Integrative and Comparative Biology

Integrative and Comparative Biology, volume 57, number 2, pp. 293–300

doi:10.1093/icb/icx069 Society for Integrative and Comparative Biology

SYMPOSIUM

Physiological and Biomechanical Mechanisms of Distance Specific

Human Running Performance
M. A. Thompson1

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Exercise Science Department, Fort Lewis College, 1000 Rim Drive, Durango, CO 81301, USA
From the symposium “The Ecology of Exercise: Mechanisms Underlying Individual Variation in Movement Behavior,
Activity or Performance” presented at the annual meeting of the Society for Integrative and Comparative Biology,
January 4–8, 2017 at New Orleans, Louisiana.

1
E-mail: [email protected]

Synopsis Running events range from 60-m sprints to ultra-marathons covering 100 miles or more, which presents
an interesting diversity in terms of the parameters for successful performance. Here, we review the physiological and
biomechanical variations underlying elite human running performance in sprint to ultramarathon distances.
Maximal running speeds observed in sprint disciplines are achieved by high vertical ground reaction forces applied
over short contact times. To create this high force output, sprint events rely heavily on anaerobic metabolism, as
well as a high number and large cross-sectional area of type II fibers in the leg muscles. Middle distance running
performance is characterized by intermediates of biomechanical and physiological parameters, with the possibility of
unique combinations of each leading to high-level performance. The relatively fast velocities in mid-distance events
require a high mechanical power output, though ground reaction forces are less than in sprinting. Elite mid-
distance runners exhibit local muscle adaptations that, along with a large anaerobic capacity, provide the ability
to generate a high power output. Aerobic capacity starts to become an important aspect of performance in middle
_ 2max is an important determinant of
distance events, especially as distance increases. In distance running events, VO
performance, but is relatively homogeneous in elite runners. VO _ 2 and velocity at lactate threshold have been shown
to be superior predictors of elite distance running performance. Ultramarathons are relatively new running events,
as such, less is known about physiological and biomechanical parameters that underlie ultra-marathon performance.
However, it is clear that performance in these events is related to aerobic capacity, fuel utilization, and fatigue
resistance.

Introduction runners who have achieved success in two different dis-

Running is one of the most popular sporting events tances. From a scientific perspective, this range of dis-
worldwide. According to Running USA’s 2015 annual tances presents an interesting diversity in terms of the
report 17,114,800 individuals participated in US sanc- parameters for successful performance.
tioned running events (Running USA 2016), while a For all running disciplines, performance is depen-
further 10.5 million runners participated in UK events dent on the time required to cover the event distance,
(Sports Marketing Survey’s Inc. 2015). The variety of which can also be expressed as the average velocity over
running related events contributes to this popularity, the event duration. Running velocity (v) is determined
with individuals being able to participate in disciplines by the ratio of metabolic power (Pmet) and energy cost
including track, road, trail, mountain, and ultra- of running (C) (di Prampero et al. 1986):
endurance running. Running events range from sprints Pmet
of 60 m to ultramarathons covering 100 miles or more. v¼ :
C
In contrast to other Olympic sports, such as swimming
and speed skating, success in multiple running events of Pmet is the sum of aerobic and anaerobic energy pro-
different distance is rare. There have been a scant few duction over the duration of the event, this

Advance Access publication August 2, 2017

Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology 2017.
This work is written by US Government employees and is in the public domain in the US.
294 M. A. Thompson

physiological variable varies substantially from sprint

events to ultra-marathons. Alternatively, C depends
on numerous factors such as acceleration, terrain,
wind speed, and fatigue. In particular several biome-
Fig. 1 Phases of the 100-m running event.
chanical factors have been shown to affect C. Thus,
this equation represents the importance of the inter-
play between biomechanical and physiological factors
and the effect on running performance.
Here, we review the physiological and biomechan-
ical variations underlying elite human running per-
formance in sprint to ultramarathon distances. For

