Scand J Med Sci Sports 2015: 25 (Suppl. 4): 110118
doi: 10.1111/sms.12573
Review
The Kenyan runners
H. B. Larsen1, A. W. Sheel2
1
The Copenhagen Muscle Research Centre, Rigshospitalet, Copenhagen, Denmark, 2School of Kinesiology, University of British
Columbia, Vancouver, BC, Canada
Corresponding author: Henrik B. Larsen, The Copenhagen Muscle Research Centre, Rigshospitalet, Copenhagen, Denmark.
Tel.: +4553744324, E-mail: hl@boag.nu
Accepted for publication 26 August 2015
Today the Kenyan dominance in middle- and long-distance
running is so profound that it has no equivalence to any
other sport in the world. Critical physiological factors for
performance in running include maximal oxygen
consumption (VO2max), fractional VO2max utilization and
running economy (energetic cost of running). Kenyan and
non-Kenyan elite runners seem to be able to reach very
high, but similar maximal oxygen uptake levels just as
there is some indication that untrained Kenyans and non-
Kenyans have a similar VO2max. In addition, the fractional
utilization of VO2max seems to be very high but similar in
Kenyan and European runners. Similarly, no differences in
the proportion of slow muscle fibers have been observed
when comparing Kenyan elite runners with their Caucasian
During the past four decades, African runners or
runners of African ancestry have produced some of
the most remarkable results in athletic events at
world-class level. While the West Africans have
excelled in short distance races (100400 m), the East
Africans, especially Kenyans, have excelled in mid-
dle-distance (8001500 m) and steeplechase, and
also, together with the Ethiopians, in long-distance
races (5000 m-marathon). The dominance of Afri-
cans has been more and more profound during these
four decades. During the same time period, the dom-
inance of European runners has been reduced dra-
matically. Thus, 38 years ago, all distances from
800 m to marathon were dominated by Europeans
(Matthews, 1987). The average proportion of Euro-
pean achievements in the six all-time top 20 lists in
the distances from 800 m to marathon including
steeplechase was 48.3% while the percentage of Afri-
can results was 26.6% of which the Kenyans pro-
duced 13.3%. Moreover, the majority of world
record holders were Europeans, and European gold
medal winners at the Olympic games and world
championships were not a rarity. Today, the propor-
tion of European achievements is reduced to 4.2%,
whereas the percentage of Africans in the top has
110
2015 John Wiley & Sons A/S.
Published by John Wiley & Sons Ltd
counterparts. In contrast, the oxygen cost of running at a
given running velocity has been found to be lower in
Kenyan elite runners relative to other elite runners and
there is some indication that this is due to differences in
body dimensions. Pulmonary system limitations have been
observed in Kenyan runners in the form of exercise-induced
arterial hypoxemia, expiratory flow limitation, and high
levels of respiratory muscle work. It appears that Kenyan
runners do not possess a pulmonary system that confers a
physiological advantage. Additional studies on truly
elite Kenyan runners are necessary to understand the
underlying physiology which permits extraordinary
running performances.
increased to 91.7%, of which 53.3% are Kenyans
(IAAF, All Time Outdoor Lists, June 2015, Fig. 1).
The markedly reduced occurrence of Europeans in
the list is not due to them necessarily running slower
now than earlier. It is simply related to the fact that
runners from East Africa in particular now run so
much faster (IAAF, 2015; Matthews, 1987). The per-
formances of Kenyan men in middle- and long-
distance events at the Olympic games, world
championships on track and world cross-country
championships underscore the Kenyan superiority.
Most of the top Kenyan performers come from a
group of eight small tribes called Kalenjin, which
number only approximately five million people.
Among the Kalenjin tribes, the Nandis have per-
formed the best although they constitute only
approximately 2% of the general Kenyan popula-
tion. The question that arises is why do Kenyan run-
ners perform so extraordinarily well in middle- and
long-distance races? This question has inspired con-
siderable interest, speculation and debate among ath-
letes, coaches, exercise physiologists and academics.
