field, central fatigue has
been presented as a decrease in percentage max-
imal voluntary activation (%VA),
gical modifications in the brain,
a modification
of motor control
or alterations in cognitive
In exercise physiology, most published articles
have defined fatigue as strength loss (i.e. a de-
crease in maximal voluntary contraction [MVC]).
Strength loss in the fatigued state is multifactorial
and is generally divided into central (i.e. above
the neuromuscular junction) and peripheral
(muscular), these two origins being interdependent
on the mediation of peripheral afferences. The
central/peripheral distinction was already pro-
posed by Bainbridge in 1931.
In this context,
central fatigue is an altered ability of the CNS to
recruit motor units at a higher discharge rate than
the frequency of tetanic fusion. In other words, a
decrease in maximal voluntary activation (i.e.
central fatigue) might be due to a de-recruitment
of motor units and/or a decrease of the discharge
frequency beyond the frequency of tetanic fusion,
both factors leading to force decline. Central fa-
tigue is variably implicated in total fatigue. It has
been shown that prolonged exercise is associated
with a large decrease in %VA, especially with
However, the role of central fatigue
and its supraspinal and spinal components in the
cessation of exercise (if the intensity is fixed) or
in performance (if the intensity is self-chosen) is
not clear. The problem is further complicated by
the fact that (i) environmental conditions such as
hypoxia and hyperthermia may exacerbate central
fatigue or perceived exertion,
two different
forms of central alterations; and (ii) that mental
fatigue another type of central alteration has
been demonstrated to play a role in performance
Although central fatigue is of great
importance, this does not imply that peripheral
fatigue is unimportant. How peripheral changes,
central fatigue, environmental conditions and a
runner’s strategies (whether cognitive, nutritional
or tactical) affect ultra-marathon performance
and regulation of speed during a race has never
been considered. Thus, the first aim of this article
is to review the central and peripheral factors that
might influence strength loss during very pro-
longed running exercise.
Describing these central and peripheral fac-
tors is essential but it does not predict how they
affect submaximal muscle function during ultra-
endurance running or how they influence speed
regulation during a real competition (i.e. including
pacing strategies). This poses the question of the
relationship between fatigue evidenced by measures
taken during maximal contractions/stimulation
and performance limitation/regulation. The second
aim of this article is then to propose a model that
integrates these different parameters, including
central and peripheral fatigue, which may help us
to understand pacing strategies and performance
during ultra-marathons. While the model is partic-
ularly well adapted to ultra-marathons, it can be
applied to any type of endurance performance.
2. Neuromuscular Alterations in
Ultra-Marathon Running
The alterations in neuromuscular function
after prolonged running, cycling and skiing were
reviewed by Millet and Lepers in 2004.
focused on the origin of muscle fatigue after
prolonged exercises lasting from 30 minutes to
several hours. The authors showed that the knee
extensors isometric strength loss increased in a
non-linear way with exercise duration when run-
ning for longer than 2 hours. Since then, several
articles have been published on this topic.
As shown in figure 1, the tendency toward no
further decrease in knee extensor strength with
increasing in running duration is confirmed.
Less is known about the decrease in peak
power after prolonged running. Nevertheless, it
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has been reported that the decrease in counter-
movement jump performance was around 4560%
of the knee extensors isometric MVC decrease after
prolonged running.
Similarly, Lepers
et al.
reported that isokinetic strength loss was
smaller when measured over a concentric contrac-
tion compared with the eccentric or isometric
2.1 Central Contribution
Methods such as the twitch interpolation
technique, the ratio of electromyogram (EMG)
signal during MVC normalized to the M-wave
response (Mmax) or the comparison of the forces
achieved with voluntary and electrically evoked
contractions, have systematically shown that cen-
tral fatigue largely contributes to muscle fatigue
during long distance running.
It is
known that the decrease in central activation oc-
curring during exercise can be caused by several
factors at the spinal (motoneurone properties,
afferent input) and/or supraspinal levels.
few studies have measured the changes in strength
of muscles not involved in the exercise to further
explore the origin of the lower central drive post-
It was hypothesized that a loss of
grip strength after running would be a good in-
dicator of supraspinal fatigue but no consistent
changes were observed in grip strength following
running exercise. Thus, this measurement did not
allow any conclusion of the existence or the absence
of supraspinal fatigue after prolonged running
because selective supraspinal fatigue may have
occurred. Ohta et al.
investigated biochemical
modifications during a 24-hour run and from
indirect measurements such as serum serotonin
and free tryptophan levels, they suggested that
this type of exercise induces some supraspinal
fatigue. It has been suggested for years that the
accumulation of serotonin in several brain regions
contributes to the development of fatigue during
prolonged exercise.
This was thought to be due
to an increase in the concentration ratio of free
tryptophan to branched-chain amino acids because
(i) branched-chain amino acids are oxidized; and
(ii) higher plasma free fatty acids during pro-
longed exercise cause a parallel increase in free
tryptophan since the free fatty acids displace
tryptophan from their usual binding sites on al-
This in turn increases the concentra-
tion of free tryptophan (the serotonin precursor)
in the brain. Meeusen et al.
went further sug-
gesting that other neurotransmitters such as
dopamine probably also play a significant role
in supraspinal fatigue. Nevertheless, to the best
of our knowledge, no study has clearly shown
any evidence of central fatigue (e.g. a depressed
%VA) with an increased serotonin/dopamine
ratio or other biochemical changes in the CNS.
Meeusen et al.
acknowledged that fatigue is
probably due to a complex interaction between
Knee extensors strength loss (%MVC)
35 *
0 500 1000
Duration (min)
1500 2000 2500
Fig. 1. Relationship between strength loss in the knee extensors expressed as a percentage of maximal voluntary contraction (%MVC) at rest
and duration of running exercise.
[18,19,23-30] *
Indicates the value is from unpublished observations.
492 Millet
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central and peripheral mechanisms. It is worth
noting that we recently observed a significant cor-
relation between %VA changes during a 24-hour
treadmill run for plantar flexor and knee extensor
which could be indicative of a com-
mon supraspinal mechanism regulating the neural
drive to the working muscle. Another possibility
is that hyperventilation lowers the arterial carbon
dioxide tension and blunts the increase in cere-
bral blood flow, which can lead to an inadequate
oxygen delivery to the brain and contribute to the
development of fatigue.
Nevertheless, this is
less likely to occur during ultra-marathons, be-
cause of the relatively low level of ventilation.
While some central activation deficit has been
observed for knee extensor muscles in cycling,
there is a lower level of central fatigue after ac-
tivities that result in less muscle damage than do
When marathon skiing
and 30-km
in similar competitive conditions and
duration were compared, the decrease in %VA
was more pronounced for running than for skiing.
Millet and Lepers
suggested that this result
indicated spinal modulation rather than cortical
alteration after the running exercise. Data from
reflex measurements, such as the Hoffmann
reflex (Hmax), provide interesting insights into
the origin of central fatigue. The Hmax/Mmax
ratio as been used as an indicator of motoneu-
ronal excitability, and more generally to evidence
modulations of neural drive at the spinal level.
This index was found to decline during a 24-hour
treadmill run and was correlated with decreases
in MVC and %VA,especiallyattheendoftheex-
ercise (personal observation). This finding con-
curs with those of Racinais et al.
who reported
depressed H-reflexes after a 90-minute run. This
could be due to reduced motoneurone excitability
or pre-synaptic inhibition. In both cases, in-
hibitory mechanisms could be limiting muscle
force production. Such inhibitory action may re-
sult from thin afferent fibre (group IIIIV) sig-
nalling, which may have been sensitized by the
production of pro-inflammatory mediators, pro-
duced during prolonged exercise.
So, while
supraspinal fatigue may play a role in reduced
neural drive after prolonged exercises, it can be
suggested that spinal adaptation, such as inhibi-
tion from type III and IV group afferents or dis-
facilitation from muscle spindles contributes to
this reduced central drive. Group IIIIV afferent
fibres may also contribute to the submaximal out-
put from the motor cortex (see Taylor et al.
Taken together, this suggests that high central
fatigue due to prolonged running is not due solely
to CNS biochemical changes but that afferent fi-
bres are probably involved. The twitch inter-
polation technique at the peripheral nerve does
not allow discrimination between central fatigue
originating from a supraspinal site and/or from
the spinal level. Researchers at the Prince of
Wales Medical Research Institute in Sydney (e.g.