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categorization we grouped distances based on the
International Association of Athletics Federations
(IAAF) defined disciplines of sprint (60–400 m),
Fig. 2 Average velocity for world record sprint performances.
middle distance (800–3000 m), distance (5000 m-
marathon), and ultramarathon (>marathon). It is
important to note that in addition to the varying considerable variation in the deceleration phase
distances there are terrain differences associated (Mero et al. 1992).
with several of these events (e.g., track vs. trail), In the shortest sprint event, the 60 m, athletes accel-
which are beyond the scope of this review. erate for nearly the entire event. The significance of the
acceleration phase of a sprint is seen by the fact that
Sprint average world record speeds are nearly identical for
the 100 and 200 m events and less for the 60 m, whereas
The IAAF defines sprint disciplines as events up to average world record velocity is also lower in the 400 m
400 m with the Olympic distances being the 100, due to anaerobic limits (Fig. 2).
200, and 400 m. The duration of sprint events is Neural-motor integration is clearly one aspect that is
under a minute in length, with the marquee sprint important in this acceleration phase, as runners must
event, the 100 m, lasting approximately 10 s. Sprint sense and respond to the starting signal as rapidly as
events have received considerable attention in the possible. Research regarding reaction times during the
literature as posing an extreme of human perfor- start of sprinting events has revealed that in all sprint
mance. It should be noted that, though generally events the reaction times of the best athletes are
similar, within the range of 60–400 m there can be <200 ms, but reaction time does not correlate with per-
considerable differences in biomechanical and phys- formance level (Mero et al. 1992). Thus, while rapid
iological factors that underlie successful perfor- reaction time is certainly an essential component of
mance. For example you may see elite runners successful sprint performance, it is rather homogenous
perform well in the 100 and 200 m, or 200 and among elite runners. In terms of differentiating perfor-
400 m, but in the modern era we have not seen an mance, measurement of force production in the blocks
elite runner with high-level performance in both the indicates that elite sprinters produce greater forces and
100 and 400 m. have greater block velocity (the horizontal velocity of
the center of gravity of a sprinter during the last contact
Biomechanical moment in the blocks) than the less skilled sprinters
A unique factor of sprint events is that due to the (Mero et al. 1983; Hunter et al. 2005; Rabita et al.
short duration the acceleration phase is an important 2015). Additionally, research suggests that step length
component. Specifically, performance in the shorter is a main feature that determines speed in acceleration
sprint events, particularly the 60 and 100 m, is highly phase of the 100 m race. In a unique study examining
dependent on the ability to rapidly accelerate, Usain Bolt’s performance at the 2008 Beijing Olympics,
whereas at longer distances the start has considerably 2009 Berlin World Championships and in 2012 London
less relevance (Moravec et al. 1988). Over the course Olympics Games, Krzysztof and Mero (2013) found
of a 100-m run, the acceleration phase constitutes that the main distinguishing feature was step length in
the first 40–60 m, and then athletes generally main- the first 10–20 m.
tain speed for the next 10–30 m, before typically It has been established that primary kinetic factors
decelerating over the final 10–20 m (Volkov and that allow for the fastest sprint speeds are high ver-
Lapin 1979; Mero and Komi 1985; Moravec et al. tical ground reaction forces (GRFs) applied over
1988) (Fig. 1). However, sprinters exhibit short periods (Weyand et al. 2000, 2010).
Distance specific running performance 295

Competitive sprinters exhibit higher vertical GRFs The ability to generate force during sprint running is
and shorter contact times than non-sprint athletes likely dependent on leg strength, power, and stiffness, as
and this finding remains not only at the maximum these factors have been shown to be correlated with
speeds of these athletes, but even over a range of sprinting performance (Chelly and Denis 2001; Bret
speeds (Clark and Weyand 1985). Moreover, force et al. 2002). The importance of leg strength is further
application technique, specifically the orientation of illustrated by the finding that age-related muscular at-
the GRF vector, is important for performance rophy and losses in muscle strength are associated with
(Morin and Sève 2011). Additionally, the horizontal longer contact times, lower GRFs, and corresponding
component of the GRF appears to be a determinant decreases in running speed (Korhonen et al. 2009).
of sprinting performance, as the horizontal propul- Skeletal muscle characteristics underlie the force de-
sive impulse has been reported to account for 57% mands with critical determinants of sprint performance