Those interested in understanding the dominance of
these athletes have wondered whether the secret
can be found in the socialization experiences and

All-time TOP 20 (%)
Asia
Kenya
53.3 South America
Africa excl. Kenya
North America
Europe
1.7
4.2
0.8 1.7
38.3
Fig. 1. Relative distribution (%) between the continents of
the all-time top 20 performances in middle- and long-dis-
tance running for men in the six major distances from
800 m to marathon incl. steeplechase in June, 2015. Black
Americans are regarded as Africans excl. Kenyans.
lifestyles of Kenyan children (e.g., the effects of run-
ning to school as children). Others point to the
impacts of living and growing up at high altitude,
and/or to the diet of these athletes. While it has been
suggested that Kenyans may benefit from biological
advantages over groups in other regions of the world
we recognize that this is controversial (Hoberman,
2004). Attempting to understand the intersecting
spheres of biology, anatomy, physiology, psychol-
ogy, sociology and how it is they explain possible
racial differences in athletic performance is con-
tentious among scholars of different disciplines [for
contrasting perspectives see: (Hoberman, 2004) and
(Pitsiladis et al., 2004)]. We are cognizant of this
controversy and have purposely narrowed the scope
of our manuscript to the physiology of Kenyan run-
ners. The study of elite athletes has long been used to
better understand human physiology and address
questions of O2 transport, muscle metabolism, car-
diorespiratory control and the neural control of
muscular contraction.
It is generally agreed upon that there are three bio-
logical prerequisites for a runner to be successful:
high capacity for aerobic energy output (VO2max),
the ability to use a high fraction of it for long periods
and a good running economy (i.e., low energy cost at
race pace throughout the race) (Joyner & Coyle,
2008) (Fig. 2). When studying athletic performance
and the physiology of running one should consider
each of the abovementioned factors. Our brief review
is concerned with the available evidence that con-
trasts Kenyan and non-Kenyan runners with respect
Kenyan runners
Maximal oxygen
uptake
Performance
Fractional Running
utilization economy
Fig. 2. Main physiological factors critical for distance run-
ning performance.
to VO2max, fractional utilization of VO2max and run-
ning economy. We also highlight recent findings,
which consider the pulmonary system as a potential
limitation to exercise performance.
Maximal oxygen uptake
Since the early work of A. V. Hill (Hill & Lupton,
1923), exercise physiologists have associated the lim-
its of human endurance with the ability to consume
oxygen at a high rate. Indeed, it is now well estab-
lished that there is a high correlation between of
VO2max and running performance in groups of run-
ners of quite different abilities. A high maximal O2
uptake during exercise represents the ability to inte-
grate the components of transport and utilization:
cardiac output, pulmonary gas exchange, hemoglo-
bin, blood flow, muscle O2 extraction, and the rate of
aerobic ATP resynthesis. Elite male athletes typically
have a VO2max between 7085 mL/kg/min with
women having values approximately 10% lower
owing to a lower hemoglobin concentration and
higher levels of body fat (Saltin & Astrand, 1967).
When groups of athletes with similar athletic perfor-
mance abilities or athletes with a relatively narrow
range of VO2max are studied VO2max becomes a less
sensitive predictor of performance (Sjodin & Sveden-
hag, 1985).
There have been several investigators that have
made direct VO2max comparisons between Kenyan
runners with runners of non-African descent (Saltin
et al., 1995b; Prommer et al., 2010; Tam et al., 2012)
while others have exclusively studied Kenyan ath-
letes and made comparisons with previously pub-
lished values (Billat et al., 2003; Foster et al., 2014;
Santos-Concejero et al., 2015). Saltin et al. (1995b)
revealed that Kenyan elite long-distance runners
studied at sea level have a very high VO2max
(79.9 mL/kg/min), but it was not higher than the
level observed in Scandinavian elite runners
(79.2 mL/kg/min). In addition, another group of
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Kenyan elite runners studied at altitude (2000 m)
had similar VO2max values when compared to Scan-
dinavian elite runners (66.3 vs 67.3 mL/kg/min).