Todd et al.
) have measured supraspinal deficit
by superimposing magnetic stimulations of the
motor cortex to voluntary contractions. Recently,
this method has been used to demonstrate the
existence of supraspinal %VA alteration in the
quadriceps after cycling exercise and prolonged
Future studies should use this new
method to further investigate central fatigue after
ultra-distance running exercises.
2.2 Alterations at the Muscle Level
Central fatigue alone cannot explain the entire
strength loss after prolonged running exercises.
Alterations of neuromuscular propagation, failure
of excitation-contraction coupling and mod-
ifications in the intrinsic capability of force pro-
duction may also be involved. To the best of our
knowledge, there has been no measure of change
in action potential conduction velocity using
high-density EMG after prolonged running ex-
ercise. Information about the propagation of the
action potentials can then only be deduced from
changes in the M-wave characteristics. This is
especially in the case of ultra-
marathons, since muscle oedema and sweat can
complicate interpretation of the Mmax. Limits also
exist for mechanical twitch responses after ultra-
in particular the fact that fully
potentiated twitches were not always used in the
past (e.g. Millet et al.
). Tetanic responses are
slight or not influenced by potentiation. A sig-
nificant but moderate (~10%) decrease in high-
frequency force response has been found for knee
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extensors after prolonged
or ultra-long
running exercise.
Indirect indices of muscle damage (creatine
kinase, myoglobin, C-reactive protein, myosin
heavy chain fragments, lactate dehydrogenase,
aspartate aminotransferase, alanine aminotrans-
ferase, cytokines) also suggest the existence of
some peripheral alterations
but from
the few results available, it appears that subjects
show wide variability in the degree of muscle
Potential explanations for this vari-
ability in inter-individual response include differ-
ences in genes, training (particularly the repeated
bout effect,
that likely occurs, particularly
during downhill running training
) flexibility,
oxidative stress,
muscle fibre type
and running
technique. No direct evidence exists to preferentially
support any one factor and it is probably a com-
bination of them all that explains the large dif-
ference in muscle damage among subjects.
Nevertheless, for the last two potential factors
(i.e. typology and running technique), it is inter-
esting to report that (i) there was no significant
relationship between knee extensor peripheral
fatigue and percentage type I muscle fibres in the
vastus lateralis;
and that (ii) ultra-long running
(from Paris to Beijing [i.e. 8500 km in 161 days,
~53 km daily]) modified the running patterns to-
wards a ‘smoother’ style in one case study.
latter point was evidenced by (i) a higher stride
frequency and duty factor; (ii) a reduced aerial
time with no change in contact time; (iii) a lower
maximal vertical force and loading rate at im-
pact; and (iv) a decrease in both potential and
kinetic energy changes at each step.
We also
measured a reduced running economy after the
trip. Thus, even if it is possible that the running
pattern changes could be linked to the decrease in
maximal strength also observed, we suggested
that running pattern modification was a strategy
to reduce the potential deleterious effects of his
ultra-long distance run rather than to decrease
the energy cost of running. Further studies should
also examine the potential influence of running
technique on muscle damage during ultra-endurance
running, particularly when running on variable
terrain (trails). Anecdotal information from qua-
lified coaches suggests that technical ability in
downhill sections might be a real determinant of
fatigue and performance.
Low-frequency fatigue (LFF; also called pro-
longed low-frequency force depression
) [i.e.
the preferential loss of force at low frequencies of
electrical stimulation] is a prominent character-
istic of exercises involving lengthening contrac-
tions of the active muscles such as eccentric- and
stretch shortening cycle-type exercises,
and has
been associated with failure of the excitation-
contraction coupling.
Contrary to what is gen-
erally observed after downhill running,
studies did not show LFF after prolonged run-
ning exercise despite several experiments that have
measured this factor during ultra-long running
exercises including the recent 24-hour treadmill
Only recently have we been able
to able to measure LFF after one of the most
extreme exercises realized by humans in race
conditions: a 166-km mountain ultra-marathon
with 9500 m of positive and negative elevation
It can then be suggested that minimal
exercise intensity is required to induce mechanical
or metabolic disturbances that can result in devel-
oping LFF. However, there is a limitation in that
it is theoretically possible that axonal hyperpolar-
ization preferentially depresses the high-frequency
response during tetanic muscle stimulation. Thus,
an absence of modification to the low- to high-
frequency ratio could have resulted from the
combined effects of LFF, which preferentially
depresses low-frequency response, and hyperpo-
larization, which preferentially depresses high-
frequency response.
3. The Flush Model
Measuring central activation changes or force/
EMG responses during MVCs or electrically
evoked stimulation after prolonged running gives
some insight into the potential factors implicated
in fatigue at the CNS and/or muscle level. How-
ever, it does not predict how this affects sub-
maximal muscle function during ultra-endurance
running or how this influences speed regulation
during a real competition when pacing strategies
are allowed. This poses the question whether there
is any relationship between fatigue evidenced by
494 Millet
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measures taken during maximal contractions/
stimulation and performance limitation/regulation.
Before answering this question, the change in
power output during ultra-marathons must first
be described.
3.1 Power Output Change in Ultra-Marathon
Only one study
has examined the changes in
efficiency/energy cost to determine whether
power output can be connected to speed. Since it
is unlikely that a large uncoupling exists between
power output and velocity, monitoring the speed
change over flat-course ultra-marathons (the ty-
pical event being the 24-hour race) gives an ex-
cellent idea of the mechanical power produced by
the runner. During a self-paced 24-hour treadmill
run where the subjects were asked to give their
best performance as they would in a normal race,
Martin et al.
showed a clear decrease in velo-
city during the first 16 hours before there was a
tendency for this to level off. Over a shorter dis-
tance (68 km) in a real competition, Utter et al.
showed that subjects reached a rating of per-
ceived exertion (RPE) similar to the subjects of
Martin et al.
(i.e. 15.4 0.4) but there was
an increase of RPE up to the end of the race.
While several studies have investigated 24-hour
we are not aware of any data report-
ing velocity changes during an official race. An-
ecdotal evidence and personal data suggest that
speed is reduced during ultra-marathons, even in
elite athletes. Also, unpublished results showed
that the speed of the top five runners in the 2007
French 24-hour championship had a tendency to
decrease but that this was less pronounced for the
winner. Over a shorter distance (100 km at the
1995 International Association of Ultra Runners
World Challenge), Lambert et al.
that the best runners (i) reduced their initial speed
less and later in the race; and (ii) they showed less
variation in their speeds than did their lesser-
performing counterparts. The general tendency
was still that these 107 runners reduced their
speed over time.
In marathon running, it has
been reported
that some elite athletes are able
to maintain their pace throughout the race but
their slower counterparts (especially young men
slow down over the distance.
It is more difficult to document power output
and speed change during ultra-trails (i.e. off-road
ultra-marathons) since the terrain is usually hilly,
even mountainous. Future studies should mea-
sure the changes in speed at given slopes using
global positioning system tools. Data from Utter
et al.
nevertheless suggest that heart rate (HR)
decreases over a 68-km ultra-marathon. Personal
data also show that HR generally decreases over
an ultra-endurance race in mountains (see figure 2a
and the first 22 hours in figure 2b) and that a
correlation exists between change in HR and
change in elevation speed (in m/hour) similar to
the relationship seen between change in HR and
speed variation after the first 10 km of a mara-
When using HR to predict speed varia-
tions, one should also consider the HR drift with
fatigue (i.e. HR increases at a given speed). This is
visible in the first 12 hours in figure 2 but the
change in HR due to cardiac drift is less than the
speed variation. However, any decrease in HR
during competitions can also only be due to a
decreased power output.
3.2 Description of the Model
Despite the lack of systematic studies on speed
change over ultra-marathons, the data presented
strongly suggest that power output/speed decreases
with time in ultra-marathon running, even in elite
athletes. Following the initial question about the
relationship between fatigue as evidenced by
measures taken during maximal contractions/
stimulations and performance limitation/regulation,
one should then ask whether the decreased speed
is due to strength loss. Based on the fact that knee
extensor and plantar flexor muscle strength de-
creased by ~3040%and since the force devel-
oped at each step is very low, it does not appear
that strength loss can directly explain this speed
reduction. Marcora et al.
suggested that the
locomotor muscles’ capacity for force production
is always well above the requirements of high-
intensity cycling, which is about 20%of MVC. In
other words, even if this point has been debated,
failure to produce the force/power required by
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the exercise despite maximal voluntary effort
does not seem to limit submaximal performance
as commonly assumed.
