Sports Med 2003; 33 (3): 167-176

CURRENT OPINION 0112-1642/03/0003-0167/$30.00/0

© Adis Data Information BV 2003. All rights reserved.

The Conscious Perception of the

Sensation of Fatigue
Alan St Clair Gibson,1,2 Denise A. Baden,3 Mike I. Lambert,2 E. V. Lambert,2
Yolande X. R. Harley,2 Dave Hampson,2 Vivienne A. Russell2 and Tim D. Noakes2
1 Human Motor Control Unit, National Institute of Neurological Disorders and Stroke, National
Institutes of Health, Bethesda, Maryland, USA
2 Research Unit of Exercise Science and Sports Medicine, Department of Human Biology,
University of Cape Town, Cape Town, South Africa
3 Department of Psychology, University of Southampton, Southampton, UK

Abstract In this review, fatigue is described as a conscious sensation rather than a

physiological occurrence. We suggest that the sensation of fatigue is the conscious
awareness of changes in subconscious homeostatic control systems, and is derived
from a temporal difference between subconscious representations of these homeo-
static control systems in neural networks that are induced by changes in the level
of activity. These mismatches are perceived by consciousness-producing struc-
tures in the brain as the sensation of fatigue. In this model, fatigue is a complex
emotion affected by factors such as motivation and drive, other emotions such as
anger and fear, and memory of prior activity. It is not clear whether the origin of
the conscious sensation of fatigue is associated with particular localised brain
structures, or is the result of electrophysiological synchronisation of entire brain
activity.

Fatigue is experienced by all individuals on a However, there is no knowledge at present of how

regular basis. There are many different definitions
of fatigue. In medical practice, fatigue is described
as a debilitating consequence of a number of differ-
ent systemic diseases or nutritional deficits.[1] In
exercise physiology, fatigue is described as an acute
impairment of exercise performance, which leads to
an eventual inability to produce maximal force out-
put as a consequence of metabolite accumulation or
substrate depletion.[2] In neurophysiology, fatigue is
described as a reduction in efferent motor com-
mands to the active muscles resulting in a decline in
force or tension as part of a centrally controlled
process.[3]
Fatigue is therefore a ubiquitous phenomenon
with different meanings in different research fields.
the conscious‘feeling’ of fatigue is created, or from
where it originates. In this article, therefore, we
examine fatigue as a sensation rather than a physical
process. The aim of the article is to discuss the
concept of fatigue from psychological, anatomical
and physiological perspectives, and to describe pos-
sible mechanisms and brain regions responsible for
the conscious perception of the sensation of fatigue.
1. Conscious Awareness of the
Sensation of Fatigue
Before examining the mechanisms responsible
for the conscious sensation of fatigue, it is necessary
to discuss briefly the more general issue of con-
sciousness itself. There are two basic theories of
168
consciousness, which have been debated for centu-
ries. The first is the theory of dualism, which sug-
gests that consciousness is a mental state which
exists autonomously and is not dependent on brain
structures for existence.[4] The second is the theory
of monism, which suggests that consciousness is a
direct product of activity in specific brain struc-
tures.[5,6] While this argument is complex and be-
yond the realm of this review, the theory of dualism
is more difficult to support from a scientific perspec-
tive, and remains a philosophical concept (Hallett
M, unpublished observation). In this review, we
therefore discuss conscious perception of the sensa-
tion of fatigue from the monistic perspective.
Consciousness has two distinct components. The
first is the ‘awareness’ of something, for example
awareness of activity occurring around an individ-
ual. The second is the ‘feeling of knowing’ of that
awareness. This ‘feeling of knowing’ allows us to
have knowledge that the awareness is specific to our
own life, and creates the special subjective ‘feeling’
each individual has which makes us aware that what
is going on around us is unique to our own individu-
al participation in that experience. This second com-
ponent makes consciousness in different individuals
a subjective phenomenon, defined as ‘qualia’,[7]
which makes the life of each different individual
unique, but which makes consciousness so difficult
to examine scientifically. An example of this would
be that the sensation of fatigue at any time point is
completely unique to the individual perceiving it.