Furthermore, three groups of junior runners studied
at altitude had VO2max values of 68.2, 65.5 and
58.7 mL/kg/min, respectively. However, as the entire
Kenyan group of elite runners encompassed six
Olympic or world champions, several silver and
bronze medalists at the world championships, one
world record holder and two junior runners who
later broke the senior world record the possibility
exists that these runners, when top-trained and stud-
ied at sea level, have values higher than the mean of
79.9 mL/kg/min. In fact, only two of the very best
Kenyan runners (top five in the world in their respec-
tive distances) were in the group of runners studied
at sea level. In addition, the results of Peter Koech
(world record holder at 3000 m steeplechase) studied
at altitude and John Ngugi (Olympic champion
in 5000 m and five times world champion in cross-
country running) (studied at sea level at submaximal
speeds) indicate that these runners would have
VO2max values close to 85 mL/kg/min. Thus, we feel
safe to conclude that Kenyan runners performing
well in long-distance running on the international
arena are characterized by VO2max values between 80
and 85 mL/kg/min although there are examples of
world-class runners like Julius Korir having 77 mL/
kg/min but still winning major races. This is consis-
tent with findings on two former Kenyan world
record holders, Kipchoge Kip Keino had a
VO2max of 84.8 mL/kg/min and Henry Rono had
84.3 mL/kg/min (Larsen, 2012). It is of note, that
these values are comparable to former non-Kenyan
world record holder Dave Bedford who had a
VO2max of 85.0 mL/kg/min and Steve Prefontaine
who had 84.4 mL/kg/min (Larsen, 2012).
It is not mandatory to have a VO2max above
80 mL/kg/min to be successful. Indeed, a study of
Kenyan marathon runners having an average best
performance time of 2:07:17 (h:min:s) has shown
VO2max values of 63.1 mL/kg/min when the athletes
were studied at altitude (2000 m) in Kenya and
67.5 mL/kg/min when studied in Italy (1300 m)
(Tam et al., 2012). In line with this, a study by
Prommer et al. (2010) has shown that Kenyan dis-
tance runners can run a 10 km in 28:29 (min:sec)
with a relatively low mean VO2max value of
71.5 mL/kg/min. In addition, several studies of
black South African elite distance runners have
shown that they can perform well without having a
very high VO2max (Bosch et al., 1990; Coetzer
et al., 1993; Weston et al., 2000). Moreover, two of
these studies demonstrated that Black South Afri-
can runners with the same race time as White
South African runners, have a lower VO2max than
the White runners. Thus, a relatively low VO2max of
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Black South African runners seems to be a com-
mon feature at a given performance level compared
to other runners in the world.
A key question is whether untrained Kenyans of
Kalenjin ancestry have a high VO2max at a young
age. To answer this question untrained adolescent
Nandi town boys were studied (Saltin et al., 1995b;
Larsen et al., 2005). When studied at an altitude of
~2000 m, Kenyan boys had VO2max values, which
were in the same range as untrained Caucasian teen-
agers studied at sea level (Andersen et al., 1987). It
can be argued that the maximal oxygen uptake of the
Kenyan boys would be higher if they were tested at
sea level. For example, VO2max is approximately
10% reduced when tested at 2000 m (Squires & Bus-
kirk, 1982). However, it appears that high-altitude
natives do not gain much in VO2max when tested at
sea level (Favier et al., 1995). Although there may be
a small difference, one also has to consider that the
body mass of the Nandi boys was only ~54 kg, which
is ~12 kg less than that of Caucasian boys of the
same age. If VO2max is normalized for differences in
body mass with the exponent of 0.75, the Kenyan
boys will actually have a lower VO2max than the Cau-
casian boys. The fact that no difference was observed
with respect to VO2max between untrained Kenyan
and Danish boys is in line with findings of others
(Boulay et al., 1988).