While it is not possible to predict to what extent
the fatigue mechanisms identified during max-
imal isometric contractions would affect perfor-
mance during an ultra-endurance running event,
it may nevertheless be hypothesized that there
is an indirect effect. As proposed by the tele-
oanticipatory system
or the central governor
the level of muscle activation (and so
the speed) is thought to be progressively reduced
to keep the RPE during running below a max-
imum tolerated level
(i.e. to maintain the body
below a homeostatistically acceptable exercise
In relation to the data presented
above regarding central and peripheral fatigue,
the last part of the present article will discuss
Altitude (m)Altitude (m)
HR (beats/min)HR (beats/min)
612 18 22 24
2 h
Race duration (h)
Race duration (h)
Fig. 2. Typical heart rate (HR) changes during two mountain ultra-marathon races: one crossing the island of la Reunion (155 km [a]) and one
around the Mont-Blanc (165 km [b]). The global tendency of heart change is indicated by the solid line. The profile of the course is also given
(grey shading).
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(i) how RPE is progressively increased with time
for a constant (or even lower) running speed; and
(ii) how in relation to changes in environmental
conditions, sleep deprivation/mental fatigue, drugs,
cognitive or nutritional strategies, this may reg-
ulate performance in ultra-marathon running. We
propose a conceptual model based on the flush
toilet (figure 3). The ‘flush model’ is based on the
central governor model proposed by Noakes
et al.
(i.e. agrees with the fact that exercise
performance is regulated by the CNS specifically
to prevent catastrophic physiological failure). How-
ever, the flush model emphasizes the importance
of peripheral fatigue that has been described in
detail in the first part of the present article. Also,
because it has been suggested that the central
governor integrate the input from various sys-
tems all related to exercise,
the flush model was
also built to take into account changes not asso-
ciated with exercise. The present model was mainly
designed to explain the role of fatigue on perfor-
mance in ultra-marathon running and consists of
four components: the ball [or buoy, (1) in figure 3]
represents RPE and can increase or decrease
based on the filling rate (2) and the water evac-
uated through the waste pipe (3). There is also a
security reserve (4), also called the emergency
which allows the subject to prevent
physiological harm.
We believe that the flush
model can help us to understand running strate-
gies during an ultra-marathon, but a few ques-
tions must first be addressed.
3.3 Feed-Forward and Feedback
Mechanisms Influence the Filling Rate
First, what influences the filling rate? At the
beginning of an ultra-marathon, running pace
is based on a control system which estimates
the optimal power output.
Depending on the
runner, his or her goal may be either simply fin-
ishing the race, finishing the race in a certain time
Security reserve
Filling rate
Waste pipe
Exercise cessation
I feel good (±)
Nociceptive information
(muscle, joint, tendon, blister,
digestive problems, etc.)
Peripheral fatigue + spinal and/or
supraspinal inhibition
Fig. 3. The flush model. Rating of perceived exertion (RPE) is assimilated to the volume of water in the tank (i.e. an increase in volume of
water signifies a higher RPE and decreasing the level of water in the tank indicates decreasing RPE). The water can get in (filling rate, 2) and
out (via waste pipe, 3) and the level of water can be detected by the ball (1). The level of water depends on the filling rate, mainly determined by
peripheral changes and central inhibition/disfacilitation (feedback and feed-forward mechanisms), but other factors such as mental fatigue,
nutritional strategies, sleep deprivation, environmental conditions and exceptional events during races can affect the level. The size of
the security reserve (4) is mainly determined by motivation. Psychostimulants and pain killers can modify the sensitivity of the RPE sensor
(i.e. the ball).
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or being well placed at the finish (for instance
being in the top ten runners). The estimation is
based on several factors such as distance, eleva-
tion, environmental conditions, training status
and runabilty (i.e. technical difficulty) of the course.
This teleoanticipation means that the runner has
a template that contains existing data on exercise
performance (the so-called ‘experience’ or per-
sonal history of running). In fact, Ulmer
siders that this template may be inborn but it is
generally accepted that optimal pacing strategy is
the result of a learning process.
The initial pace
gives an initial filling rate of the tank and the
higher the initial speed for a given running, the
faster the filling rate. This is mainly due to both
feed-forward and feedback mechanisms.
Feedback mechanisms have been widely docu-
mented in the literature. For instance, Amann
et al.
nicely showed that by attenuating the
ascending activity of nociceptive and metabo-
receptive Ad(group III) and C fibres (group IV),
somatosensory feedback from the locomotor mus-
cles influences central motor drive. Even if the
this experimental study has been criticized for
lack of proper placebo procedures,
the authors
concluded that locomotor muscle afferent feed-
back, which also facilitates performance through
optimizing muscle oxygen delivery,
exerted an
inhibitory influence on the determination of cen-
tral motor drive during high-intensity exercise.
While acidosis or inorganic phosphate accumu-
lation is unlikely to occur in ultra-marathon run-
ning, other biochemical mediators, such as the
accumulation of extracellular potassium
or cyto-
kines (especially interleukin-6 and its antagonist
IL-ra) due to structural muscle damage
section 2.2 about peripheral fatigue), could trig-
ger group III/IV afferent fibres and mediate the
sensation of fatigue.
Nociceptive information
is much more complicated; there is a potential
role for pain arising not only from the muscles
but also from other sites such as joints and tendons.
Even blisters (cutaneous afferences
) or diges-
tive problems
could all potentially play a role.
For shorter distances (i.e. higher intensity and
ventilation rate), dyspnoea may also be involved
in nociceptive information
but probably not in
ultra-marathon running. It should also be noted
that a modest temporary reduction in pressure
pain perception was observed after a 100-mile
(161-km) trail run, only in the faster runners.
Although the subject is debated,
feedback from
the locomotor muscles probably plays a major
role in central motor drive regulation in ultra-
marathons. As stated in section 2.1, the down-
regulation of group III/IV afferents at the spinal
and supra-spinal levels
is a probable explana-
tion for why maximal %VA is lower after running
than after cycling or skiing for similar intensity/
While afferent feedback certainly has a key
function, the regulation of central motor command
is complex and also depends on the environment.
For example, altitude and elevated temperature
are two conditions frequently encountered by
ultra-marathon runners (e.g. races in Nepal or in
deserts). Regarding altitude, using a sub-maximal
test until exhaustion in hypoxia/normoxia while the
muscles were maintained in identical complete
ischaemic conditions, we showed that (i) inhibitory
mechanisms from working muscles play a major
role in the cessation of the exercise in hypoxia and
that (ii) a minor but significant direct effect of
inspired oxygen fraction on the CNS could po-
tentiate this limiting mechanism and explain why
performance was slightly reduced in hypoxia.
Similarly, Amann et al.
showed that peripheral
fatigue measured with femoral nerve magnetic
stimulation at task failure was substantially less
severe in hypoxia compared with normoxia or
moderate hypoxia. This was attributed to brain
hypoxic effects on effort perception, leading the
subjects to stop earlier. Similar conclusions can
be deducted from hypoglycaemia
or hyper-
thermia experiments. Regarding this latter factor,
it is worth noting that high temperature does not
alter performance over brief contractions but
does cause reductions during sustained contrac-
or prolonged exercises.
Thus, sensory feedback contributes to central
fatigue and effort perception, presumably through
its indirect projection into the anterior cingulate
Other authors have argued that sensory
signals from peripheral receptors do not contribute
to perception of effort but generate other sensa-
tions experienced during exercise (e.g. muscle
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pain and thermal sensation
). In all cases, the
increase in RPE with fatigue is not abolished by
spinal blockade of somatosensory feedback from
the muscles; there must be other mechanisms.
Besides triggering inhibitory mechanisms, peri-
pheral fatigue implies that greater muscle activa-
tion is required for a given mechanical power to
be produced in the fatigued condition. Indeed, at
a given force or power output, the onset of fatigue
is usually concomitant with a rise in neuromus-
cular cost (EMG signal amplitude), which points
to the recruitment of additional motor units and/
or a higher discharge rate in order to compensate
for peripheral alterations. It is known that RPE
changes and the increase in muscle activity dur-
ing a constant-load exercise are correlated.
