exertion and pleasure from an evolutionary perspective
TRANSCRIPT
Exertion and Pleasure from an Evolutionary Perspective
Michel Cabanac, PhD
Our knowledge of the world, including ourselves, is filtered twice – once by the narrow
physical or chemical window of our sense, and again by the biological or cultural programming
our brains. Both filters are the result of evolution, that is they have been passed down to us
because they proved their worth in our ancestors. Such filtering might therefore affect the way
we sense and perceive our own bodies.
But, first what do words like” sense” and “perceive” mean? Sensation is the irruption into
consciousness of any nervous message carried to the brain by an afferent pathway. From it, the
brain creates a mental object that has four mental dimensions: Quality, or the kind of stimulus;
intensity, or how strong it is; hedonicity, or how useful or noxious it is and duration or how
long it lasts. Quality, intensity and duration are positive and multiplicative. Only hedonicity is
additive , and it can be positive, negative or nil. Such a definition of sensation is simple and has
two advantages:
1. It lumps all the different categories of sensations into one category , whereas classical
categorization would list many different sorts of sensation with different attributes.
2. It points to a fundamental unity of sensory input to the central nervous system.
More complex than sensation, perception is the simultaneous entry of several afferent messages
including those retrieved from memory into consciousness ( Cabanac,1995 ).
This book is about humans. Our species, however, evolved from earlier life forms. And
this chapter addresses the origin of conscious creatures, their behavior resulting only from
tropisms and reflexes. So when, in phylogeny, did consciousness emerge ? my research team has
sought the answer in the evolutionary origin of emotional responses.
When we gently handled mammals, birds ( Gallus domesticus [ Cabanac & Aizawa,
2000] ) , and reptiles ( lizard, tortoises [Cabanac &Bernieri, 2000 ]), their body temperature rose,
producing an emotional fever. This response was produced in reptiles only through behavioral
means and lacking altogether in amphibians ( Cabanac & Cabanac,2004 ) and fish. Gentle
handling also accelerated heart rate , another sign of emotion, in mammals, birds (Cabanac &
Aizawa, 2000), tortoise (Cabanac &Bernieri, 2000), and lizards, but not in frogs (Cabanac &
Cabanac,2000). Because emotional fever and emotional tachycardia exist in mammals, birds, and
reptiles, but not in amphibians, the mental experience of emotion may have emerged in
evolutionary lineage between amphibians and reptiles.
Just as ancient is “ hedonicity “ that is the capacity tobassociate different sensation with
pleasant or unpleasant responses. Mammals, birds, and reptiles learn to avoid the flavor of
a novel food when digestive illness follows ingestion. Such learning is absent from amphibians
( Paradis & Cabanac, 2004 ). Again, the transition from amphibians to reptiles seems to have
been a critical evolutionary threshold.
These results are consistent with the hypothesis that specific mental capacities emerged
with reptiles. According to Darwinian theory, sensory messages should have become conscious
because consciousness had proved useful to the organisms first acquiring it. To be useful, these
sensations had to describe the quality, the intensity, and above all the usefulness of
environtmental stimuli; therefore, it is likely that they were multidimensional from the outset, as
defined earlier. The very persistence of consciousness over such a long time span demonstrates
its formidable selective advantage. It provided the first sensorially conscious animals with a
decisions – making edge by optimizing their behavior and by feeing them from the need for an
infinitely complex network of hardwired reflexes.
It consciousness has a single phylogenetic origin , it is likely that conscious events all
share the same basic four – dimensional structure (Cabanac,1996). We may now turn to a
specific case: the self – optimizing nature of feelings aroused by muscular exertion
(Ulmer,1996). All that we have defined in the general case of sensation should also hold true
here.
SENSATIONS FROM MUSCULAR EXERTION
Borg has extensively studied the feeling aroused by muscular excertion and has proposed
a scale from 6 to 20 called “ rating of perceived exertion “ ( RPE ) ( Borg, 1962,1982 ). The
feelings measured by Borg’s RPE scale are global perceptions arising from various parts of the
body; they are not broken down into their various inputs. Where, then do they originate? During
exertion, nervous messages emerge into consciousness not only from the working muscles,
tendons, and joints but also from other loci in the body.