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of variance in sprint velocity (Hunter et al. 2005) consisting of muscle mass, fiber type composition, and
and net horizontal force was reported to correlate fascicle length (Costill et al. 1976; Mero et al. 1981;
highly with sprint performance (Morin et al. 2011a). Kumagai et al. 2000). The high mechanical output re-
The need to generate high forces in a small amount of quired for elite sprint running performance is achieved
time indicates that the force-velocity relationship is the by a large number and cross-sectional area of type II
most important contractile property of muscle in terms fibers in the leg extensor muscles (Mero et al. 1981,
of limits to maximum sprinting speed. Greater stride 1983), which has recently been associated with the
frequencies require the legs to move through the stride ACTN3 R577R and ACE I/D gene variants
cycle at faster rates and the muscles must shorten and (Papadimitriou et al. 2016). Additionally, elastic energy
lengthen more rapidly, hence the force–velocity rela- storage and return is important for sprinting perfor-
tionship of skeletal muscle is a primary limiter of max- mance (Cavagna et al. 1971; Alexander 1991).
imum sprinting speed (Miller et al. 2012). In particular,
the gastrocnemius and soleus muscles are largely re- Middle distance
sponsible for the vertical GRF, and from a force–veloc-
ity perspective, shorter ground contact times mean that Middle distance events consist of the 800, 1500, and
3000 m, though there is some controversy regarding if
these muscles must contract with increased shortening
velocities, resulting in decreased peak forces (Schache the 3000 m should be considered as a distance or
middle-distance event. The duration of middle distance
et al. 2014). While the ability to generate these high
forces in rapid amounts of time certainly dictates run- events ranges from slightly <2 min to 8 min. Middle
distance running performance is an area that has re-
ning speed, the mechanics of the swing phase are also
important as rapid repositioning of the legs is required ceived limited attention in the scientific literature, per-
haps due to the variety of factors that are related to
(Weyand et al. 2000). Anthropometrically, sprinters
generally have long legs and relatively small calf circum- performance over this range of distances.
Performance in middle distance events is unique in
ference, which is thought to be associated with de-
creased inertia associated with swinging the leg that it is characterized by intermediate factors of bio-
mechanics and physiology, with the challenge being to
(Rahmani et al. 2004; Vucetic et al. 2008).
run at high velocities while still maintaining economical
movement. Middle distance events possibly create a
Physiological realm where different intermediates in terms of biome-
chanics and physiology may be associated with high-
While sprinting speed is clearly dependent upon the
level performance. The relative importance of mechan-
mechanical ability to apply high forces in short con-
ical power output and energy use likely varies across
tact times, several physiological factors underlie this
different middle distance events, as well as over the
ability to generate force during sprint running.
course of specific middle distance events. For example,
Sprint events rely heavily on anaerobic metabolism
power output likely plays a more dominant role in the
to support the high force output. The shortest of
800 m than in the 3000 m, and it is suggested that early
sprint events (duration <15 s) rely predominantly
in the 800 m race mean power output plays a predom-
on the ATP-PCr system to supply the ATP needed
for muscle contraction, whereas anaerobic glycolysis inant role, whereas energy use is more important later in
the race (Kadono et al. 2007).
provides a greater percentage of the required ATP as
distance increases (Cheetham et al. 1986; di
Prampero et al. 2015). Thus, individuals with higher Biomechanical
levels of anaerobic functioning will likely have better In terms of the biomechanical factors associated with
sprint running performance. high-level middle distance performance, the relatively
296 M. A. Thompson

fast velocities require a high mechanical power out- accumulation, have been found to have a significant
put, though GRFs are less than what is seen in relationship with middle distance running perfor-
sprinting (Nilsson and Thorstensson 1989). mance (Yoshida et al. 1990).
Mechanically we see that middle-distance runners
run in a similar manner to sprinters, but differ Distance
from distance runners at both maximal and submax-
Similar to sprint running, distance running has re-
imal velocities. The differences between middle dis-
ceived considerable attention in the scientific litera-
tance and distance runners include increased stride
ture. Distance events consist of the 5000 m, 10,000
length, reduced contact time, increased knee flexion
m, and marathon distances, although some consider
during swing, and greater center of mass oscillation
the 3000 m to be a distance event. The duration of
(Cunningham et al. 2013). Changes in running me-