Marcora et al.
showed that the reduced loco-
motor muscle force after drop jumps resulted in
a higher RPE at a given power output and a re-
duced time to exhaustion during high-intensity
constant-power cycling. They suggested that the
effects were mediated by the increased central
motor command required to exercise with weaker
locomotor muscles,
which increased the percep-
tion of effort probably throughout its corollary
discharge to sensory areas of the brain.
Gagnon et al.
tested the effects of pre-induced
quadriceps fatigue (using electrostimulation) on
endurance performance of healthy individuals
and patients with chronic obstructive pulmonary
disease. These authors demonstrated that endurance
time significantly decreased by 2025%in the
experimental condition in both groups. There
has been hot debate about the role of afferent
feedback from fatigued locomotor muscles as
an important determinant of endurance exercise
Gagnon et al.
that that the enhanced metaboreflex is not the
main mechanism through which exercise toler-
ance was reduced in the fatigued state in both
study populations.
Moreover, a change in muscle efficiency after
ultra-marathon running
can affect muscle recruit-
ment to maintain a given task. Spinal inhibition
and/or disfacilitation (see section 2.1) after ultra-
marathon running could also necessitate higher
central command (i.e. reinforcing feed-forward
mechanisms) from supraspinal sites. Finally, the
gain of motoneurones decreases in fatigued con-
ditions such that additional synaptic drive at a
premotoneuronal level is required to maintain a
constant firing rate
(i.e. a larger descending drive
is needed to continue exercise at the same power
output). So, as well as the nociceptive signal coming
from the peripheral receptors, these feed-forward
mechanisms (also called the ‘sense of effort’
could also partly explain the RPE drift.
Interestingly, RPE for the same exercise could
vary with environmental conditions and is not
necessarily associated with a decrease in MVC
(e.g. at altitude).
In summary, the initial pace
and the adjustments made in response to these
feed-forward and feedback mechanisms
affect filling rate. RPE is probably affected by
both central command output and muscle affer-
In other words, as stated by Smirmaul,
an interaction between the sense of effort and the
sensations obtained from afferent sensory feedback
that is probably the ultimate regulator of exercise
performance. We suggest that for ultra-marathons
performed over hilly terrain, feed-forward and feed-
back mechanisms are mainly implicated in uphill/
flat sections and downhill sections, respectively.
3.4 Apart from the Filling Rate, Which Factors
Influence the Quantity of Water?
The second question is whether the volume of
water (absolute RPE) depends only on the filling
rate? The answer is clearly no. It has recently been
argued that it is an interaction between the sense
of effort and the sensations obtained from affer-
ent sensory feedback that is probably the ultimate
regulator of exercise performance.
while these two factors certainly play a crucial
role, it is, for example, possible to start an ex-
ercise with more water in the tank than usual (i.e.
with a higher RPE at the beginning of exercise
than is normally the case). Indeed, it is possible to
feel some fatigue without any exhaustive physical
load, for example, after a stressful day. More
importantly, Marcora et al.
reported that the
time to exhaustion at 80%of peak power output
was significantly reduced by ~15%after 90 minutes
of a demanding cognitive task. This was asso-
ciated with a higher RPE at the beginning of the
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cycling exercise compared with the control con-
dition. Since RPE increased similarly over time in
both conditions, mentally fatigued subjects reached
their maximal tolerated RPE and disengaged from
the cycling exercise earlier than did the controls.
Similarly, although one night of sleep depri-
vation does not usually affect MVC or intense
several studies have demonstrated a
deleterious effect on endurance performance.
Interestingly, a higher RPE for a given load has
been observed after sleep deprivation.
Some authors
showed that, after sleep de-
privation, subjects ran a shorter distance during a
30-minute self-paced treadmill exercise than did
their controls, yet their perception of effort was
similar. The authors suggested that altered per-
ception of effort may account for decreased en-
durance performance after sleep deprivation.
Other data show that RPE is not only dependant
on sensory information and cortical output. For
example, unknown or unexpected running exercise
duration may affect RPE,
suggesting that
RPE has an affective component. Also, changing
the tempo of music that cyclists were listening to
influenced their self-chosen power and cadence.
In summary, effort perception probably involves
the integration of multiple signals from a variety
of perceptual cues. Alternatively, as suggested in
section 3.6 for the amphetamines or pain killers,
it can be argued that perception is a complex
neurocognitive process that does not depend only
on the intensity of the sensory signal because
sensory signals are processed at brain level and
interpreted by the subject. It is then possible that
mental fatigue, music and sleep deprivation affect
the processing of sensory signals rather than
provide additional sensory signals. Nevertheless,
in that case, the rate of increase in RPE (filling
rate) would be changed rather than RPE at the
beginning of the exercise, as is the case, for ex-
ample, for mental fatigue.
3.5 The Waste Pipe
The third question is whether rest is the only
way to decrease the level of water in the tank. It is
the most obvious but probably not the only one.
For example, it may be possible to reduce the
water level while running using suitable psycho-
logical strategies.
Various psychological routines
can be used by runners to attenuate the discomfort
of intense physical exertion. These strategies, la-
belled ‘dissociative thoughts’ (i.e. the runner dis-
tracts him-/herself by thoughts of a more external
nature) are performed to diminish the sensations
of pain during a marathon.
Nevertheless, it
has been reported that the best marathon runners
adopt associative cognitive strategies (i.e. are
centered on their own sensations),
with the goal of maintaining optimal running
technique/efficiency and so decreasing peripheral
fatigue (and the filling rate). Another way to de-
crease the level of water in the tank might be
nutritional strategies. Chambers et al.
showed that rinsing the mouth with solutions
containing glucose and maltodextrin could im-
prove cycling performance. The authors suggested
that this could be due to activation of brain re-
gions involved in reward and motor control since
functional MRI measurements showed that these
regions believed to mediate emotional and be-
havioural responses to a rewarding sensory stimulus
were activated. Thus, in addition to its peripheral
action (i.e. slowing the tank filling rate), glucose
ingestion could have some central effects (i.e. de-
creasing the water level). Unidentified oral recep-
tors in the mouth could counteract the increase in
RPE, permitting higher central command and
power output.
Interestingly, the suggestion of downhill cycling
through hypnotic manipulation decreased RPE
without altering exercise HR or blood pressure
One could also suggest that redu-
cing maximal neural drive (i.e. central fatigue, a
decrease in %VA) may reduce the size of the tank.
In this context, Søgaard et al.
that central fatigue can only be demonstrated
during MVCs, but these authors suggested that
adecreasein%VA may have contributed to the
increase in RPE during sustained low-intensity
contractions (i.e. 15%MVC for 43 minutes). They
based this speculation on the fact that central fa-
tigue was among the factors that predicted RPE
changes. To our knowledge, no neurophysiogical
basis exists regarding the potential role of central
fatigue to explain at least in part the RPE changes.
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3.6 The Security Reserve
The fourth question is why exercise stops (in
the case of a time to exhaustion) or why the runner
decides to adjust his or her speed/power output
(in the case of a time trial). Assuming a constant
level of motivation, exercise cessation appears
to occur at the same RPE whatever the rising
and the starting level.
et al.
have incorporated Brehm’s theory a
general motivation theory that does not specifi-
cally refer to exercise to propose a psychobio-
logical model of endurance performance. Marcora
et al.’s model postulates that subjects decide to
withdraw effort (i.e. disengage) when an exercise
is perceived to be either too difficult or the effort
demanded exceeds the upper limit of what people
are willing to invest. Alternatively, it has been
proposed that exercise terminates when the feelings
of discomfort overwhelm the potential rewards of
continuing to exercise.
This was reviewed in
2003 by Kayser.
It is interesting to report that
an opiate antagonist, naloxone, leads to signif-
icant reductions in exercise performance when
compared with control trial.
The authors of
the article concluded that working capacity was
limited by the individual RPE, which can be at-
tenuated by endogenous opioids rather than by
physiological fatigue. Thus, it is not task failure
but task disengagement that sets the exercise limit
before reaching the security reserve. Even in highly
motivated competitors, task disengagement always
occurs before there is a threat to life.
do not usually exceed their security reserve. There
are a few exceptions where dramatic loss in body
homeostasis was reached causing collapse such as
during marathons or triathlons (e.g. Gabriela
Andersen-Schiess in the 1984 Olympics or Julie
Moss in the 1982 Hawaii Ironman), especially
in (i) hot environments with subjects not always
familiar with these environmental conditions (i.e.
unadapted template/sensor efficiency); or (ii) under
the effect of psychostimulants (e.g. Tom Simpson
who died).