MUSCLES AND TENDONS
In the working apparatus itself , the muscles and tendons, local conditions are modified
by exertion. Conscious message are sent via mechanical (Bloomstrand&Essen-
Gustavsson,1987;Gandevia&McCloskey, 1977 ; Roland, 1975 ), chemical ( Starkie et.al., 1999 ),
and temperature signals ( Saltin,Gagge,&Stolwijk,1968 ) in relation to the degree of exertion by
the muscles. Sensation of muscular exertion thus originates peripherally ( Sanes &
Shadmehr,1995; Wade,2003 ).
Muscles temperature is an important signal of fatigue but, unexpectedly , not of heat.
Cooling significantly shortens the time to reach fatigue and more than halves the work capacity.
This is unexpected because cooler muscles require less effort, make less demand on reserves, and
create lower concentrations of waste products and by- products. It is not yet understood why
fatigue occurs at a particular point or why local cooling reduces work capacity ( Wade et al,
2000) .
HEART AND RESPIRATION
Many authors have shown that rates of perceived exertion and fatigue are independent of
peripheral stimuli. Perceived exertion correlates most highly with blood pressure and not at all
with electromyogram readings ( Kilbom et al, 1983 ). Knibestol and Valbo ( 1980 ) conclude that
the signal for perceived exertion is central, not peripheral. For Pandolf 9 1978 ), muscular fatigue
is fundamentally based on a central signal : Cardiorespiratory stress. Indeed, dyspnea provides a
conscious signal that reliably describes the underlying phsyological state ( Mahler & Horowitz,
1994 )
Although heart rate is critical to perceived exertion, it is not the only factor. Heart rate is
significantly raised by sleep deprivation even though perceived exertion remains unchanged
( Martin& Haney, 1982 ). The findings of Jackson and colleagues ( 1981 ) do not support a
central control model that links perceived exertion solely to cardiovascular stress.
OTHER INTERNAL INFLUENCES
If neither peripheral inputs nor cardiorespiratory signals can explain perceived exertion.
Other factors may be involved. An important one seems to be lactacidemia : When subjects
inhale air with less oxygen, lactacidemia rises with perceived exertion while endurance falls
( Hogan& Welch, 1984 ). Conversely, intravenous glucose perfusion lowers heart rate and
respiratory quotient while lowering perceived exertion ( Tabata& Kawakami, 1991 ).
Hyperthermia raises lacttacidemia and in turn logarithmically increases perceived exertion (Berg
et al., 1986 ), a result also obtained by Kozlowski and colleagues (1985).
Core body temperature is thus a limiting factor that may increase discomfort and limit
exertion . Temperature has a real but indirect influence. Hyperthermia does not seewm to affect
the activation pattern of the muscles. Rather, the linear correlation among core temperature ,
electroencephalographic recording (EEG), and perceived exertion indicates that changes in
cerebral activity may be associated with hyperthermia-induced fatigue during prolonged exercise
in hot environment ( Nybo&Nielsen,2001 ). A rise in ambient temperature increases oxygen
consumption foR a constant workload and raises the anaerobic fraction. Thus, heat stress cause
some blood flow to be dedicated to the thermolysis rather than to muscles ( Dimri et.al, 1980 ).
Finally, as may be expected, the sensation of exertion depends on the type of work
performed by the muscle. At the start of exercise, sensation is a function of the muscle’s
resistance to the force and is largely independent of whether the work is static or dynamic
( Cafarelli,1982 ). When the work is extended overtime, the sensation of the exertion is of course
a function of duration ; the relationship between workload and work duration is hyperbolic when
subjects maintain a steady sensation of exertion ( Cafarelli, Cain&Stevens, 1977 ). Different type
of work arouse different sensation and also modify the internal environment differently : for an
identical workload, light and prolonged aerobic pedaling or weightlifting modifies ventilator
flow, heart rate, and level of lactate, cortisol, insulin and blood glucose less than intense but
intermittent work ( VanHelder et.al., 1985 )
CONCLUSION
The conscious signal is neither purely muscular nor purely central. It is combination of
both ( Lollgen, Graham&Sjogaard, 1980; Robertson, 1982 ). There are sensation of the working
muscles and from the heart, yet perceived exertion is not a single sensation but rather an overall
perception, as defined earlier. It is worth nothing that the rating of perceived exertion , that is, a
mental signal, measures exercise intensity as least as well as heart rate, that is a Physiologycal
Index ( Eston, Davies,& Williams,1987; Ueda & Kurokawa, 1991 ). In numerous experimental
studies, the subjects’ rating of perceived exertion predicted their relative metabolic demand,
especially at higher workload ( Noble,1982 ). This mental signal reliably adapts behavior to
Phsyological capacity.
