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Routledge Handbook of Ergonomics in Sport and Exercise

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Neuromuscular Adaptations in Running

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William A. Braun
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3
NEUROMUSCULAR
ADAPTATIONS IN RUNNING
William A. Braun
SHIPPENSBURG UNIVERSITY, SHIPPENSBURG

Introduction
Run training, as with all forms of physical exercise training, stimulates the development of
adaptations that should benefit the performance of the task. In the case of distance running,
these adaptations often result in improved economy of transport (i.e. lower energy cost to
maintain a fixed running speed). Mechanisms behind training-related improvements in running
economy fall primarily into two broad categories: metabolic adaptation and neuromuscular
adaptation. The focus of this chapter will be on factors that may contribute to neuromuscular
adaptations and improvements in running economy primarily in distance runners. In conjunc-
tion with this, common training practices will be explored, including: (1) incorporation of
resistance training as a means of supporting the performance of running; (2) adaptations that
result from chiefly ‘run-only’ training; and (3) possible influences of barefoot running versus
shod running on neuromuscular adaptations that develop from running. The chapter begins
with a general exploration of the relationship between endurance run training and neuro-
muscular factors.

Neuromuscular influences of endurance run training

The specificity of training principle would suggest that slow, continuous aerobic exercise, which
is commonly performed by distance runners, will compromise high-velocity force production
and power. Endurance-only training also impedes gains in strength and can cause reductions in
power due to shifting of fibre-type characteristics towards Type I fibre-type and disuse atrophy
effects on Type IIa and IIx muscle fibre (Thayer et al., 2000). In addition, functional adaptations
that result from endurance training may be limited to metabolic outcomes (Cohen et al., 2010).
As a result, distance runners often incorporate interval-type training and/or strength training as
methods for supporting the power-based needs that are often important for success in their sport.
Plyometric training has also shown positive benefits for use in distance runners by supporting
improvements in running economy (Markovic and Mikulic, 2010; Saunders et al., 2006; Turner
et al., 2003). Neuromuscular adaptations that result from these types of training interventions
may positively influence motor unit activation and recruitment patterns and alter more commonly
assessed factors such as stride length and frequency. Bonacci et al. (2009) compiled a comprehen-
sive review of neuromuscular adaptations related to various types of training to which the reader
is referred. However, the focus of this chapter will remain on running adaptations.

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The quality of running economy may be related to existing (or changes to) elastic properties
of the leg muscles in addition to metabolic adaptation. For example, it has been shown that
the oxygen cost of running is inversely related to lower limb flexibility in elite marathon
runners (Jones, 2002). Those with greater leg stiffness (poorer flexibility) were found to be
more economical. Craib et al. (1996) made similar observations in a group of sub-elite runners,
hypothesizing that greater leg stiffness might confer an economy advantage by improving storage
and return of elastic energy generated during the eccentric phase of the stretch-shortening
cycle. These findings are also corroborated by Dalleau et al. (1998), who found better RE in
those runners who possessed greater leg stiffness. The investigators hypothesized that increased
leg stiffness develops as a neuromuscular adaptive response to endurance run training. However,
stiffness can be acutely affected as well. Stiffness may be increased transiently, leading to a lower
energy cost, by avoiding over-flexing the knees during running or by running on a less
compliant surface (Dalleau et al., 1998). An adaptive response to running that results in increased
leg stiffness may seem counterintuitive, however, as a more rigid muscle might be expected
to be more prone to injury resulting from repetitive impact.
Heise et al. (2008) found that muscle coactivation of biarticular muscles (rectus femoris and
gastrocnemius) during the stance phase leads to increased joint stiffness, which may allow for
more efficient use of stored elastic energy. They theorized that coactivation of these muscles
may be an adaptive response found in more economical runners. In a group of 16 female
runners, muscle coactivation duration was able to explain 45 per cent of the inter-individual
variability in running economy; an inverse relationship between running economy and duration
of muscle coactivation was noted (Heise et al., 2008). It was also theorized that EMG-based
biofeedback might be useful for educating runners about the coactivation process and, thereby,
driving higher economy that may result in performance gains.
Few studies have been conducted to explore neuromuscular adaptations that result specifically
from endurance training. However, Cohen et al. (2010) sought to assess whether such
adaptations might be observed in endurance trained adult males and boys. Different aged
populations were assessed to gauge whether age-related effects on neuromuscular adaptations
could be identified. Trained and untrained subjects from the two age cohorts were assessed
for isometric strength of elbow and knee extensors and flexors and EMG activity of the
associated agonist and antagonist muscle groups. While age-related and training-related effects
were noted for some measures (i.e. higher rate of muscle activation and rate of torque
development in adults), there was no clear evidence of neuromuscular adaptation when
comparing trained to untrained cohorts. As a result, the authors concluded that endurance
training elicits chiefly metabolic adaptations (Cohen et al., 2010).
Indices of neuromuscular adaptation in runners have been reported by other investigators.
For example, Paavolainen et al. (1999c) showed that 5 km race time was related to
neuromuscular characteristics in a homogenous group of elite male orienteers. Specifically, the
ability of runners to generate force rapidly during maximal and submaximal running was related
to race performance. Similarly, ground contact time was shorter in those with the better race
performance. Spending less time during ground contact, especially during the braking phase
of contact would, presumably, conserve energy. In support of this, shorter ground contact time
was also related to better RE (Paavolainen et al., 1999c). This tends to support the theory
proposed by Kram and Taylor (1990) that suggests the energy cost of running is a function of
mass and ground contact time. They noted an inverse relationship between the rate at which
energy is used during running and the amount of time the foot generates force against the
ground. Thus, those runners with better economy tend to also have shorter ground contact
time.