While the level of sensation of fatigue associated
with different levels of activity can be fairly well
quantified in different individuals, at present observ-
ers cannot ‘feel’ the ‘exact’ feeling of the person
being studied when they describe that individual’s
sensation of fatigue. However, recent work, which
will be discussed later in this article, has begun to
shed light on the subjective nature of conscious
perception.
In relation to fatigue, the first component of
conscious perception is easier to define. The con-
scious sensation of fatigue would occur when altera-
tion to a physical state induced by physical activity
leads to changes in activity in brain neural networks,
© Adis Data Information BV 2003. All rights reserved.
St Clair Gibson et al.
which in turn cause ‘awareness’ of those alterations.
This origin of the conscious perception of fatigue
may therefore be determined by examining changes
in different brain structures associated with the de-
velopment of awareness of the sensation of fatigue.
2. Brain Structures Involved in the
Sensation of Fatigue
There have been several theories of how the
sensation of fatigue is activated in neural networks
in different brain structures. In the peripheral model
of fatigue, changes in a certain factor in the peri-
pheral organs during physical activity, such as mus-
cle metabolite accumulation,[8-11] substrate deple-
tion[12-16] or cardiac and respiratory limitations,[17-19]
stimulate mechanoreceptors[20-22] or metaborecep-
tors[23-25] resulting in afferent feedback to the brain
which induces the sensation of fatigue.[26-28] As there
is no conclusive evidence to support any single
factor as being directly responsible for the onset of
the sensation of fatigue,[29] it is also suggested that a
number of different afferent inputs, together with
other non-sensory inputs such as psychological and
motivational factors, are integrated in brain struc-
tures and the ensemble leads to the development of
the sensation of fatigue which arises directly from
these integrative brain structures.[20,29-32]
This latter hypothesis is supported by three dif-
ferent observations. Firstly, the sensation of fatigue
can be altered by hypnosis while individuals main-
tain constant work output.[33-35] Secondly, the sensa-
tion of fatigue is affected by expectations of task
demands, being greater at the same absolute time-
point in races of similar intensity but shorter dura-
tion compared with those of longer duration (Baden
DA et al., unpublished data) and lower when actual
exercise duration is shorter than expected.[36] Third-
ly, in the clinical condition known as chronic fatigue
syndrome, an excessive and debilitating sensation of
fatigue is present at rest, and is not related to altera-
tion in physical activity levels.[37-39] These observa-
tions suggest that the origin of the sensation of
fatigue must be associated with anatomical brain
structures responsible for conscious perception,
Sports Med 2003; 33 (3)
Perception of Fatigue
rather than from any single peripheral physiological
perturbation or altered metabolic state.
The sensation of fatigue is itself a complex phe-
nomenon. It is associated with the conscious percep-
tion of changes in body functions, such as breath-
lessness from increased ventilation, pounding of the
heart from cardiac output increase, the sensation of
being hot and sticky from temperature increases and
sweating, and the sensation of increased muscle
activity associated with increased power generation
caused by increased physical activity levels.[29]
There are also mental cognitive functions associated
with the sensation of fatigue. These include level of
motivation at the time of the activity, memory of
prior exercise sessions and an associated decision-
making component based on the relationship be-
tween the current sensorimotor input and memory of
previous events.[32]
Because of the complex nature of the sensation of
fatigue, the regions of the brain from which the
sensation of fatigue originates have been difficult to
assess, even with the development of modern
neuroscience techniques, such as functional magnet-
ic resonance imaging, dynamic electroencepha-
lography (EEG) and repetitive transcranial magnetic
stimulation (figure 1). This may be because a num-
ber of different brain regions are involved, or be-
cause general processes involving energy changes
throughout the brain are responsible for the genera-
tion of the sensation of fatigue.
The development of the sensation of fatigue is
associated with changes in motor activity.[40] As
motor function is controlled by the motor and
premotor cortex, supplementary motor cortex, basal
ganglia and cerebellum,[41-43] changes in neural net-
work activity in any of these areas may be responsi-
ble for the generation of the sensation of fatigue.
Indeed, it has been suggested that the ‘effort of
will’[44] or ‘sense of effort’[45] is corollary discharge
associated with motor activity.[40,46] However, more
recent work has suggested that the sensation of
effort and the sensation of fatigue may not be gener-
ated in the motor cortex and rather originates in
motor command centres ‘upstream’ from the motor
cortex.[40,47]
© Adis Data Information BV 2003. All rights reserved.