The Borg’s scale of overall perceived exertion effectively describes what is taking place
overall in the main of a person exerting effort. Is it possible to push this analysis further? By
dissociating hedonicity from overall perception, we may find a tool with which to analyze
perception. If the hedonic dimension of perception is what motivates and optimizes behavior this
will be especially obvious in the case of muscular exertion and the perceptions it arouses.
HEDONICITY OF MUSCULAR EXERTION
Humans, unlike animal subjects, can verbally describe their sensations. Human
experimentation thus enables us to explore the mental experience of muscular exertion and
especially to analyze the hedonic content of the perceptions it arouses. To this end, we recorded
human sensation in the chest (variable x ) and the lower limbs ( variable y ) and how human
perception combine in subjects simply walking on a treadmill with five slopes (x) and five
speeds (y) ( Cabanac, 1985 ). The subjects separately estimated displeasure for chest and leg
sensations. Figure 6.1 plots these ratings as isohedonic lines ( left box: displeasure in the lower
limbs; middle box: displeasure in the chest; right box: sum of the two ratings). These ratings
were then compared with actual behavior in other session when the subjects had to climb 300m
( 328 yd ) on the treadmill at varying speeds and slopes. In these new sessions, when a set speed
was imposed, the subject could adjust slope, and vice versa. The dots on figure 6.1 show the
actual behavior of one subject. The dots fall along the lines, though not generating them, and
were from different sessions. Behavior (dots ) was strikingly adapted to the sum of perceived
displeasure in the chest and in the lower limbs.
Thus, in the situation explored, chest versus lower limbs, behavior was repeatedly
consistent: in the bidimensional sensory situation imposed by the experimenters, the subjects
described maps of bidimensional pleasure in session investigating pleasure and tended to move
to areas of minimal displeasure on the maps in sessions investigating behavior.
Figure 6.2 compares the behavioral choice ( dots 0 at the end of a session with the
theoretical time ( Lines ) needed to climb 300 m ( 328 yd ) as directed, the subject manipulating
speed or slope. It can be seen that behavioral choice coincided with the 40 min isochronal curve;
that is the subject tended to walk with constant time for a constant work. He exercised at a
constant power for the various combinations of slopes and speeds. This makes sense from the
standpoint of physiology . the result confirm that perception of muscular exertion (figure 6.1 )
integrates all afferent sensory inputs in addition, behavior is not only motivated by the hedonic
dimension of perception, but also optimized through minimization of displeasure ( figure 6.2 ).
These conclusions were repeatedly borne out by the findings of other research teams.
When subjects performed a mechanical task, that is, pointing, at a target with their arm in
various more or less functional positions, their precision and efficacy were maximal when their
posture was most comfortable ( figure 6.3 ). Performance deteriorated with increasing discomfort
( Rossetti, Meckler, & Prablanc, 1994 ).
When subjects performed a short ( 10 min )incremental exercise on a electrically
stabilized exercise bicycle, their oxygen uptake, leg effort and dyspnea varied significantly with
different pedaling frequencies . Oxygen uptake was minimal at 60 rpm and increased at both
higher and lower pedaling frequencies. Both leg effort and dyspnea were minimal at 80 rpm; leg
effort intensified at higher and lower pedaling frequencies., and dyspnea was most intense at
100 rpm . thus, there was some conflict between minimization of energy expenditure and leg
effort at expenditures less than 180 W. leg effort was minimized at the cost of increased energy
expenditure ( Chen, Jones, and Killian,1999 ), but the general trend was similar to that of our
other experiment.
Pleasure aroused by muscular exertion decreased when the intensity of exertion reached
maximum aerobic power and bordered on anaerobiosis ( Acevedo et.al., 2003). The same
conclusion have also been reached by Ekkekakis, Hall, & Petruzzello ( 2005 ) : when exertion
became anaerobic, formerly pleasurable muscular effort now aroused displeasure. Oxygen
consumption tended spontaneously to be minimal when subjects could select their own pace
(Zarrugh, Todd, & Ralston,1974); indeed, that was what they did ( Zarrugh& Radcliffe, 1978 ).