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Acute neuromuscular effects

Neuromuscular control appears to be modified during the course of sustained running.
Paavolainen et al. (1999b) found that braking and propulsion components of ground contact
times were longer immediately following completion of a 10 km time trial. In addition to
these changes, peak and vertical ground reaction forces were significantly reduced and there
was a reduction in muscle activation during a 20 m maximal sprint performed after the time
trial. The investigators hypothesized that repetitive stretch-shortening cycles reduced the ability
of the neuromuscular system to maintain force production and to tolerate impact forces. It was
suggested that a decline in muscle stiffness, coupled with a longer transition time between
braking and propulsion, led to less conservation of elastic energy. These effects were
corroborated by a reduction in IEMG activity during the maximal 20 m runs (Paavolainen
et al., 1999b). Thus, acute and high-repetition SSCs have been shown to transiently influence
neuromuscular control properties and force production, both of which are important factors
in RE. Those runners who were able to better maintain running performance under fatiguing
conditions were also found to have higher relative pre-activation and lower relative agonist
IEMG-activity during the propulsion phase than the lower caliber runners. This supplies further
evidence for the importance of neuromuscular control and conservation of elastic energy via
maintenance of muscle stiffness during distance running.
Neuromuscular characteristics were also attributed to success in 5 km race performance
(Paavolainen et al., 1999c). Specifically, the ability to produce force quickly was an important
factor in maximal and submaximal running. The faster runners tended to demonstrate shorter
ground contact time, braking phase time and propulsion phase time than their slower, matched
counterparts.

Running economy
Running economy (RE) is a term used to reflect steady-state oxygen uptake under specified
conditions (i.e. fixed submaximal treadmill speed at zero grade, controlled environment)
(McCann and Higginson, 2008). It has also been defined as the energy demand for a given
submaximal velocity (Saunders et al., 2004a). More economical runners will maintain steady-
state running at a lower oxygen uptake. Further, when RE is improved through training, the
effect is evidenced by a reduction in the oxygen uptake needed to sustain a fixed submaximal
running speed. Thus, an increase in RE is reflected by a reduction in total energy cost.
Numerous factors have been reported to influence RE. Saunders et al. (2004b) provided a
schematic that identifies 17 different variables that may impact RE. These variables can be
broken into five subsets that include: training, environment, physiology, biomechanics and
anthropometry. Much attention has been given to many of these subsets, but teasing apart
specific effects and the magnitude of such effects is a complex process. While scientists may
be keenly interested in specific slices of the overall outcome and strive to characterize
mechanisms for adaptation, trainers and the athletes themselves are more likely to view the
big picture outcomes that training provides (i.e. improved performance, better pace
maintenance, quality of training). Since RE has been reported to be a better predictor of race
performance than aerobic power among elite runners (Costill et al., 1973), it has garnered a
great deal of attention from the scientific community. In the following sections, consideration
will be given to a variety of training interventions that have been investigated as a means of
improving RE and, in some instances, endurance run performance.