Fractional utilization of VO2max
The ability to sustain a high percentage of VO2max
during competition has long been identified as a pre-
dictor of endurance performance. Is the success of
Kenyan runners explained by a high fractional uti-
lization of VO2max? The answer to this question is
difficult as the number of data points in the literature
are few. It was shown in one runner, Kip Keino, that
he utilized his whole VO2max when running 5000 m
and above 9798% of VO2max in a 10-km race
(Karlsson et al., 1968). This original observation is
consistent with more recent findings where Kenyan
runners are able to sustain 9396% of the velocity
associated with VO2max at 10 000 m speed during
treadmill running (Billat et al., 2003). Moreover, in
the above-mentioned study by Tam et al. (2012) the
fractional utilization of VO2max were calculated and
compared with European athletes of almost the same
performance level (2:08:24 in the marathon). This
study indicated that the fractional utilization of
VO2max sustained during the marathon was extre-
mely high but similar in the Kenyan and European
runners. Finally, when the average heart rate during
a 5000 m competition was recorded in Nandi town
and village boys they utilized 97.3% of their maximal
heart rate (Larsen et al., 2005). Here, we present a
summary of published values in Kenyan runners that

are physiological indices of a high fractional utiliza-
tion of VO2max.
Muscle fiber type
The majority of factors that may explain superior-
ity in exercising at a high fractional utilization of
VO2max are related to the features of the muscles
involved in the action of running. There is a mod-
erate-to-strong relationship between distance run-
ning performance and the proportion of type I
muscle fibers (Costill et al., 1973; Sj
odin & Sveden-
hag, 1985) which also appears to be the case in
well-trained cyclists (Coyle et al., 1988). It has been
suggested that the percentage of type I muscle
fibers may be an indicator of the potential train-
ability of the musculature (Sj odin et al., 1982).
However, extreme endurance training has been
demonstrated to induce a similarly high mitochon-
drial oxidative capacity of type I and type II mus-
cle fibers, in line with the fact that the contractile
characteristics of a fiber and transformation of the
main fiber types are not so easily affected by train-
ing, whereas the metabolic capacity is; also in the
type II fibers (Jansson & Kaijser, 1977). Both Ken-
yan and Scandinavian elite runners have a high
proportion of type I muscle fibers (Saltin et al.,
1995a). Kenyan junior runners have a proportion
of type I muscle fibers that is high and not much
different from Kenyan elite runners (~70% type I
fibers). South African distance runners appear to
differ from Kenyan runners, having less type I
fibers (Coetzer et al., 1993). In sum, there is contra-
dictory evidence to suggest that Kenyan elite dis-
tance runners may have a higher proportion of
type I muscle fibers relative to other runners. These
conflicting results are difficult to resolve and we
emphasize that the number of muscle biopsies
obtained from Kenyan runners to date have been
remarkably few and additional study is required.
Leg muscle oxidative enzymes
Muscle mitochondrial enzyme capacity is linked with
the substrate metabolism during exercise. The more
mitochondria, the larger the muscle lipid consump-
tion during exercise and in turn, the level of blood
lactate at a given work rate is lower, the higher the
activity of the oxidative enzymes (Ivy et al., 1980).
Thus, a direct link exists to the oxidative potential of
the muscles engaged in running and the performance
level, as a running speed has to be high before blood
lactate starts to become elevated (Sj odin & Jacobs,
1981). There is a positive correlation between the
activity of three oxidative enzymes, citrate synthase
(CS), succinate dehydrogenase, and glycerol-3 phos-
phate dehydrogenase, and performance in running.
Kenyan runners
When comparing the CS activity of Kenyan and
Scandinavian elite runners, no differences were
found regardless of whether the comparisons were
made between the vastus lateralis or the gastrocne-
mius muscle in the two groups of runners (Saltin
et al., 1995a).