169
Motor
cortex
Cingulate cortex
Frontal
cortex
pus
am
i p poc dala
H my g
A
Cerebellum
Brain stem
Spinal cord
Fig. 1. Putative brain regions associated with the development of
the sensation of fatigue.
Fatigue is also related to motivation and emotion-
al state.[48,49] Indeed, it has previously been suggest-
ed that fatigue is an emotion itself.[50] Emotional
states such as anger, fear and possibly also fatigue,
are not free standing psychological concepts, but are
distinct collections of chemical and neural responses
to particular emotional stimuli that have related
physical symptoms, which in some cases can be
partially quantified.[50,51] Emotional responses are
thought to be triggered by the amygdala and ventro-
medial prefrontal cortex, and are executed by the
hypothalamus and brainstem nuclei.[50] The main
targets of the emotional responses are the viscera
and musculoskeletal system, but other targets in the
brain are also affected including monoaminergic
neurons in the brainstem that modulate behaviour
and memory formation.[52] Emotional states such as
anger and fatigue involve adjustments in homeostat-
ic balance and peripheral physiological changes,
such as heart and respiratory rate which can be
scientifically measured.[50] Feelings of anger or fa-
tigue may therefore be the mental representations of
the physiological changes which characterise these
emotions. Functional imaging studies have revealed
Sports Med 2003; 33 (3)
170
increased or decreased activity in several brain areas
including the orbitofrontal cortex, anterior cingulate
cortex and insular cortex, when normal individuals
experience emotions such as sadness, happiness,
fear or anger.[52-57] Therefore, as motivation and
drive are affected by all these different emotional
states, fatigue may originate in any of these different
brain areas associated with emotional responses.
Previous experience and planning of current ac-
tivity are also important components of the sensa-
tion of fatigue. Previous memory of fatigue will
allow one to estimate one’s reserves and tolerance
levels, and allow decision making to occur as to
whether to continue, reduce activity or halt the phys-
ical activity. Memory formation and decision mak-
ing involve representations of sequences of events
(episodic, personal experiences) within a framework
of general semantic knowledge (facts and places),
linked by their common events and places.[58] The
hippocampus, together with the parahippocampus
and surrounding areas of temporal lobe neocortex,
synthesise these episodic representations into
unique single experiences, such as a specific exer-
cise activity.[58] The transition from immediate to
long-term memory is associated with activity in the
prefrontal cortex.[59] In the prefrontal cortex, current
activity is compared with previously stored memory
of previous activity, and the current activity would
be altered appropriately if differences between these
events were unacceptable as part of decision-mak-
ing processes in the prefrontal cortex. Working
memory is active only for as long as cells in the
prefrontal cortex are firing. Therefore, it would be in
these prefrontal cortical systems, perhaps, that the
origin of the conscious perception of the sensation
of fatigue arises from.
Recently, Parvizi and Damasio[60] have suggested
that the brainstem and spinal cord may be the area
where consciousness is modulated. The reason they
proposed that this area was responsible for the origin
of consciousness was, firstly, because the reticular
activating system, which sets the level of elec-
trophysiological activity in the cortex and most oth-
er areas in the brain, arises from the reticular nuclei
of the brainstem, and individuals with damage to the
© Adis Data Information BV 2003. All rights reserved.
St Clair Gibson et al.
brainstem region have impaired consciousness. Sec-
ondly, the brainstem is the site of arrival of afferent
input from nociceptors, the vestibular system, mus-
culoskeletal system and integrative interneurons,
and therefore could be an area where integration of
these different signals occurred. Thirdly, there are a
number of connections between the hypothalamus
and brainstem nuclei, and through these connections
knowledge of homeostatic perturbations could be
integrated. Fourthly, brainstem nuclei are involved
in the dopaminergic, cholinergic and other neuro-
transmitter systems which alter the mode of process-
ing of other brain structures and areas responsible
for mood, memory or perceptual changes. There-
fore, these areas in the brainstem and spinal cord
may also be associated with the sensation of fatigue.