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Run training-imposed changes in economy

Beyond neuromuscular and metabolic adaptations that result from run training, tissue-specific
changes have been reported as well. For example, Fletcher et al. (2010) sought to investigate
whether run training coupled with isometric training leads to increased triceps-surae tendon
stiffness, which is associated with improved running economy. These investigators imposed
isometric plantarflexion exercise (working at 80 per cent of MVC) 3 days per week for
8 weeks. Stiffness of the triceps-surae and MVC were assessed every 2 weeks, with RE mea-
sured at baseline and upon completion of the 8-week intervention. The group of highly trained
middle- and long-distance runners maintained their run training throughout the study. A non-
intervention control group of runners was used for comparison. While the training intervention
failed to significantly affect tendon stiffness and RE, a significant inverse relationship was found
between RE and stiffness of the triceps-surae tendon. This relationship occurred irrespective
of group assignment. The investigators observed that acute changes in tendon stiffness and RE
can occur and that these changes may happen in tandem. They suggest that less compliant
muscle would lower the energy transfer to the tendon, resulting in a lower cost of the muscle
activity. Conversely, a more compliant tendon would lead to more muscle fibre shortening
and/or an increased velocity of shortening for a given joint movement, which would raise the
cost of muscle activity. Whether an increase in tendon stiffness is a function of chronic
adaptation or acute high-intensity training is not evident. It is possible that isometric training
would be more likely to elicit positive effects on tendon stiffness in moderately trained runners
who are not as well adapted as the runners used in this study.
Brisswalter and Legros (1995) applied training overload (300 per cent volume increase over
2 weeks) on well-trained middle-distance runners to gauge impact on running economy and
running pattern. Two trends were found to develop: improved economy (~ 9 per cent) was
observed only at the running speed that corresponded with the volume overload (which
occurred at 80 per cent intensity); the change in economy was associated with individual stride-
rate adaptation although stride length was not altered. The improvement in economy was
attributed to an adaptation in foot pattern, as well as an improvement in foot stability at the
training pace.
Other differences have been reported in mechanical factors that are important to run
performance and economy based on sex and training emphasis (i.e. middle distance versus long
distance) (Chapman et al., 2012). Chapman et al. (2012) found that elite female runners showed
significantly shorter ground contact times when running at the same speeds as males. They
also had greater stride frequency and shorter stride length compared to male counterparts.
However, these differences appeared to be a function of height as when the data were
normalized for height, the differences disappeared. The only exception to this was with stride
frequency, which remained higher in women across all running speeds. When comparing
middle-distance to long-distance trained males, the rate of change in ground contact time
differed significantly as the running speed was increased. The long-distance runners showed a
significantly larger decline in contact time as running speed increased compared to the middle-
distance runners. This effect translated into a 31.1 per cent increase in metabolic cost for the
LD runners versus a 25.4 per cent increase for the MD runners when moving from 5 to 7
m.s–1. Based on these findings, the authors suggest that gait differences between distance
specialists may be a function of preferred and/or competition speeds and that the preferred
speed among LD runners could be driven by economy factors. For information related to
neural adaptations and sprint training, the reader is referred to a review by Ross et al. (2001).
For a more applied review of power training and sport, readers are referred to Young (2006).

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Resistance training: effects on run economy