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these events for elite runners ranges from slightly
chanics have been noted to occur during longer mid-
over 12 min to over 2 h.
dle distance events, such as the 3000 m, with
decreases in stride length, flatter foot placement,
Biomechanical
and forward trunk placement observed toward the
end of longer middle distance events (Elliott and While aerobic capacity and in particular running econ-
Roberts 1980). omy (to be discussed subsequently) are essential factors
for distance running performance, an obvious link ex-
ists between distance running biomechanics and econ-
Physiological omy. It is evident that economical running stems from
Elite mid-distance runners exhibit local muscle ad- executing optimal mechanical patterns that consist of
aptations (e.g., developed capillary network and an applying forces of appropriate magnitude, direction,
increased metabolic potential of muscle cells), which, and timing, without nonproductive movements. In
along with a high anaerobic capacity, generate the terms of distance running biomechanics there are
high mechanical power required for these events. both kinematic and kinetic variables that have been
Mechanical power output is critical for the fast ve- identified to influence running economy (Williams
locities observed in middle distance events (di and Cavanagh 1987). In regard to kinematics, stride
Prampero et al. 1993). The importance of velocity length has been identified as having a significant effect
can be seen in the fact that velocity at anaerobic on running economy, with most individuals self-
threshold, ventilatory threshold, and VO _ 2max have selecting a stride length that is optimal in terms of run-
been identified as a predictors of middle-distance ning economy (Moore 2016). As individuals deviate
running performance in novice and elite athletes from this point, with either shorter of longer stride
(Maffulli et al. 1991; Zacharogiannis and Farrally lengths, running economy worsens (Hogberg 1952;
1993; Abe et al. 1998). The relatively great muscle Knuttgen 1961; Cavanagh and Williams 1982; Heinert
force requirements of middle distance events are et al. 1988). As far as other kinematic varibles, to date an
considerably higher than the capacity of the athlete understanding of economical distance running perfor-
to provide using purely aerobic metabolism; thus, mance remains elusive. In terms of kinetics, it has been
there is a strong anaerobic component particularly shown that impact peak magnitudes, which are associ-
in the 800 and 1500 m events (Lacour et al. 1990b; ated with initial ground contact, and decreased vertical
Yoshida et al. 1990; Deason et al. 1991; Brandon impulses are associated with improved running econ-
1995). Whereas sprint events rely primarily on an- omy (Williams and Cavanagh 1987; Heise and Martin
aerobic metabolism, in middle distance events there 2001). Lastly, high total vertical impulse and net vertical
is an increasing demand for aerobic metabolism. The impulse (Heise and Martin 2001), and increased
maximal velocity that can be sustained under aerobic anterio-posterior braking force (Kyröl€ainen et al.
conditions has been identified as being significantly 2001) have been associated with poor running
correlated with middle distance running performance economy.
(Lacour et al. 1990a). Further, elite middle distance
runners have been found to exhibit high VO_ 2max and Physiological
_VO2max has been found to be more highly correlated In general, the primary determinants of distance
with running performance in middle distance run- running performance are strongly related to aerobic
ners when compared with distance runners (Foster capacity. In distance events maximum oxygen uptake
et al. 1978; Boileau et al. 1982; Camus 1992). _ 2max) is an important determinant of perfor-
(VO
Additionally, blood lactate variables, specifically lac- mance, with elite distance runners exhibiting high
tate threshold (LT) and onset of blood lactate _ 2max values (Foster 1983; Noakes et al. 1990;
VO
Distance specific running performance 297

Billat et al. 2001). Factors that contribute to the high are relatively new to the spectrum of running events,
_ 2max values observed in elite distance runners in-
VO given this and the relatively small numbers of partici-
clude increases in blood volume, capillary density, pants, less is known about the biomechanical and phys-
and mitochondrial density, with a primary factor be- iological parameters that underlie ultramarathon
ing increased stroke volume (Kanstrup and Ekblom performance. However, it is clear that these events
1984; Krip et al. 1997; Martino et al. 2002). rely heavily on aerobic functioning, fuel utilization,
However, VO _ 2max has been found to be relatively and fatigue resistance.
homogeneous in elite runners, and within elite pop-
ulations race times have been shown to have only a Biomechanical
low to moderate correlation with VO _ 2max.
There is a scarcity of research examining the biome-
Alternatively, VO_ 2 at LT and velocity at LT have
chanics of ultramarathon runners. Given the close rela-