It is therefore difficult to show that the sensation
of fatigue is localised to a single area of the brain, as
a number of different cognitive functions and in-
tegrative brain regions may be involved with
processes that lead to the initiation of the sensation
of fatigue. It has also been suggested that no region-
al classification of the brain is correct for any func-
tion, as all areas of the brain are involved during all
activities, albeit to different degrees.[61] This concept
is encapsulated by the ‘binding problem’, which
describes the problem of binding together represen-
tations of different properties of an object or physi-
cal state such as it’s colour, form or location,[62] and
suggests that large areas of the brain must be in-
volved in complex cognitive tasks or motor activi-
ties. It has been suggested that large scale syn-
chronisation of oscillatory electrical activity in the
neural circuitry of different brain areas or in local
brain areas,[63,64] controls the temporal sequences of
task activity. This phase synchrony, or coherence of
activity in different brain regions is also found to
occur in EEG and electromyogram activity during
motor tasks, generally in the gamma rhythm range
(25–70Hz),[62-64] indicating that central and peri-
pheral neural systems may communicate using syn-
chronisation processes or activity. It is not clear how
fatigue influences this synchronisation of electrical
activity, or whether alterations in synchronisation
patterns induce the sensation of fatigue during func-
Sports Med 2003; 33 (3)
Perception of Fatigue
tional activities, and further work is necessary in this
field.
Alterations in neurotransmitter concentrations in
various brain structures have been suggested to
cause fatigue. It is not clear whether neurotransmit-
ter concentration changes are caused by changes in
neurotransmitter precursor concentrations in peri-
pheral tissue which cross the blood-brain barri-
er,[65,66] or whether activation of brainstem struc-
tures which control release of neurotransmitters in
the higher cortical structures are altered or initiated
by fatiguing activity and lead to the release of exci-
tatory or inhibitory neurotransmitters which may
alter cognitive function or perception of the fatigue
state. For example, monoaminergic nuclei of the
reticular formation in the brainstem are responsible
for serotonin, dopamine and noradrenaline changes
in the cortical mantle, and these neurotransmitters
are involved in the modulation of global activity in
the cortex, and lead to changes in attentiveness and
behavioural responses to external stimuli.[60] Neuro-
transmitters, such as acetylcholine, have also been
found to modulate synchronisation and generation
of cortical electrical activity in a concentration-
dependent manner.[62] Therefore, neurotransmitter
concentration changes may alter the cognitive re-
sponse to or be involved in the subconscious fatigue
generating mechanisms.
Alterations in substrate availability in brain tis-
sue, associated with increased metabolic demands
and caused by increased mental activity during exer-
cise, have also been suggested to cause fatigue. For
example, it has been postulated that during maximal
activity, increased cognitive activity increases brain
metabolism, leading to energy demands exceeding
production of energy substrates such as brain glu-
cose and lactate.[67,68] However, if brain substrate
depletion is the cause of the sensation of fatigue, one
would expect there to be a period of time after
intense physical activity when all brain function is
impaired. As this does not occur,[40] it is more likely
that if substrate depletion occurs it would modulate
rather than cause the perception of the sensation of
fatigue.
© Adis Data Information BV 2003. All rights reserved.
171
In summary, if the sensation of fatigue is a com-
ponent of cognitive decision making or active set-
ting of exercise intensity as a part of the working
memory system and based on prior experience, the
most likely region involved would be the prefrontal
cortex.[59] If fatigue is a specific type of emotional
state, the amygdala or hippocampus or a combina-
tion of these areas may be involved.[50,58] If the
sensation of fatigue is part of an integrative met-
abolic regulatory system balancing afferent input
with efferent command, the hypothalamus or brain-
stem and spinal cord regions may be the site of
origin.[43,60] Alternatively, fatigue as a sensation may
occur as activity processes in the brain which are
currently not determined and which involve many
different areas of the brain at any one timepoint.
3. Conscious Perception from
Subconscious Activities
While we have examined putative structures and
processes that may be responsible for the origin of
the sensation of fatigue, we have not yet discussed
how the sensation of fatigue originates from the
underlying neural networks which regulate most
functional processes subconsciously. A number of
theories have been proposed to explain how con-
sciousness develops, such as the global work-
space[69] and neural oscillation theories,[7] but no
scientific evidence has been able to clearly substan-
tiate any of these theories and a detailed discussion
of each of these is beyond the realm of this review.