Despite seemingly divergent outcomes based on the principle of specificity, resistance exercise
training is a commonly accepted practice among distance runners. Numerous benefits of
resistance training for runners have been identified, including better maintenance of form and
economy, improved leg power and speed (Paavolainen et al., 1999a), improved sprint ability
(Mikkola et al., 2011), attenuation of cardiovascular drift (Hayes et al., 2004), enhanced
neuromuscular characteristics (Mikkola et al., 2011) and reduced injury risk. Muscular strength
endurance has been linked to better fatigue resistance during distance running as well (Hayes
et al., 2011). It has also been suggested that development of power could be impeded as a
result of continuous focus on aerobic training in some athletes, which may adversely affect
neuromuscular adaptation as well as heighten the risk of overtraining (Elliott et al., 2007).
Tanaka and Swensen (1998) hypothesized that resistance trained distance runners may be
better able to sustain performance at fixed submaximal running speeds by either enabling force
contribution from each myofibre to be reduced or by requiring fewer myofibres to sustain the
work. It was further suggested that resistance training leads to a lesser need for contributions
from lower efficiency Type II fibres (Tanaka and Swensen, 1998). These benefits would be
achieved by improved quality of the Type I fibre content that may accompany resistance
training.
Taipale et al. (2010) examined the effects of differing strength training interventions on
measures related to distance running performance, including running economy and VO2max
velocity (vVO2max). Strength training was performed concurrently with normal run training
over a 28-week period in a population of recreational runners. They found that explosive
strength training and maximal strength training were more effective than circuit weight training
for improving RE and vVO2max. These benefits were associated with greater strength gains
and neuromuscular adaptations (measured via EMG activity of the vastus lateralis and medialis)
in the two groups.
Ferrauti et al. (2010) failed to identify improvements in RE or muscle coordination in
response to 8 weeks of concurrent run and resistance training in recreational marathon runners
despite significant gains in leg strength. Strength training was performed twice weekly and
focused on the trunk (for muscular endurance) and lower extremity (high intensity). The high-
intensity strength training of the leg muscles was designed to promote motor unit recruitment
pattern improvements while minimizing hypertrophic gain in an effort to improve RE. Ground
contact time revealed a significant interaction when compared to a run endurance group such
that contact time tended to increase in the concurrently trained group while tending to decrease
in the run-only group. Despite this interaction, RE remained the same. Stride length and stride
frequency, indices of muscle coordination, were unaffected by concurrent training. However,
stride length was significantly decreased in the run-only group. Thus, some of the alterations
may have been a function of changes to run volume or intensity that occurred in the subjects
as they were building towards a marathon competition. The investigators hypothesized that a
larger subject pool or longer study duration may have revealed more profound effects (Ferrauti
et al., 2010).
Paavolainen et al. (1999b) introduced a 9-week explosive strength training regimen on
top of normal endurance training in elite distance runners to study the effects on 5 km race
performance. Explosive training included plyometric-type jumping exercises, leg press and leg
extension/flexion exercises with low loads and maximal velocity repetitions, and sprint sets.
Lifting loads fell below 40 per cent of 1 RM. Significant improvements in 5 km race time
(3.1 per cent faster) and in RE were associated with reduced ground contact time during

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constant velocity running in the strength-trained group. This occurred despite no significant
effects on stride rate, stride length or ground reaction force during constant velocity running.
Strength training was associated with significant gains in maximal isometric force production
of the leg extensors and maximal anaerobic run test velocity (VMART). The improvement in
RE (8.1 per cent lower oxygen cost) correlated with the improvement in VMART. VO2max
was not affected by the training intervention. Thus, the gains in performance were attributed
to improved neuromuscular function that led to gains in muscle power and a lower cost of
steady state transport (Paavolainen et al., 1999b).

Plyometric training and running economy

Plyometric training has been widely used to support and enhance performance in a variety of
sport activities (Markovic and Mikulic, 2010). Commonly, this type of training has been
advocated for sport tasks that incorporate jumping, agility and explosive power generation.
The training technique revolves around the stretch-shortening cycle (SSC) of muscle activation.
By challenging the neural and musculotendinous systems that are involved via repeated SSCs,
it is believed that more force can be generated over less time, allowing for an increase in power
production by the muscle. Thus, plyometric training would theoretically stimulate neuro-
muscular adaptations that could have application for many types of movement tasks, including
distance running. One of the draws for endurance athletes is that plyometric-type training has
been reported to induce positive neural adaptations and mechanical advantages with an
attenuated hypertrophic response that is commonly associated with heavy resistance training
(Häkkinen et al., 1985; Saunders et al., 2006). Plyometric training has also been found to
stimulate neuromuscular adaptations (better maintained muscle recruitment pattern) that support
the bike-to-run transition in triathletes (Bonacci et al., 2011). However, the neuromuscular
adaptations that developed did not translate into improved economy upon completion of the
bicycling phase (Bonacci et al., 2011).
SSC training involving plyometrics has focused primarily on the lower body musculature.
Saunders et al. (2006) examined the effects of 9 weeks of plyometric training on running
economy in a population of highly trained distance runners. Plyometric exercises employed
fast concentric/eccentric movement patterns and included activities such as straight-leg jumps,
squat jumps for height, fast feet drills (where minimum ground contact time and high force
production were emphasized), bounding, and high-skipping among others. Running economy
(fixed speeds of 14, 16 and 18 km.h–1) was assessed prior to and after 5 and 9 weeks of plyometric
training. RE was unchanged at the lower running speeds for any time-point. However, a
significant improvement in economy was noted after 9 weeks of plyometric training at the
highest running speed (18 km.h–1). The oxygen requirement for this speed was reduced by
4.1 per cent in the plyometric group, while no change was found in the control group. No
other cardiorespiratory variables were affected by the training intervention during RE testing.
Likewise, no changes in max were found. However, there was a trend for the VO2 versus
running speed slope to be lower in the plyometric group after 9 weeks of training. The
plyometric group also tended to show improvement (14.7 per cent) in average power
production for a five-jump plyometric test (p = 0.11) at 9 weeks. Improved RE was attributed,
in part, to improved locomotor muscle metabolism (i.e. improved efficiency of ATP use) and
improved recovery of elastic energy (Saunders et al., 2006). It was also speculated that an
improvement in whole-body mechanics occurred only with the fastest speed under evaluation,
which was deemed more representative of normal training and competition running speed.
Finally, since cardiorespiratory substrate use measures were largely unaffected, the improved