The same type of of optimization also took place in swimming; the stroke frequency
spontaneously selected to achieve nmaximum speed was indeed the optimal one for oxygen
consumption ( Swaine& Reilly, 1983 )
When the intensity of treadmill work rose, the perception of activation also rose from 2 to
5 ( scale 1 to 6 )and shifted from agreeable to disagreeable, +3 to -1.5 ( scale -5 to +5 ). The
working muscles produced a clearly negative alliesthesia of -2 to -3 points. The same stimulus,
that is, muscular exertion , can thus arouse either pleasure or displeasure according to the
circumstances. When exertion is interrupted, the general perception becomes immediately
pleasurable. Thus, hedonicity is actually the means whereby the muscular system is optimized
and whereby sentient organisms become aware of challenges to homeostasis ( Hall, Ekkekakis,
& Petruzzello, 2002 ). This is the very definition of alliesthesia ( Cabanac, 1971 )
Exercise intensity beyond the point of transition from aerobic to aerobic to anaerobic
metabolism is accompanied by an exponential decline in affective valence. This change in affect
may be a useful guide in helping exercisers recognize their phase of metabolism and thus more
effectively self monitor and self regulate the intensity of their efforts ( Ekkekakis, hall, &
Petruzzello,2004 ). Ekkekakis distinguishes five types of hedonic responses to the muscular
exertion:
1. Brief episodes of pleasure, even at high intensity
2. Strong interindividual differences when exertion is prolonged
3. Universal clear-cut pleasure when exertion ends
4. Universal clear-cut displeasure when exertion borders on exhaustion
5. Universal clear-cut pleasure when intense exertion is interrupted ( Ekkekakis, 2003 )
When mediated by perceptions of exertion, the pursuit of pleasure optimizes
behavior , as assessed by physiological criteria. Pleasure indexes usefulness. Thus,
muscular exercise may be intrinsically motivated; that is, muscular exertion in itself
might be rewarding. Some daily walking does improve mood ( Thayer et.al, 2004 ).
Muscular exertion might also be rewarding because, though unpleasant, it procures
another rewards that offsets the unpleasantness of fatigue, for example, fighting the cold
by heating oneself through physical exercise.
If hedonicity is indeed the signal for behavior optimization, as shown in figure 6.2
and 6.3, it is worth nothing that constant muscular exertion was governed by thresholds
of negative hedonicity. Thus behavior was optimizes by minimizing bidimensional
displeasure. Such a result does not change the conclusions: Hedonicity is still the
optimizer. Yet one may wonder why our subjects would indulge in unpleasant behavior.
The answer is that they were motivated to do so. They hate the pleasure of being useful
participants in research and receiving a modest financial compensation. Whatever the
motivations, they were motivated to experience displeasure. The hedonicity of exertion
had to compete inside them with other hedonicities. We will now turn to the problem of
conflicting motivations.
HEDONICITY IN MOTIVATIONAL CONFLICTS
An organism must rank its priorities because behavior is a final common path: it is
not possible to dine and sleep at the same time. To compare motivations and rank them
in order of priority, one needs a common currency ( McFarlan& Sibly, 1975 ). I have
proposed elsewhere that pleasure is this common currency ( Cabanac, 1992 ). The
perception of pleaure, as measured operationally and quantitatively by behavioral choices
( in the case of animals ) or by ratings of the intensity of pleasure ( in the case of humans
), can serve as such a common currency for various motivations. Decisions would be
made simply through maximization of the sum of different hedonic values. Is this true in
the case of muscular exertion when fatigue is pitted against other motivation ?
THERMAL DISCOMFORT VERSUS FATIGUE
The hypothesis was verified in the treadmill versus ambient temperature experiments
in which a clear cost was involved, either fatigue or cold discomfort. In these
experiments, the subjects were placed in a bidimensional ( x,y ) sensory space and had to
make a tract – of. The result so that there behavior tended to place them in pleasurable
areas of this space. The perception of thermal environment was pitted again that of
walking on a treadmill ( Cabanac & LeeBlanc, 1983 ). Thermal comfort ( x ) could be
improved at the cost of fatigue ( y ). Dressed in swimsuits and tennis shoes, the subjects
walked at 3km/hr ( 1.9 mi/hr ) on a treadmill in a climatic chamber. In an initial series of
measurements, the treadmill ‘s slope was varied from 0% to 24%, and this conditions was
combined with an ambient temperature ranging from 25oC to 5oC in a 25-node matrix.
The subjects separately rated the pleasure or displeasure evoked by ambient temperature
and by exercise. Actual ratings of pleasure / displeasure of x and y where obtained an
ambient temperatures 5o,10o, 15o,20o and 25oC in combination with 0%,, 6%, 12%, 18%,
and 24% slopes. The ratings of x and y were totaled . figure 6.4 give the sum of the two
ratings and isohedonic lines interpolated between the nodes. The figure is a map of
pleasure in a bidimensional sensory space ( exertion vs. ambient temperature ).