One could pose the question whether elite
athletes finish with a lower security reserve. While
the common belief is that better athletes can ‘dig
deeper’ and work relatively harder than their less
successful counterparts,
we are not aware of
any scientific study supporting this concept. One
indirect argument has been proposed by Esteve-
Lanao et al.
who found that the pattern of
percentage maximum heart rate (%HR
) res-
ponse during an event was very similar in athletes
with large differences in running performance.
These authors concluded that better runners are
faster due to their underlying physiological ca-
pacity rather than to their ability to put greater
relative effort into their competition. Studies in-
vestigating the percentage of maximal oxygen
uptake ( .
) sustained during competition in
function of the level of performance are contra-
dictory. It was found that the faster running speed
of the more trained runners over 1090 km was
not due a higher %.
during competition
but was due to their superior running economy.
In contrast, performance was significantly related
to the specific endurance (i.e. the average speed
sustained over a 24-hour running exercise ex-
pressed in %.
Nevertheless, a higher
or %HR
sustained during competi-
tion does not necessarily mean that the elite ath-
letes can ‘dig deeper’ since this may be due to
physiological differences in terms of endurance.
Further studies must examine this interesting
A very important factor in the flush model
is that the sensor may be deregulated (i.e. the
interpretation of the incoming signal may be
In other words, the processing of
sensory signals is affected rather than additional
sensory signals provided. This is true in two op-
posing cases: amphetamines
(or more generally
when dopaminergic system is manipulated
and pain killers,
both of which induce higher
peripheral fatigue and/or metabolic disruptions.
For example, higher lactate/HR
or lower
peak doublet response to a magnetic stimulation
at the cessation of exercise has been reported.
Interestingly, while more anecdotal, Amann et al.
reported that all their subjects needed assistance
in disembarking the cycling ergometer after in-
jection of intrathecal fentanyl. In the evening (i.e.
several hours after exercise cessation), their sub-
jects reported continuing problems with ambu-
lation and muscle soreness, which had never been
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observed in any of the many other studies re-
questing exhaustion conducted in that laboratory.
It should be noted that psychostimulants (e.g.
amphetamines, cocaine) could act by providing a
pleasure sensation (i.e. water leak throughout the
waste pipe rather than sensor deregulated). Some
runners, even elite athletes, use local anaesthetics/
anti-inflammatory drugs (e.g. Tissugel
in France),
particularly applied to the knee joint. While this
is not prohibited by the World Anti-Doping
Agency policy, the flush model suggests that this
may be beneficial for improving performance in
ultra-marathon running.
According to the flush model, there is always
a reserve for muscle recruitment (the security re-
serve) that can be used for the so-called ‘end
when the runner is at his or her highest
level of peripheral fatigue. In ultra-marathons,
this is clearly illustrated in figure 2b representing
the HR data of an ultra-marathon runner per-
forming a race around the Mont-Blanc. In this
example, very particular race conditions (from
11th to 6th place with opponents regularly an-
nounced as potential targets) led the runner to
accelerate at the end of the race when his muscle
fatigue was higher than at ~22 hours. This further
illustrates that central regulation is not totally
based on peripheral changes. Since RPE was not
recorded, it is not possible to say whether this was
related to the enjoyment of overtaking several
opponents near the finish line, which counter-
acted the sensation of fatigue (i.e. the same level
of water despite increasing the power output with
some water being evacuated through the waste
pipe) and/or a decrease in the security reserve due
to increased motivation.
While mental processes are important in all
sports, it is probably particularly true for ultra-
marathon runners. Weir et al.
suggested that
the central governor is most applicable to endurance
exercise since the decline in muscle performance
under intensely fatiguing exercise conditions can
be directly attributed to peripheral fatigue. It is
further possible that for shorter distances, the
organism is equipped with a system to protect its
integrity, but at a peripheral level.
For ex-
ample, glycogen depletion may alter excitation-
contraction coupling, which in turn limits muscle
contraction capacity and so restricts muscle da-
mage. Also, it has been suggested that for exercise
of several minutes duration, RPE is increased
by sleep deprivation but when it is as short as
30 seconds, sleep deprivation causes only a small
change in the perception of exercise intensity
and thus no reduction in performance. Marino
et al.
recently argued that events such as the
marathon and, for example, walking 1000 km are
both quantitatively and qualitatively different,
and not simply because the 1000 km race is longer.
They proposed that it could be that the limits to
performance in a 1000-km race are predominantly
mental, while the limits to performance in the
marathon are predominantly physiological. Thus,
it can be suggested that ultra-marathon running
is an interesting model to study central regulation
of exercise. We believe that all interventions
designed to manipulate RPE and the sensation
of discomfort are of particular interest in this
4. Conclusion
As Marino et al.
recently pointed out,
Mosso concluded in his book La fatica (fatigue)
published 120 years ago,
that two phenomena
categorize fatigue, the diminution of muscular
force and the sensation of fatigue: ‘‘That is to say,
we have a physical fact which can be measured
and compared and a psychic fact which eludes
measurement.’’ Thus, the study of fatigue should
address both the perception of effort and the
decline in force that occurs during sustained ex-
The aim of the present article was to re-
view these two aspects of fatigue and to propose a
model that integrates the ‘neuromuscular’ and
‘physiological’ factors of fatigue (responsible for
maximal force reduction) in ultra-marathon
running to explain the regulation of performance.
It has been argued
that fatigue can be under-
stood as a highly regulated strategy conserving
cellular integrity, function and, indeed, survival.
The flush model, dedicated to integrating the fati-
gue mechanisms in ultra-marathon performance
(and more generally to any type of endurance
performance), using a holistic approach, is fully
compatible with this statement.
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The author would like to thank Professor Ken Nosaka for
his valuable comments on the manuscript and Wanda Lipski
for English language correction.
No sources of funding were used to conduct this study or
prepare this manuscript. The author has no conflicts of in-
terest that are directly relevant to this article.
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Correspondence: Professor Guillaume Millet, Laboratoire de
Physiologie de l’Exercice (EA 4338), Me
´decine du Sport-
Myologie, Ho
ˆpital Bellevue, 42055 Saint Etienne, Cedex 2,
506 Millet
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... The sense of effort is associated with neural or neuromuscular fatigue (Noakes, 2004). Neuromuscular fatigue is usually defined as strength loss after prolonged running exercises (Millet, 2011;Millet et al., 2003;Millet et al., 2011). ...
... Changes in running patterns due to fatigue can explain, at least in part, the alteration in running economy. While neuromuscular alterations may have consequences on the running pattern (Millet, 2011;Millet et al., 2011). Previous studies have investigated the effect of maximum fatigue, induced by a running protocol, have reported modifications to gait mechanics including increases in ankle and knee flexion during the stance phase of gait (Bazuelo-Ruiz et al., 2018;Derrick et al., 2002) and changes in vertical ground reaction forces (VGRF) (Bazuelo-Ruiz et al., 2018). ...
... Changes in running patterns due to fatigue can explain, at least in part, the alteration in the running economy, while neuromuscular alterations may affect the running pattern (Millet, 2011;Millet et al., 2011). Running-induced fatigue may increase vertical GRF as stresses on the tibia increase throughout a prolonged treadmill run (Rice et al., 2019). ...
Running can be defined as propulsion on the ground from one leg to the other. Ground reaction forces (GRF) are a critical element acting on the joints of the lower limbs, and the presence of fatigue has been shown to modify running biomechanics. This thesis aimed to identify, assess, and synthesize the current evidence on the effects of fatigue on GRF-related variables in long-distance runners. The PRISMA 2020 guidelines were followed. PubMed, Scopus, SPORTDiscus, and Web of Science were consulted in July 2022, followed by a search within reference lists and expert suggestions. Eligibility criteria: health competitive or recreational long-distance runners, performing running before and after an exercise-based fatigue-inducing protocol, and assessing GRF-related variables during running using a treadmill or overground with a force plate. Twelve studies (n = 299 participants, 177 ♀ & ♂122) were included. There was considerable heterogeneity regarding age, competitive level, distances covered, nature of interventions, comparators, and the testing methods, which inhibited performing a meta-analysis. Moreover, the included studies investigated different GRF-related variables (e.g., loading rate, active peak force, and peak ankle plantarflexor). The GRF-related variables showed fatigue-related changes in eleven studies, but not all changes were significant. Most consistently across four of twelve studies, loading rate increased (p = .01; p = .06; p = .04; p = .008), and impact peak force (Fpeak) significantly decreased (p < 0.05; p = 0.31[ES 0.40]; p < 0.005) post-fatigue. Three studies provided data for peak plantarflexor moment and, a significant decrease was observed in one study following the long run (p < .001; d 4.93 and CI [10.40 - 15.96]). Risk of bias in the studies was moderate to high, and certainty of evidence was judged as very low. Currently, the effects of fatigue on GRF-related variables in long-distance runners are uncertain, and it is unclear to what extent they may affect performance. Further studies should examine homogeneous samples to perform sensitivity analyses and investigate the potential effects of fatigue-inducing protocols on the GRF. Keywords: biomechanics; fatigue; ground reaction forces; kinetics; running.