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RE at the fastest speed was primarily attributed to gains in muscular power and conservation
of elastic energy and/or from better gait coordination and technique that accompanied
plyometric training (Saunders et al., 2006). While mechanics and changes in gait patterns can
be evaluated and quantified, methods for quantifying changes in elastic energy remain elusive
(Saunders et al., 2004b). Notably, it has been theorized that those most likely to benefit from
interventions designed to boost economy will be the lesser-trained runners (McCann and
Higginson, 2008). Thus, plyometric training may be found to elicit more profound and possibly
multifaceted effects in less well-trained runners.
For a comprehensive review on plyometric training and performance adaptations, see
Markovic and Mikulic (2010).

Barefoot running and neuromuscular adaptation

Barefoot running has gained considerable attention in the past several years. A recent PubMed
search using ‘barefoot running’ revealed that 28 publications have been produced since January
2011 in this general area. Despite the recent heightened attention, a number of sports medicine
investigations were conducted in the early to mid 1980s involving barefoot running (Burkett
et al., 1985; Robbins and Hanna, 1987; Rodgers and Leveau, 1982). Early investigators explored
factors including the potential for barefoot running to help prevent running-related injury
(Robbins and Hanna, 1987), and effects of barefoot running on limb kinematics (Burkett
et al., 1985). Robbins and Hanna (1987) proposed that running shoes impede sensory input
and induce more foot rigidity. These effects of shod running could be associated with a greater
injury frequency.
More contemporary investigators have also investigated kinematic adaptations related to
barefoot or minimalist running (Divert et al., 2005b; Lieberman, 2012; Perl et al., 2012). Perl
et al. (2012) studied the effects of footwear on running economy and kinematics. It is commonly
found that shod runners are more prone to rearfoot strike while barefoot runners are more likely
to show forefoot strike patterns. These differences may influence kinematic and economy factors
during steady state running. Perl et al. (2012) compared minimalist footwear (Vibram
FiveFingers™) to standard running shoes (Asics Gel-Cumulus 10™). Two trials were completed
using each style of footwear: forefoot strike (FFS) and rearfoot strike (RFS). Trial order was
randomized and expired gases were collected while at steady state (trials were a minimum of
5 minutes). Stride frequency and foot mass were matched between footwear types. Footstrike
style (FFS versus RFS) was not found to affect economy within each type of footwear.
However, economy was influenced by footwear type within a strike style such that during FFS
running, the minimally shod condition was 2.4 per cent more economical. RFS was found to
be 3.2 per cent more economical when performed with minimalist footwear. There was wide
variability within subjects when minimally shod for economy, ranging from 9.7 per cent more
economical to 7.3 per cent less economical; though most subjects were more economical with
minimalist footwear. While the mechanisms are yet to be soundly confirmed, economical
advantages associated with minimalist footwear were attributed to: (1) greater storage of elastic
energy and recoil in the longitudinal arch; (2) less knee flexion during gait (a possible energy
conservation adaptation); and (3) increased elastic energy storage in the Achilles tendon (Perl
et al., 2012). Further work is needed to investigate each of these proposed mechanisms for
enhanced economy in minimalist footwear. Likewise, given the broad variability within subjects
under the different conditions, further examination of individual variation is warranted.
Divert et al. (2005a) explored stiffness adaptations under shod and barefoot conditions during
treadmill running in male runners. They found that vertical stiffness and leg stiffness decreased