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been shown to be superior predictors of distance tion with distance running it can be assumed that the
running performance (Farrell et al. 1979; Allen principles related to distance running biomechanics
et al. 1985; Bird et al. 2003; McLaughlin et al. 2010). also apply to ultramarathon performance. Specifically,
Running economy, which is defined as oxygen up- it can be assumed that the selection of economical stride
take at a submaximal running velocity, has been shown length, as well as lower impact peak magnitudes and
to be one of the most important determinants of dis- decreased vertical impulse are related to ultramarathon
tance running performance (Conley and Krahenbuhl performance. However, it has yet to be determined if
1980; Daniels and Daniels 1992; Roecker et al. 1998). distance and ultramarathon runners run in mechani-
Running economy has been shown to correlate highly cally similar manners. What research has been con-
with and be a predictor of distance running perfor- ducted regarding the biomechanics of ultramarathon
mance (Morgan and Pollock 1977; di Prampero et al. running indicates that runners alter their gait, perhaps
1986; Weston et al. 2000). Athletes with good running in an effort to adapt to fatigue induced changes or to
economy use less oxygen than athletes with poor run- avoid injury. These gait changes consist of a higher
ning economy at the same steady state velocity stride frequency and duty factor, and a reduced aerial
(Saunders et al. 2004). Economy can vary by as much time without a change in contact time (Morin et al.
as 30% in athletes with similar maximal oxygen uptake 2011b). Additionally, ultramarathon runners exhibit
values, which may explain performance differences lower maximal vertical GRFs and loading rate at impact
(Daniels 1985; Heise and Martin 2001). This variation (Morin et al. 2011b). Further, it has been noted that
in running economy is likely due to storage and return ultramarathon runners exhibit considerable gait vari-
of elastic energy in tendon. It has been determined that ability, which is likely dependent on terrain, but further
the amount of energy stored and returned in a tendon research in this area is needed (Giandolini et al. 2015).
during a given movement depends more primarily on
moment arm of the tendon, with the amount of stored Physiological
energy increasing as the size of the moment arm de-
Similar to distance running, running economy is also
creases (Carrier et al. 1994). In distance running there
a determinant of ultramarathon performance.
is a strong correlation between the moment arm of the
Specific variables that have been identified as predic-
Achilles tendon and running economy (Scholz et al. _ 2max,
tors of ultramarathon performance include VO
2008).
fractional utilization, and mean energy cost of run-
ning (Lazzer et al. 2012), as well as peak treadmill
Ultramarathon velocity during maximal exercise testing and velocity
at lactate turn point (Noakes et al. 1990). Given the
Ultramarathon events, also known as ultra distance or
extreme duration or ultramarathon events, fatigue
ultra running events, consist of any event over the mar-
resistance is clearly an important determinant of per-
athon distance. There are a wide variety of ultramara-
formance. Millet (2011) developed what he termed
thon events ranging from 50 and 100 km track races to
the flush model, which explains the relationship be-
multi-day cross country events. At the short duration
tween neuromuscular fatigue, running strategies, and
end of the ultramarathon spectrum are 50 km events
environmental factors and how these factors interact
that are just slightly longer than marathons with elite
to affect ultramarathon performance.
times around 3 h, whereas continuous events may last
48 h or longer and there are many multi-day events.
Additionally, ultramarathon events may be conducted Other factors
over specified distances or for specified times, for ex- While this review has focused on the biomechanical
ample distance covered in 24 h. Ultramarathon events and physiological factors that underlie human
298 M. A. Thompson

running performances at different distances, there aimed at further understanding these factors. Lastly,
are several other factors that affect performance ei- thus far it is understood that physiologically ultra-
ther on their own or via physiology and biomechan- marathon performance is dependent on high run-
ics. These factors may vary considerably across ning economy and fatigue resistance. The
individuals and/or disciplines. A primary factor biomechanical factors of increased stride frequency
that influences running performance is injury avoid- while maintaining contact times, and lower maximal
ance. Running has one of the highest rates of inju- vertical GRFs and loading rate at impact are poten-
ries, with the number of injuries ranging from 2.5 to tially related to the fatigue resistance that is associ-
33.0 per 1000 h of running (Videbæk et al. 2015). ated with successful ultramarathon performance.
Many elite runners have had careers interrupted or Expanding research in this area will further elucidate
shortened by overuse injuries. Additionally, particu- the physiological and biomechanical variables related

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larly in longer events, strategies such as pacing and to successful performance.
drafting are important performance factors
(Hoogkamer et al. 2017). Footwear and track sur-
faces have also been identified as factors that can
affect performance (Franz et al. 2012). While track
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