... Pour investiguer la fatigue supra-spinale, la mesure de l'activation volontaire par l'utilisation de la stimulation magnétique transcranienne (Transcrannial Magnetic Stimulation, TMS) est possible. Le principe reste le même que la technique de twitch interpolation sauf que la stimulation au repos est réalisée à l'aide d'une extrapolation à partir d'une régression linéaire entre la stimulation surimposée et la force volontaire à l'aide de différentes contractions sous-maximales (Millet et al. 2011). La diminution de l'activation corticale volontaire, illustrée par l'augmentation de la force surimposée, serait indicative d'une incapacité de la commande descendante émanant des voies corticales motrices à activer l'ensemble des unités motrices (Todd et al. 2016). ...
... Cependant, pour un exercice identique, ne trouvaient pas de différence entre les groupes de PaSEPs fatigués et non fatigués, à la fois pour la fatigabilité et la RPE. Ces résultats suggèrent que lors d'un exercice corps entier (marche, vélo), la RPE pourrait être dépendante de la fatigabilité comme il a été suggéré en ultra marathon (Millet 2011) ou en cyclisme lors de courses contre la montre (Thomas et al. 2015). De ce constat, nous estimons que le schéma revisité de Enoka et al. (2021) ne fait pas mention d'un aspect sur lequel il nous semble important de mettre l'accent. ...
... Cependant, un autre courant de pensée explique que la RPE est modifiée par l'ensemble des afférences (ou « feedback ») provenant des muscles et autres organes (de Morree et. Ces deux visions continuent à animer les débats bien que récemment, ces deux concepts aient été groupés sous le concept de « tolérance sensorielle limite » (« Sensory tolerance limit »)(Hureau et al. 2018) et considérer dans le « modèle de la chasse d'eau » (« Flush model »)(Millet 2011). Dans les études sur la SEP, les avis divergent. ...
Full-text available
Multiple sclerosis (MS) is an autoimmune disease of the central nervous system, chronic fatigue being its main symptom. Despite its prevalence in MS, fatigue etiology remains incompletely understood. Moreover, exercise remains the only effective treatment for fatigue, but the effectiveness of a personalized intervention has not been thoroughly investigated. The aim of this PhD project was to characterize the causes of chronic fatigue in people with MS (PwMS) and to compare the benefits of an individualized exercise intervention to a conventional one in fatigued patients. In a first study, we demonstrated the high prevalence of fatigue in PwMS and the link with physical activity, disability or unemployment status. This first study reinforced the need to investigate fatigue etiology through a more comprehensive list of objective and subjective variables. In the second study, fatigued PwMS showed an increased fatigability, associated with a higher perception of effort, during dynamic whole-body exercise when compared to less fatigued PwMS. Finally, in a third study, we examined the effectiveness of an individualized vs a traditional exercise intervention in fatigued PwMS. Our results showed that the individualized program induced greater benefits, particularly in regards to exercise capacity, than the traditional program. This PhD thesis provides new knowledge on fatigue in MS and suggests potential clinical applications to improve the quality of life of PwMS.
... This decrease in muscle force has been shown to have both central (i.e., proximal to the neuromuscular junction) and peripheral (i.e., within the muscle) origins. The magnitude of isometric maximal voluntary force reduction seems to increase with the exercise duration during endurance running and tends to plateau around 1500 to 2000 min of exercise, with maximum values of strength loss around 35% to 40% in both KE and PF (16,17). Neuromuscular fatigue studies after multistage races are scarce, even when considering other sports than running. ...
... Neuromuscular fatigue. It has been reported that muscle strength loss increases with distance and exercise duration in endurance running until reaching a plateau (16,17). Despite the plethora of external parameters to consider in mountain ultramarathon, for example, weather, temperature, altitude, race profile and trail conditions, the decrease in muscle strength observed in this study on both muscle groups was similar to previous results for a similar duration (12). ...
... The SSR exhibited a greater decline in VA than MSR in KE at the end of the race (23% ± 17% vs 7% ± 12% for SSR and MSR respectively). The central component of fatigue has previously been described as a protection mechanism acting to limit fatigue and muscle damage (17,25). Although Temesi et al. (14) showed that VA measured by transcranial magnetic stimulation was reduced after a 110 km ultramarathon, the contribution of spinal modulation is still unknown (26). ...
... Obtaining perceived exertion is relatively easy, but practitioners often neglect some critical methodological issues extensively addressed when the scales are developed, which can subsequently compromise the validity of perceived exertion assessment [3][4][5]. In addition, several theoretical models have proposed that perceived exertion plays a role in explaining endurance exercise performance [8][9][10][11][12][13][14]. These models rely on assumptions about the origin of the neural signals responsible for generating the perceived exertion [8,11,13,15,16]. ...
... The intercept between exercise intensity and perceived exertion slopes approximates the critical power ( Figure 5). Importantly, this method of critical power approximation does not require exercise until exhaustion, as the perceived exertion slopes can be obtained from intermediate levels (11)(12)(13)(14) on the RPE scale) of perceived exertion [87]. ...
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The perceived exertion construct creation is a landmark in exercise physiology and sport science. Obtaining perceived exertion is relatively easy, but practitioners often neglect some critical methodological issues in its assessment. Furthermore, the perceived exertion definition, neurophysiological basis, and practical applications have evolved since the perceived exertion construct’s inception. Therefore, we revisit the careful work devoted by Gunnar Borg with psychophysical methods to develop the perceived exertion construct, which resulted in the creation of two scales: the rating of perceived exertion (RPE) and the category-ratio 10 (CR10). We discuss a contemporary definition that considers perceived exertion as a conscious perception of how hard, heavy, and strenuous the exercise is, according to the sense of effort to command the limbs and the feeling of heavy breathing (respiratory effort). Thus, other exercise-evoked sensations would not hinder the reported perceived exertion. We then describe the neurophysiological mechanisms involved in the perceived exertion genesis during exercise, including the influence of the peripheral feedback from the skeletal muscles and the cardiorespiratory system (i.e., afferent feedback) and the influence of efferent copies from the motor command and respiratory drive (i.e., corollary discharges), as well as the interaction between them. We highlight essential details practitioners should consider when using the RPE and CR10 scales, such as the perceived exertion definition, the original scales utilization, and the descriptors anchoring process. Finally, we present how practitioners can use perceived exertion to assess cardiorespiratory fitness, individualize exercise intensity prescription, predict endurance exercise performance, and monitor athletes’ responses to physical training.
... The elevation of OS and the inflammatory response during long-term exercises are closely related to increased pain (Ndengele et al., 2008) and muscle fatigue (Goulart et al., 2020, Theofilidis et al., 2018. The lower limbs receive a great load of activity during the running and there are many reports of muscle pain (Rowlands et al., 2012) and influence the increase in the withdrawal rate of competitions (Millet, 2011). Nonsteroidal antiinflammatory drugs (NSAIDs), such as ibuprofen (IBU), are commonly believed to reduce pain and muscle damage among trail runners in an attempt to increase their running performance (de Souza et al., 2020). ...
Up to 75% of marathon runners ingest non-steroidal anti-inflammatory drugs (NSAIDs) during competition. Despite the doubt whether or not they contribute to performance, the effect of NSAID in endurance sports is unclear. We evaluated the effect of ibuprofen (IBU) use on oxidative stress, muscle damage, physical performance, and vertical jump of runners participating in a 42-km-trail running. The sample consisted of 12 men randomly divided into 2 groups: a placebo group (placebo) and an ibuprofen group (IBG). A 400-mg IBU capsule was administered to the IBG 15 min prior to the start of the trial and during the course after 5 h. In the intergroup analysis, placebo 70.1% increase (p < 0.0001; Cohen's d = 4.77) of the thiobarbituric acid reactive substances (TBARS); the IBG exhibited a 31.46% increase of the sulphhydryl groups (SH) (p = 0.024, Cohen's d = 0.27), 55% of squat jump (SJ) (p < 0.01; Cohen's d = 1.41) with no significant effect on creatine kinase (CK), pace, speed, and finish time. In summary, IBU had positive evidence on oxidative stress and muscle fatigue, but had no effect on physical performance and muscle damage. Keywords: Ibuprofen; muscle damage; oxidative stress; performance; trail running.