However, recently, Damasio[51] has suggested a
model for how conscious perception develops from
subconscious processes, which may be particularly
relevant as a mechanism to describe how the sensa-
tion of fatigue originates. In an attempt to describe
the association between consciousness and subcon-
scious control processes, Damasio and Parvisi[51,60]
suggested that consciousness is underpinned by a
subconscious ‘proto-self’. In this model, the ‘proto-
self’ controls or maps the physiological changes
associated with emotions, such as increased heart
rate and sweating, and although this proto-self also
operates at a subconscious level, these changes gen-
erate ‘feelings’ which are our conscious response or
Sports Med 2003; 33 (3)
172
manifestation of these emotional changes. In this
model of consciousness, the proto-self is a coherent
collection of neural patterns which map the state of
physical and physiological structures of the organ-
ism. This proto-self becomes the homeostatic set-
point of the organism and is a permanent first-order
map. When changes in the external environment or
internal physiological milieu occur, these changes
become a further first-order map. When this altered
environment is compared with the proto-self, the
differences between the two become second-order
descriptive maps which describe the relationship
between the proto-self and the altered environment.
This second-order map becomes a ‘mental’ image,
which resides at a subconscious level. For the
mental image to become a conscious image, the
mental image needs to induce a ‘feeling’ or emotion,
which allows our conscious cognitive processes to
be aware that a change has occurred which is rele-
vant and requires avoidance behaviour if the emo-
tion or feeling is negative, such as fear or fatigue.
The hypothesis of the proto-self thus centres on a
relationship between the changing state of an organ-
ism and the sensorimotor first-order maps, or the
proto-self, which describes an individual’s internal
physiological and physical set-point. As changes in
the individual occur, another level of second-order
mapping creates an account of the events taking
place as a consequence of the proto-self/changed
environment interaction. The essence of conscious
knowledge of these changes occurs when a ‘feeling’
or emotion, such as fatigue, is induced by this sec-
ond-order map, which allows responses to the
changed state to be initiated. In this model, there-
fore, because conscious knowledge originates in
neural structures associated with the proto-self rep-
resentation of body states, consciousness therefore
is in effect the ‘feeling’ itself, such as fatigue or
other emotional states.
This model may be relevant for describing how
the sensation of fatigue originates. For example, at
the beginning of an exercise bout, the proto-self
first-order map would describe the homeostatic set-
points for all metabolic activity at rest in the body,
and would serve as the reference point for the activi-
© Adis Data Information BV 2003. All rights reserved.
St Clair Gibson et al.
ty to follow. As exercise begins, the change in
metabolic conditions causes a different first-order
map to be created, and the difference between these
two maps creates a second-order map which creates
a conscious perception of change and thus ‘aware-
ness’ of the activity. As exercise continues, further
second-order maps are created of the ongoing
change in metabolic profile, and these metabolic
maps must create a cumulative perception of change
that eventually is recognised as the conscious per-
ception of the sensation of fatigue (figure 2).
This model is interesting also in that it explains
both the ‘awareness’ of conscious sensations such as
fatigue, and the ‘feeling of knowing’ of the aware-
ness of theses conscious sensations, which as de-
scribed previously, has traditionally been the more
difficult concept to explain. In Damasio’s[51] theory
of emotion underlying consciousness, physiological
changes result in the ‘feeling’ of emotions, and these
feelings ultimately represent the ‘feeling’ which we
attach to consciousness. Therefore, what gives qua-
lia their particular identity are the specific physio-
logical changes which occur at the exact timepoint
when a second-order representation is perceived.
Damasio’s theory may therefore be relevant for un-
derstanding how the conscious perception of the
sensation of fatigue arises. It is, however, at present
still a theoretical model and further work is needed
to assess its validity.