Capillaries
Capillarization is one of the key factors determining
the oxidative profile of the musculature and it is clo-
sely related to VO2max. Capillary density has also
been shown to correlate positively with the running
velocity at which blood lactate begins to accumulate
(Sj
odin & Jacobs, 1981). A moderate correlation is
found between muscle capillarity and the mean
marathon running velocity during competition
(Sj
odin & Jacobs, 1981). When comparing the capil-
larization of Kenyan vs Scandinavian elite runners
there appears to be only a tendency for a higher
capillarization of the Kenyan elite runners (Saltin
et al., 1995a).
Plasma lactate and ammonia responses
The blood lactate response to submaximal running is
a good predictor of endurance running performance
and primarily reflects the local metabolic response in
the running muscles. Kenyan elite runners have
lower blood lactate levels, both at altitude and at sea
level, when running at a given exercise intensity com-
pared to other elite runners. The difference is most
pronounced at high exercise intensities. With the
accumulation of lactate in the blood the ammonia
concentration usually also increases. This is also true
for Kenyan elite runners but only at very high exer-
cise intensities and then to a lower extent than other
elite runners (Saltin et al., 1995a). In addition, the
peak ammonia concentration following a maximal
test was only half to one third in the Kenyan runners
as compared to other elite runners. Following a per-
iod of endurance running, the blood ammonia level
was reduced in Kenyan town and village boys (Lar-
sen et al., 2005). These blood ammonia data warrant
further exploration, as they may explain some meta-
bolic regulations in the muscles of Nandi runners, in
turn affecting critical fatiguing factors, either periph-
erally in the muscle or at a more central level.
Running economy
Running economy is expressed as the steady state
submaximal oxygen uptake at a given running veloc-
ity. The lower the VO2 at a given submaximal run-
ning speed, the better the running economy. Studies
from the 1930s were the first to suggest differences in
the amount of oxygen that different athletes require
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Larsen & Sheel
when running at the same speed (Dill et al., 1930)
and the submaximal oxygen requirement (mL/kg/
min) has been shown to vary considerably between
subjects (Svedenhag & Sj odin, 1984). The oxygen
cost of running at a given running velocity normal-
ized for differences in body mass has been found to
be lower in Kenyan elite runners relative to other
elite runners (Saltin et al., 1995b). In addition, when
comparing some of the most efficient Kenyan run-
ners (Julius Korir and John Ngugi) without normal-
izing for differences in body mass these runners are
still superior compared to top-Swedish runners
(Saltin et al., 1995b). In fact, their running economy
is very comparable with the one found in two of the
most efficient Caucasian runners (Derek Clayton
and Frank Shorter) ever measured (Pollock, 1977).
Furthermore, their data are in line with or slightly
better compared to the best ever economy curves
(Joyner, 1991).
Other researchers have found no association
between running economy and running performance
in Kenyan runners which was interpreted to mean
that running economy is compensated for by other
factors to afford distance running success (Mooses
et al., 2015). It should be noted that others have cri-
tiqued this work [see (Santos-Concejero & Tucker,
2014)] and suggested that the importance of running
economy in Kenyan runners cannot necessarily be
downplayed.
Classical studies of human locomotion (Cavagna
et al., 1964) show that the work of moving the limbs
comprises a substantial part of the metabolic cost of
running, just as load-carrying experiments have
shown that adding a few grams of mass on the feet/
ankle evokes an increase in the metabolic rate
(Myers & Steudel, 1985). Some have speculated that
successful Kenyan runners are characterized as hav-
ing slender limbs with low masses allowing them to
run with a minimal energy used for swinging the
limbs. Evidence to support this postulate comes from
studies which show that elite Kenyan runners had
longer legs (approximately 5%) relative to Scandina-
vian runners and they had thinner and lighter calf
musculature and that Kenyan boys have a low body
mass and a very low BMI relative to boys of similar
age from other continents (Saltin et al., 1995b; Lar-
sen et al., 2004). If slimness and low body mass
indeed is important prerequisites for becoming a
world elite runner it is probably easier to produce
this kind of athletes in Kenya than most other places
in the world due to a much higher prevalence of peo-
ple having this body shape in Kenya.