In a second series of measurements, one variable was imposed, either treadmill slope
or ambient temperature , and the subject could manipulate the other one. The results are
also given on figure 6.4 as dots. The subjects reciprocally adjusted exertion intensity and
ambient temperature. When a steep slope was imposed, they selected a low ambient
temperature, and when walking on a level slope was imposed, they selected a lukewarm
ambient temperature. When the subjects were allowed to adjust treadmill slope, with
various ambient temperatures being imposed, they selected steep slopes at low ambient
temperature and zero slopes at high ambient temperature. Quite strikingly, the dots
showing the finally selected experimental conditions ( in quasi- steady states at the end
of 1 hr sessions ) are in the white areas indicating bidimensional pleasure. . operant
behavior was guided by a tendency to minimize displeasure ( or maximize pleasure ) in a
bidimensional space.
Heat production in working muscles may be estimated be three times the amount of
mechanical energy produced by the muscle due to the relatively low rentability of the
muscle engine. When the subjects walked on the treadmill , the heat production was
inversely proportional to the ambient temperature. Behavioral heat production was
proportional to the need to offset heat loss and was therefore optimal for temperature
regulation. Taken separately from verbal reports of pleasure / displeasure , these results
are consistent with animal observations ( Krebs& Davies, 1981) or experiments
( Collier& Rovee- Collier, 1981 ). Which show a good fit between animal behavior and
physiological need. This has been repeatedly demonstrated and needs no further
demonstration. Here, however I have gone beyond simple behavior measurement to
compare variations in behavior with variations in sensory pleasure, as judged from
ratings obtained in separate sessions. Behavior and pleasure follow the same patterns.
Thus, the tendency to maximize sensory pleasure serves the purpose of physiological
regulation not only in the perception of exercise but also in motivational conflict,. In both
cases, pleasure coincides with a clearly adaptive physiological aim. This strongly
suggests that pleasure is the key to optimal behavior and that maximizing pleasure leads
to optimal physiological performance.
If sensory pleasure is the common currency that mediates competing motivations for
behaviors with physiological outcomes. It may also mediate other, purely mental
motivations. Pleasure may be felt even when no physiological need is being addressed.
This was tested in a conflict between exertion and money.
STATIC EXERTION VERSUS MONEY
In this experiment, human volunteers, could earn money by simply exposing
themselves to unpleasant, painful sensations from isometric contractions in their thighs
( Cabanac, 1986 ). They sat with their backs against a wall and their legs at 90o angles,
without any chair or stool for support. The longer they remained and endured the pain,
the more money they earned,. In several sessions, the rate of pay was varied. It was found
that discomfort or pain, as rated by the subjects, increased linearly as a function of time.
The subjects also tolerated more intense pain for a longer time when the monetary reward
was higher (Figure 6.5). this finding is consistent with common sense. It can be assumed
that the subjects decided to end a session just when the displeasure of the sensation
became greater than the pleasure of the anticipated monetary reward. The relation
between reward and duration of tolerated displeasure was logarithmic. The hypothesis
may thus cover behavioral motivations other than simple sensory hedonicity. The pursuit
of pleasure may involve non physiological motivations.
The hedonic dimension of sensation seems to be common currency that mediates the
ranking of priorities. The sensations and perceptions aroused by muscular exertion are
among many that may be mediated ( Cabanac, 1986 )
ANIMAL STUDIES
This conclusion is supported by several animal experiments in which animals had to
work for their reward. The amount of work an animal did for an increasingly infrequent
reward was used to measure its rewarding effect. Three groups of rats were trained for
this experimental design, the rewards being food pellets containing 1%, 10%. And 95%
sucrose. When the food pellets were provided continuously, the 10% sucrose ones
maintained the higher work responses rates. When, however the food pellets were
provided less snd less often, performance was consistently related to sucrose
concentration ( Cheeta, Brooks, & Willner, 1995 ).