It is not clear when the exact point when aware-
ness of change, as described by second-order maps,
becomes a conscious awareness of fatigue. It has
recently been suggested that vocalisation processes
may be necessary for this acknowledgement of al-
tered mapping or evolving second-order states.[70] In
this model, inner self-talk alerts one to significant
changes in proto-self maps, and this self-talk is
associated somehow or as part of a second-order
neural network that involves the parts of the cerebral
cortex responsible for vocalisation. Further work is
needed to examine this theory, looking at these
cortical areas specifically using functional imaging
techniques during fatiguing activity.
Finally, it must be noted that while the sensation
of fatigue represents our conscious knowledge of
Sports Med 2003; 33 (3)
Perception of Fatigue
Second order map 1
Proto self
First order map
Fig. 2. The relationship between the ‘proto-self’ and physiological changes associated with increased physical activity and the generation of
the conscious perception of the sensation of fatigue.
underlying physical activity, the majority of activity
is controlled at a subconscious level. For example,
when we are running we are not generally aware of
individual placement of the feet on the ground, or of
co-ordination of limb activity. While a number of
different motor tasks are processed by the brain at
any one time, we only have a single dominant
‘stream’ of conscious thought. This may be related
to the idea that subconscious neural network activity
controlling all subconscious functions such as motor
activity occurs in a massively parallel manner, while
conscious perception is a completely linear occur-
rence, and at any timepoint we only have one con-
scious perception of reality. In the fatigue model
therefore, this serial mechanism may be responsible
for the dissociation between the conscious sensation
of fatigue and underlying subconscious changes in
pacing strategy.
4. Future Directions
In this article we have examined briefly both the
anatomical origin of the sensation of fatigue and a
© Adis Data Information BV 2003. All rights reserved.
173
Second order map 2
Awareness
of activity Awareness
of fatigue
Longer activity
Early activity
First order map First order map
possible mechanism of how it may originate from
subconscious processes in these anatomical areas or
functional processes, from a monistic perspective.
As described previously, neuroscience techniques
have been developed which allow us now to ex-
amine brain function in greater spatial and temporal
detail. We propose that future research should use
these techniques to examine, in detail, changes that
occur in different brain regions, or as generalised
alteration in brain activity, in order to elaborate on
the processes involved in the generation of the sen-
sation of fatigue and the ‘feeling of knowing’ that
we personally feel when fatigued. Fatigue as a
model may be relevant therefore to study the
broader problem of consciousness, because unlike
other emotions such as anger and fear which are
difficult to quantify, fatigue has generally been
shown to be a reproducible sensation in the laborato-
ry environment[71-73] which can be generated and
regulated in the human model using routine exercise
testing. Therefore, exercise physiology may have an
important role to play in answering questions that
Sports Med 2003; 33 (3)
174
have traditionally been the domain of philosophers
and neuroscientists.
5. Conclusion
In conclusion, we have examined fatigue as a
sensation rather than a physiological occurrence.
Fatigue is described as conscious awareness
originating from subconscious control structures. A
model for the development of the sensation of fa-
tigue is described, where the sensation of fatigue
originates from a difference between subconscious
representations of baseline physiological homeostat-
ic state and the state of physiological activity in-
duced by physical activity, which creates a second-
order representation which is perceived by con-
sciousness-producing structures as the sensation of
fatigue. In this model, fatigue has a teleological
function, and can be classified as an emotional con-
struct. It is not clear whether the origin of the fatigue
process is in a particular brain structure, is the result
of an integrative process involving a number of
different brain regions, or is the result of elec-
trophysiological synchronisation of entire brain ac-
tivity. Further work is required to determine from
which neural processes or brain structures the sensa-
tion of fatigue originates.
Acknowledgements
The Medical Research Council of South Africa, National
Research Foundation of South Africa, Technology and
Human Resources for Industry Programme of South Africa
and the Harry Crossley Research Funds of the University of
Cape Town provided financial assistance for studies de-
scribed in this review. The authors have no conflicts of
interest that are directly relevant to the content of this manu-
script.
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© Adis Data Information BV 2003. All rights reserved.
St Clair Gibson et al.
Correspondence and offprints: Alan St Clair Gibson, Re-
search Unit of Exercise Science and Sports Medicine, De-
partment of Human Biology, University of Cape Town,
SSISA, Newlands, 7725, South Africa.
E-mail: agibson@sports.uct.ac.za
Sports Med 2003; 33 (3)