With respect to body dimensions the study by
Tam et al. (2012) raises several important points.
The Kenyan and European marathon runners who
had a comparable running economy also had similar
body dimensions (height, weight, and BMI). In
114
contrast, the study by Saltin et al. (1995b) demon-
strated that the Scandinavian elite runners having
inferior running economy compared to Kenyan elite
runners were taller, heavier and had a higher BMI
compared to the Kenyans. This lends support to the
hypothesis that body dimensions are crucial for run-
ning economy. Tam et al. (2012) also noted that
Kenyan and European runners who performed
equally well in the marathon achieved the results
with a very similar combination of VO2max, frac-
tional utilization of VO2max and running economy.
Pulmonary system limitations to exercise
It is commonly held that the structural capacity of
the normal lung is over-built and exceeds the
demand for pulmonary gas transport in the healthy,
exercising human (Dempsey, 1986). However, the
adaptability of the pulmonary system structures to
habitual physical training is substantially less than
other links in the O2 transport system (Dempsey &
Wagner, 1999). Accordingly, in some highly fit indi-
viduals the lungs diffusion surface, airways, and/or
chest-wall musculature are underbuilt relative to
the demand for maximal O2 transport (Dempsey,
1986). Two specific pulmonary limitations to exercise
performance in the highly trained have been
reported: (a) exercise-induced arterial hypoxemia
(EIAH) secondary to excessive widening of the alve-
olar-to-arterial O2 difference, inadequate hyperventi-
lation and metabolic acidosis; and (b) highly
fatiguing levels of respiratory muscle work which
effectively steals blood flow from locomotor muscles
via a sympathetically mediated reflexes and heightens
perceptions of limb and respiratory discomfort.
While these limitation have been shown by several
research groups to occur in highly trained athletes, it
is unknown if Kenyan runners experience these pul-
monary limitations during exercise. Here, we briefly
summarize the available data, which speaks to pul-
monary system limitations in Kenyan runners.
Exercise-induced arterial hypoxemia
Arterial O2 desaturation of 315% below resting
levels has been observed to occur at or near maxi-
mum exercise intensities. EIAH has been reported in
highly trained young males, as sedentary males do
not experience EIAH. Of note is that the relationship
between VO2max and the severity of EIAH is modest.
We emphasize that the true prevalence of EIAH and
susceptibility remains unresolved. EIAH is usually
viewed as occurring only in very heavy or maximum
exercise, it is underappreciated that many of those
experiencing EIAH begin to show decrements of
PaO2 even in submaximal exercise. PaO2 is reduced
below resting levels due to a widened alveolar-to-

arterial difference in O2 (AaDO2) and/or insufficient
increase in alveolar PO2. HbO2 desaturation is fur-
ther exaggerated by a rightward shift of the dissocia-
tion curve due to metabolic acidosis and increased
temperature. The precise cause(s) of EIAH have not
been fully identified exercise-induced ventilation/per-
fusion inequality most likely contributes to a
widened AaDO2. During heavy exercise a diffusion
limitation and perhaps even stress-failure of the
alveolar-capillary interface remains plausible, but
evidence for their existence is indirect in exercising
humans. From an athletic performance perspective,
it has now been well established that naturally
occurring reductions in arterial oxygenation have a
significant effect on exercise performance.