... A pesar de que el RC puede ser establecido previo a la competición, éste no dejará de estar influenciado directamente durante la carrera por diversos factores como la capacidad de termorregulación del deportista (Cheuvront & Haymes, 2001), el nivel de deshidratación, la depleción de los depósitos de glucógeno (Rauch et al., 2005), la deriva pulsativa o cardiac drift (Billat et al., 2020), la fatiga neuromuscular (Millet, 2011), el incremento de la percepción subjetiva del esfuerzo ...
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El ritmo de competición (RC) es sumamente importante para establecer un óptimo pacing behaviour (PB) y así alcanzar un mayor rendimiento en la carrera a pie. Por lo tanto, el objetivo del presente estudio fue determinar posibles diferencias en el PB desarrollado en base al grupo etario y sexo durante la Maratón de Berlín. 43.970 atletas formaron parte del estudio, divididos en grupos de edad en base a lo establecido por la IAAF. Para determinar el PB, diez segmentos de distancia fueron tenidos en cuenta (0–5 km, 5–10 km, 10–15 km, 15–20 km, 20–21,1 km, 21,1–25 km, 25–30 km, 30–35 km, 35–40 km, 40–42,2 km). En los hombres se dió un grado de asociación significativo entre PB y los diferentes grupos de edad (X2 (26) = 380; p <0,01; CC=0,111). El negative pacing (NP) fue el perfil dominante en los grupos de edad masculinos. Las mujeres mostraron asociaciones significativas entre el grupo de edad y el PB desarrollado (X2 (24) = 178 p <0,01; CC=0,115), siendo el NP el más utilizado en casi todos los grupos etarios. Finalmente, los atletas más jóvenes tienen una mayor tendencia al even pacing (EP), mientras que los grupos de edad más avanzada tienen preferencia por el NP. No se observaron diferencias entre los perfiles más utilizados por sexo o grupo etario. Sin embargo, se pudo constatar que los corredores más rápidos desarrollaron un EP, para casi todos los grupos de edad y sexo.
... An important implication of these investigations is unveiling human physiological and psychological responses to the unique and extreme demands placed by an ultra-endurance exercise task. The majority of existing ultra-endurance studies have focused on running (28,32,35,39,40) and cycling (7,26,33) tasks, with very few studies investigating events compiling ultra-endurance of upper limbs as the chief exercising muscle group. Some existing works have investigated low-load repetitive International Journal of Exercise Science ...
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While neuromuscular fatigability has been previously characterized after running and cycling, no study has investigated an ultra-endurance upper body task. In preparation for a world record attempt, three pacing strategies to perform 1980 pull-ups in 6 hrs were compared during independent sessions: fast pace, long recovery (FL), fast pace, multiple short recoveries (FMS), and slow pace, no recovery (SN). Elbow flexion maximal voluntary contraction (MVC) force, grip strength, peripheral fatigue, and biceps brachii electromyography were quantified every 330 pull-ups and during recovery, alongside heart rate, perceived effort, and arm muscle pain. In all conditions, MVC force decreased rapidly within the first set of 330 pull-ups, with the greatest depression observed in FL (-29.1%) and more gradual declines in FMS (-18.6%) and SN (-8.6%). Similarly, FL displayed the greatest decline in potentiated single twitch (FL: -75.0%; FMS: -53.9%; SN: -41.8%) and high-frequency doublet forces (FL: -63.3%; FMS: -29.2%; SN: -41.8%) following the first set, as well as higher heart rate, effort, and pain throughout the task. Following 24 hrs, MVC force recovered slowest in FL and grip strength recovered fastest in SN. Therefore, for the world record attempt, a strategy with a continuous workload at slower pace should be used.
... Noakes ou le modèle de la chasse d'eau de Guillaume Millet (Millet, 2011)). À cet égard, plusieurs évidences démontrent que l'hyperthermie a un impact sur le système nerveux central, en augmentant la perception de l'effort et donc, la fatigue, ce qui réduirait la motivation à poursuivre l'exercice. ...
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(Thesis written in French) Dehydration, especially when ≥ 2% body mass, is usually associated with an alteration of physiological functions, subjective perceptions as well as endurance performance and cognitive functions. More specifically, it is frequently mentioned that together, the cardiovascular, thermal, metabolic impacts as well as subjective effects induced by dehydration could contribute to increase the perception of effort, which in turn would act as a mediator of endurance performance. While several studies have observed an exacerbation of the perception of effort with dehydration, evidence is contradictory, and some studies do not observe such an effect. In addition, evidence suggests that some individuals may better tolerate dehydration, while others are more widely affected. In this regard, a hypothesis has recently been put forward, according to which repeated exposure to dehydration could somewhat attenuate certain effects, in particular on subjective perceptions, including the perception of effort, and therefore reduce its subsequent impact on performance. In addition to being a possible key mediator in the relationship between exercise-induced dehydration and endurance performance, perception of effort could also play a central role in the process of habituation to the dehydration. However, the idea that humans can become habituated to dehydration has been discussed superficially and anecdotally. Several questions remain unanswered and must therefore be studied.
... The differences in metabolic profile between children and adults could also give some clues to the observed differences in fatigability. The exercise duration has a direct influence on fatigability, with longer exercise inducing greater decrement muscle in performance (Millet, 2011). Then, it would have been incorrect to analyze fatigability data for experimental designs that used exercises of different duration. ...
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The transition from childhood to adulthood is characterized by many physiological processes impacting exercise performance. Performance fatigability and time to task failure are commonly used to capture exercise performance. This review aimed to determine the differences in fatigability and TTF between youth (including both children and adolescents) and young adults, and to evaluate the influence of exercise modalities (i.e., exercise duration and type of exercise) on these differences. Medline, SPORTDiscus and Cochrane Library were searched. Thirty-four studies were included. The meta-analyses revealed that both children (SMD −1.15; p < 0.001) and adolescents (SMD −1.26; p = 0.022) were less fatigable than adults. Additional analysis revealed that children were less fatigable during dynamic exercises (SMD −1.58; p < 0.001) with no differences during isometric ones (SMD –0.46; p = 0.22). Children (SMD 0.89; p = 0.018) but not adolescents (SMD 0.75; p = 0.090) had longer TTF than adults. Additional analyses revealed 1) that children had longer TTF for isometric (SMD 1.25; p < 0.001) but not dynamic exercises (SMD −0.27; p = 0.83), and 2) that TTF differences between children and adults were larger for short- (SMD 1.46; p = 0.028) than long-duration exercises (SMD 0.20; p = 0.64). Children have higher endurance and are less fatigable than adults. These differences are influenced by the exercise modality, suggesting distinct physiological functioning during exercise between children and adults. The low number of studies comparing these outcomes between adolescents versus children and adults prevents robust conclusions and warrants further investigations in adolescent individuals.
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Background Over the past decade, numerous advances have been made in the research on electrical stimulation of skeletal muscle. However, the developing status and future direction of this field remain unclear. This study aims to visualize the evolution and summarize global research hot topics and trends based on quantitative and qualitative evidence from bibliometrics.Methods Literature search was based on the Web of Science Core Collection (WoSCC) database from 2011 to 2021. CiteSpace and VOSviewer, typical bibliometric tools, were used to perform analysis and visualization.ResultsA total of 3,059 documents were identified. The number of literature is on the rise in general. Worldwide, researchers come primarily from North America and Europe, represented by the USA, France, Switzerland, and Canada. The Udice French Research Universities is the most published affiliation. Millet GY and Maffiuletti NA are the most prolific and the most co-cited authors, respectively. Plos One is the most popular journal, and the Journal of Applied Physiology is the top co-cited journal. The main keywords are muscle fatigue, neuromuscular electrical stimulation, spinal cord injury, tissue engineering, and atrophy. Moreover, this study systematically described the hotspots in this field.Conclusion As the first bibliometric analysis of electrical stimulation of skeletal muscle research over the past decade, this study can help scholars recognize hot topics and trends and provide a reference for further exploration in this field.