When fed a tasty diet, rats tend to ingest more. When quinine, a bitter substance, is
added to their food, they tend to ingest less and lose weight. In an experiment in which
rats had to work for their food, their body weight was the same as when quinine was
added to their food and indirect indication that work was aversive ( Peck, 1978 ). When
rats had to run on a treadmill for sweet drinks, drinking was related to speed and, to a
lesser degree to distance. Thus, the rats must have been comparing information from their
own bodies ( rate of energy expenditure ) with their use of a commodity ( sweet reward )
( Gannon, Smith, & Tierney, 1986 ). Other experiments pitted reproduction
(insemination, lactation, nursing ), with costs energy , against food intake, which was also
costly. As the rats had to earn food by working on a treadmill ( Perrigo, 1987 ).
These experiments provide convincing , albeit indirect, evidence that animals optimize
their behavior through hedonic signals. Is it possible to gain more direct evidence that
animals will seek sensory pleasure just for the sake of it and even will be ready to trade
off some displeasure of it ?
Light work must be pleasant to animals, as experiments show that rats will run
indefinitely on a treadmill if it is made available ( Jonsdottir et. Al, 1996 ) even when no
special reward is obtainable trough running. It may be, then, that work can be pleasurable
in itself. As intensity or duration increases, however, work becomes aversive. When
monkeys had to pull a chain for heat in a cold environment, they obviously linked the
muscular exertion to the warm reward : The increasing force requirement was met with
increasing tolerance for larger air temperature fluctuations and longer interval between
responses. Eventually they stopped and just sat and shivered ( Adair & Wright, 1976 )
In the obstruction method use by Warden ( 1931 ), rats could reach a bait at the cost of
stepping on an unpleasant electrified meshed floor. In a similar but more natural
situation, rats were trained to feed in a nest placed at one end of a 16 m (17.5 yd )zigzag
alley ( Cabanac & Johnson , 1983 ). The nest was provided with water and food in excess
of the rats’ needs. On the day of an experimental session, the nest was heated and tasty
food ( Shortcake, peanut butter ) was placed 16 m away, but the environment outside the
warm nest was cooled to -15oC, a temperature potentially lethal to rats. The rats run the
cold feeder for the palatable bait, not out of necessity ( chow was provided at no cost in
the warm nest ) but for the pleasure of ingesting the tasty food. The number of trips was
related to the tastiness of the food. When chow was placed in the cold feeder instead of
shortcake, the rats went to it once and did not return. The rats were quantitatively
comparing the pleasure of eating the food with the displeasure of enduring the cold.
Other mammals thus seem to behave as humans do in situations in which conflicting
motivations have to be resolved, with a central role for pleasure and hedonicity. If such a
mechanism exists in mammals, which we can more easily accept as having
consciousness, would it also exist in reptiles?
The role of sensory pleasure in decision making was verified in iguanas placed in a
motivational conflict. The iguanas had to leave a warm refuge, provided with standard
food, to reach a tasty but unnecessary bait ( lettuce ) in a cold environment. They
nonetheless ventured out into the cold, trading off the coldness of the outside
environment for the tastiness of the bait, but only in mild cold. When ambient
temperature was 0oC, they remained under the infrared lamp. Like humans, the iguanas
seemed to be minimizing their sensory pleasure; thus, they optimized their behavior
( Balasko & Cabanac, 1998 ).
CONCLUSIONS
Information on our physiological status is transmitted via signas to the human mind.
These signals are critical to survival and must be analyzed, prioritized, and adjudicated to
produce an appropriate behavioral response, such as during muscular exertion. This is the
role of hedonicity in sensation and perception : to select a behavioral response that is
appropriate to both the organism’s needs and its physiological capacities. Pleasure/
displeasure is the optimizer of behavior.
Hedonicity must have been an efficacious solution to the problems of life, as it has
been retained by successive reptilian and mammalian lineages down to us.because
behavior is a final common path, all motivations, including the autohedonicity of exertion
itself, compete to access for it. If a motivation to accomplish a given task is given so high
a priority that other hedonic messages are ignored, the resulting overexertion may lead to
injury or even death, as with Pheddipides , the famous marathon runner ( Cabanac &
Bonniot- Cabanac, 1997 ). In most cases however, the inconvenience of fatigue is simply
less than the rewards of work. Work is disliked but done nevertheless for the rewards that
flow from it. This was the very point of the verse in Genesis.
Conversely, acute and chronic physical activity influence brain function, resulting in
changes to brain morphology, neuronal firing rates, cellular metabolism, neurotransmitter
concentrations and release, number and sensitivity of receptors and level of gene
transcription and protein production ( Hoomissen, 2004 ). In the absence of activity,
boredom itselfbecomes hedonic motivation ( Mageau, Green-Demers & Pelletier, 2000 ).