Foster et al. (2014) observed significant gas
exchange impairments in Kenyan runners that would
be of sufficient magnitude to compromise O2 deliv-
ery. Young Kenyan runners (10M, 4F; 25 years)
were instrumented with a radial artery catheter and
esophageal temperature probe for determination of
blood gases and core temperature. Participants com-
peted in national and international running events
with a range of racing accomplishments. Several sub-
jects had principally competed at local, modest high
altitude (15002150 m) in Kenya. One female partic-
ipant was former medalist at the Jr. World Cross-
Country Running Championships, while two male
participants competed internationally at major
marathons. Subjects performed incremental tread-
mill running to exhaustion at 1545 m, which corre-
sponds to an FIO2 of approximately 0.172 at sea
level. Significant decreases in the PaO2 and oxyhe-
moglobin saturation (SaO2) and a widening of the
Kenyan runners
alveolar-to-arterial difference in O2 from rest to peak
exercise were observed (Fig. 3). At maximal exercise,
the reduced SaO2 was caused by a combination of
reduced PaO2 (~60%) along with acid shifts (~40%)
in arterial blood and increases in core temperature.
We emphasize that there were notable between-sub-
ject differences in gas exchange during submaximal
exercise where several subjects had reductions in
PaO2 during the initial treadmill stage, whereas
others maintained PaO2 values close to resting levels
until near-maximal exercise intensity.
Humans who are native to high altitude (at least 3
generations) appear to exhibit an enhanced lung
structure and function relative to their lowland coun-
terparts where pulmonary gas diffusion during exer-
cise appears enhanced (Dempsey et al., 1971).
Moreover, it has been consistently shown that mod-
erate high-altitude exposure accelerates developmen-
tal growth of the mammalian lung. However, to date
it is unknown if these purported altitude-induced
lung enhancements are present in elite Kenyan run-
ners of high-altitude ancestry.
High levels of respiratory muscle work
The ventilation associated with heavy exercise causes
significant increases in the total work of breathing
including inspiratory and expiratory muscle work as
well as the resistive and elastic work of breathing.
The elastic work of inspiration is particularly high if
expiratory flow limitation occurs and the subject is
made to breathe in a relatively hyperinflated state.
The metabolic and circulatory costs of the hyperven-
tilation of heavy exercise are substantial and have

(a) (b) (c)

Fig. 3. Arterial blood gas and gas exchange values in Kenyan runners at rest and incremental percentages of maximal work-
load. (a) PAO2, alveolar partial pressure for oxygen and PaO2, arterial partial pressure for oxygen. (b) SaO2, arterial oxygen
saturation. Values for SaO2 were determined in three ways; (1) direct measure from blood gas analyzer (closed circles); (2) cal-
culated based on current PaO2 applied to an ideal dissociation curve (pH = 7.40, temperature = 37 C, PaCO2 = 40 mmHg)
(open triangles); and (3) individual resting PaO2 applied to a curve shifted due to current pH, temperature and PaCO2 (open
diamonds); (c) A-aDO2, alveolar-to-arterial difference in oxygen and PaCO2, arterial partial pressure for carbon dioxide. *Sig-
nificantly different from rest (P < 0.05). From Foster et al. (2014).

115
Larsen & Sheel
been estimated to represent 10% of the VO2max in
the untrained and upwards of 1516% of VO2max
and of maximum cardiac output in highly trained
males (Sheel & Romer, 2012). Highly trained sub-
jects are capable of achieving very minute ventila-
tions but the maximum flow-volume envelope is
unchanged from that of the untrained subjects. As
such, the expiratory flow during normally occurring
tidal breathing intersects with the maximum flow-
volume envelope. End-expiratory lung volume subse-
quently forced upward in order to increase flow rate
further. However, this increase in flow is accom-
plished at the expense of lung hyperinflation and
high work of breathing. It is now known that the
high work of breathing during sustained exercise
>85% maximum results in diaphragmatic fatigue.
Exercise performance can be affected, whereby
the fatigued respiratory muscles trigger a metabore-
flex from, which in turn causes a decrease in O2
transport to locomotor muscles in response to a fati-
gue-induced increase in sympathetic vasoconstrictor
outflow.