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Nine experienced endurance runners performed individual marathon runs that involved several tests of neuromuscular performance before, during and after the marathon. The tests were performed with special force platform and dynamometer techniques. The results showed an overall decrease in performance from the marathon. The maximal sprint velocity decreased parabolically during the marathon, reaching the final value of 84% of the pre-marathon one. Similarly, the other test results after marathon indicated that maximal isometric knee extension torque was 78%, the performance in a special rebound test (drop jump) 84% and the 5-jump performance 92% of the pre-marathon values. These reductions were accompanied by alteration in the ground reaction force curves in the sprint and jump tests, suggesting reduced tolerance to stretch load as well as loss in the recoil characteristics of the muscles.
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The unique physiological attributes of marathoners have long been recognised, but until the pioneering research of Morgan and Pollock (1977) little was known about their psychological characteristics. Their work revealed marathoners have significantly better mental health compared with non-athletes, with desirable mental health variables being most pronounced in elite competitors. It was also found that during competition, elite marathoners typically utilise a unique cognitive strategy labelled ‘association’, whereby they regulate pace based upon bodily sensations including pain and effort. More recent research indicates there are considerable individual differences in the psychological responses of marathoners to the stressors associated with training and competition, and in some cases negative emotions traditionally presumed to be harmful actually benefit performance. This brief review will highlight findings of psychological research involving marathoners and other endurance athletes, distinguishing between characteristics common among groups (i.e. nomothetic) with those particular to individuals (i.e. ideographic) or sub-groups of elite and non-elite competitors.
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The purpose of this study was to determine if runners who completed a 100 km ultramarathon race in the fastest times changed their running speeds differently compared to those runners who ran an overall slower race. Times were taken from the race results of the 1995 100 km IAU World Challenge in Winschoten, Netherlands. Race times and 10 km split times were analyzed. Runners (n = 67) were divided into groups of ten with the last group consisting of seven runners. The mean running speed for each 10 km segment was calculated using each runner's 10 km split times. Mean running speed was calculated using each runner's race time. The first 10 km split time was normalized to 100, with all subsequent times adjusted accordingly. The mean running speed for each group at each 10 km split was then calculated. The faster runners started at a faster running speed, finished the race within 15 % of their starting speed, and maintained their starting speed for longer (approximately 50 km) before slowing. The slower runners showed a greater percentage decrease in their mean running speed, and were unable to maintain their initial pace for as long. It is concluded that the faster runners: 1) ran with fewer changes in speed, 2) started the race at a faster running speed than the slower runners, and 3) were able to maintain their initial speed for a longer distance before slowing. Key PointsFaster runners in the 100 km race;ran with fewer changes in running speed compared to the slower runners;started the race at a faster running speed than the slower runners;were able to maintain their initial running speed for longer distances than slower runners.
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The influence of an individual marathon race was investigated among 7 subjects who performed 2 types of unilateral isometric strength tests before and after the race. The tests included maximal isometric voluntary actions (MVA) of the knee extensor muscles and isometric endurance of 60% of pre-marathon MVA. The results showed dramatic reductions of the maximal isometric torque (26 ± 14%) and endurance time (39 ± 9%). Maximal integrated electromyographic activity (IEMG) of the vastus medialis muscle decreased by 36 ± 26% and that of the vastus lateralis muscle by 42 ± 25%. In the submaximal endurance test, the post-marathon isometric knee extension started at a higher level of IEMG and lower mean power frequency (MPF) for 6 of 7 subjects. These data suggest that the repeated stretch-shortening cycles during the marathon race affected the force production by reducing neural input to the muscles and deteriorating the efficacy of the contractile mechanism.
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The aim was to test the hypothesis that one night of sleep deprivation will impair pre-loaded 30 min endurance performance and alter the cardio-respiratory, thermoregulatory and perceptual responses to exercise. Eleven males completed two randomised trials separated by 7 days: once after normal sleep (496 (18) min: CON) and once following 30 h without sleep (SDEP). After 30 h participants performed a 30 min pre-load at 60% [VO(2 max) followed by a 30 min self-paced treadmill distance test. Speed, RPE, core temperature (T(re)), mean skin temperature (T(sk)), heart rate (HR) and respiratory parameters VO(2 max), VCO(2), VE, RER pre-load only) were measured. Less distance (P = 0.016, d = 0.23) was covered in the distance test after SDEP (6037 (759) 95%CI 5527 to 6547 m) compared with CON (6224 (818) 95%CI 5674 to 6773 m). SDEP did not significantly alter T(re) at rest or thermoregulatory responses during the pre-load including heat storage (0.8 degrees C) and T(sk). With the exception of raised VO(2) at 30 min on the pre-load, cardio-respiratory parameters, RPE and speed were not different between trials during the pre-load or distance test (distance test mean HR, CON 174 (12), SDEP 170 (13) beats min(-1): mean RPE, CON 14.8 (2.7), SDEP 14.9 (2.6)). In conclusion, one night of sleep deprivation decreased endurance performance with limited effect on pacing, cardio-respiratory or thermoregulatory function. Despite running less distance after sleep deprivation compared with control, participants' perception of effort was similar indicating that altered perception of effort may account for decreased endurance performance after a night without sleep.
The original central fatigue hypothesis suggested that an exercise-induced increase in extracellular serotonin concentrations in several brain regions contributed to the development of fatigue during prolonged exercise. Serotonin has been linked to fatigue because of its well known effects on sleep, lethargy and drowsiness and loss of motivation. Several nutritional and pharmacological studies have attempted to manipulate central serotonergic activity during exercise, but this work has yet to provide robust evidence for a significant role of serotonin in the fatigue process. However, it is important to note that brain function is not determined by a single neurotransmitter system and the interaction between brain serotonin and dopamine during prolonged exercise has also been explored as having a regulative role in the development of fatigue. This revised central fatigue hypothesis suggests that an increase in central ratio of serotonin to dopamine is associated with feelings of tiredness and lethargy, accelerating the onset of fatigue, whereas a low ratio favours improved performance through the maintenance of motivation and arousal. Convincing evidence for a role of dopamine in the development of fatigue comes from work investigating the physiological responses to amphetamine use, but other strategies to manipulate central catecholamines have yet to influence exercise capacity during exercise in temperate conditions. Recent findings have, however, provided support for a significant role of dopamine and noradrenaline (norepinephrine) in performance during exercise in the heat. As serotonergic and catecholaminergic projections innervate areas of the hypothalamus, the thermoregulatory centre, a change in the activity of these neurons may be expected to contribute to the control of body temperature whilst at rest and during exercise. Fatigue during prolonged exercise clearly is influenced by a complex interaction between peripheral and central factors.
At least 30 minutes of moderate-intensity physical activity accumulated on most, preferably all days is considered the minimum level necessary to reduce the risk of developing cardiovascular disease. Despite an unclear explanation, some epidemiological data paradoxically suggest that a very high volume of exercise is associated with a decrease in cardiovascular health. Although ultra-endurance exercise training has been shown to increase antioxidant defences (and therefore confer a protective effect against oxidative stress), an increase in oxidative stress may contribute to the development of atherosclerosis via oxidative modification of low-density lipoprotein (LDL). Research has also shown that ultra-endurance exercise is associated with acute cardiac dysfunction and injury, and these may also be related to an increase in free radical production. Longitudinal studies are needed to assess whether antioxidant defences are adequate to prevent LDL oxidation that may occur as a result of increased free radical production during very high volumes of exercise. In addition, this work will assist in understanding the accrued effect of repeated ultra-endurance exercise-induced myocardial damage.
Striding bipedalism is a key derived behaviour of hominids that possibly originated soon after the divergence of the chimpanzee and human lineages. Although bipedal gaits include walking and running, running is generally considered to have played no major role in human evolution because humans, like apes, are poor sprinters compared to most quadrupeds. Here we assess how well humans perform at sustained long-distance running, and review the physiological and anatomical bases of endurance running capabilities in humans and other mammals. Judged by several criteria, humans perform remarkably well at endurance running, thanks to a diverse array of features, many of which leave traces in the skeleton. The fossil evidence of these features suggests that endurance running is a derived capability of the genus Homo, originating about 2 million years ago, and may have been instrumental in the evolution of the human body form.