Foster et al. (2014) found that during maximal
treadmill exercise most male and female Kenyan run-
ners demonstrated significant flow limitation where
the tidal breath intersected with the expiratory side
of the maximum flow-volume curve or impending
flow limitation where their tidal flow-volume loop
approached the maximal curve and developed its
characteristic shape, but did not fully intersect. Sub-
jects were instrumented with an esophageal balloon-
tipped catheter for the determination of the work of
breathing. Throughout exercise there was a progres-
sive and curvilinear increase in the work of breathing
which was commensurate with values seen in non-
Kenyan athletes.
It is unknown if the diaphragm of maximally exer-
cising Kenyan runners fatigues. However, many of
the predisposing factors for fatigue of the diaphragm
appear present including: expiratory flow limitation,
a high work of breathing and significant metabolic
acidosis. Furthermore, Kenyan runners appear to
demonstrate expiratory flow limitation and high
levels of respiratory muscle work that are consistent
with observations other aerobically trained athletes.
The flow limitation observed was of sufficient magni-
tude to restrain the increase in ventilation during
exercise. These findings suggest that pulmonary sys-
tem limitations are present in runners of Kenyan
ancestry. Based on the available evidence, it appears
that Kenyan runners do not possess a pulmonary
system that confers a physiological advantage.
Perspectives
Different geographical regions including Finland,
Australia and New Zealand have dominated interna-
116
tional running races over the last 100 years. Runners
from East Africa, and in particular Kenya, currently
dominate international competition. As argued by
Joyner and Coyle (2008) this geographical diversity
argues against a simple genetically-based set of
answers to the complex problem of elite endurance
performance. On the other hand, the above-men-
tioned countries have far from been as dominant as
the African countries are today. Until the legendary
Kipchoge Keino made his breakthrough in 1965,
the Kenyans did not run for competition. Keino did
not only show the world that Kenyans could run but
he also showed the Kenyans that they could run.
When studying the Kenyan phenomenon in
middle- and long-distance running there is no doubt
that the interactions between biology, motivational
and sociological factors must be considered. Much of
what we know about Kenyan runners, from a physi-
ological perspective, is based upon a limited number
of studies with small sample sizes and difficulties in
study-design. The pertinent question is; what is the
appropriate control group? In order to compare
Kenyan runners born/raised at modestly high alti-
tude the comparative group must be fully acclima-
tized across physiological systems (e.g., ventilatory,
hematological, metabolic and cardiovascular). We
stress that when it comes to conclusions between
Kenyan and non-Kenyan runners it must be consid-
ered that most studies may not have made the desired
comparison. For example, comparing top-level
European marathon runners with Kenyans with an
equivalent marathon time (e.g., 2:08:00) is difficult as
the Kenyan runners cannot be characterized as top-
level. The mean performance time in the two groups
of runners corresponds to number 16 in the all-time
best performance list of European marathon runners
(IAAF, June, 2015) but to number 162 among Ken-
yan marathon runners. Future studies need to con-
sider the above factors when attempting to
understand the current success of Kenyan runners.
Key words: Elite runner, oxygen consumption, pul-
monary system, running economy.
Acknowledgements
Twenty-five years ago, Bengt Saltin initiated the studies in
Kenya of the physiology of Kenyan runners. Being a very
strong runner himself, the physiology of running had for
many years been one of his main interests. Therefore, after
first the Ethiopians and as the Kenyans had shown a still
increasing dominance in middle- and long-distance running,
Bengt Saltin had a strong desire to study the underlying
physiological reasons for this. As he said: If you are
interested in running of course you would like to know
why the Kenyan runners are the best. There is only one
way to find out and that is to go there and study them.
Therefore, he took the initiative and established a small
laboratory in Kenya in the area from where the most

successful runners originates. Since then studies from differ-
ent research groups have been going on in Kenya. Without
his great enthusiasm and scientific curiosity these studies
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