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during 4 minutes of running (3.61 m.s–1) in the shod condition, while in the barefoot condition
stiffness was stable. A significant difference between conditions was noted for stride frequency
(5 per cent higher in barefoot), while vertical displacement, leg compression and maximal
vertical force were significantly lower (15, 5 and 3.5 per cent, respectively) in barefoot running.
The lower mean and vertical leg stiffness measures in the shod condition were attributed to
the notion that musculotendinous stiffness was not increased adequately to compensate for
shoe compliance during the shod running condition (Divert et al., 2005a) Alteration of shoe
properties (both acute and chronic) should be considered when examining kinematic and
economical measures of runners.
de Koning (1992) suggested that neurologic adaptation may result from barefoot running.
He noted significant differences between shod and barefoot running in EMG activity of the
tibialis anterior muscle, with higher activity observed during barefoot running. He suggested
that this difference may be due to an attempt of the neuromuscular system to attenuate the
ground impact force by controlling plantar flexion and/or foot pronation during landing.
Barefoot running was also found to elicit a lower landing velocity than shod running. Whether
these adaptations are more about protection, comfort or fatigue remains to be answered (de
Koning, 1992). Based on early and more contemporary research, it is evident that barefoot

Table 3.1 Summary of key training-specific neuromuscular adaptations and associated economy and
performance-related benefits

Training method Neuromuscular adaptation Benefit Reference

Run-only training Increased leg stiffness
Improved biarticular muscle
coactivation in stance phase
Increased rate of force
generation and reduced
ground contact time
Concurrent Increased leg power, leg
resistance and speed and sprint ability
run training
Increased EMG activity of
vastus lateralis and medius
Improved Type I fibre
quality(?)
Reduced ground contact
time with constant run
velocity
Concurrent Improved neuromuscular
plyometric and control
run training
Improved leg power,
locomotor muscle
metabolism, recovery of
elastic energy
Improved storage of elastic Craib et al. (1996)
energy and lower metabolic Dalleau et al. (1998)
cost
Increased leg stiffness and
higher economy
Improved 5 km race
performance
Improved economy
maintenance
Improved economy and
vVO2max
More efficient fibre
performance
Improved 5 km race time
and economy
Better transition from
cycle to run in triathletes
Improved economy at
higher run speeds, but not
at slower run speeds
Heise et al.(2008)
Paavolainen et al.
(1999c)
Paavolainen et al.
(1999a)
Taipale et al. (2010)
Tanaka and Swensen
(1998)
Paavolainen et al.
(1999b)
Bonacci et al. (2011)
Saunders et al. (2006)

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running will influence gait pattern, lower extremity muscle activation patterns and leg stiffness,
economy and potentially injury incidence. However, neuromuscular adaptations that develop
from introduction of barefoot running are yet to be clearly established. Further, whether
adaptive responses to barefoot running develop as a result of protection from injury (i.e. shock
attenuation) or more from an inherent drive for higher economy of transport is unknown. For
further information on the evolutionary influences of barefoot running and its medical
implications, Lieberman (2012) provides a review of these topics.

Summary
Numerous factors have the potential to alter running patterns. As with any form of physical
training, adaptation will occur. The adaptive response to run training appears to generate several
positive outcomes: (1) reduced cost of transport; (2) attenuated impact forces as a means of
minimizing injury occurrence; and (3) delayed fatigue development. Neuromuscular adaptations
that contribute to these advantages appear to be related to developing increased leg stiffness
and reducing knee flexion during gait, which both help to conserve elastic energy. Other
neuromuscular factors that may contribute to adaptation include increased stiffness of the
Achilles tendon and the tendon surae. Further beneficial adaptations may result from
incorporation of resistance, plyometric training or barefoot running. As above, neuromuscular
adaptations that result from these types of training are also commonly linked to stiffness changes.
Other factors that should be investigated more thoroughly include the effects of eccentric
loading of muscle on neuromuscular adaptations, running surface rigidity and gait alterations,
and the effects of footwear rigidity on economy, adaptive response and forces applied to the
lower extremity. In addition, clothing that has been designed to support run performance (i.e.
elastic compression stockings) should also be studied more thoroughly to clearly elucidate their
effects on mechanical and energetic factors of locomotion.

References
Bonacci, J., Chapman, A., Blanch, P. and Vicenzino, B. (2009) ‘Neuromuscular adaptations to training,
injury and passive interventions: implications for running economy’, Sports Medicine, 39(11): 903–21.
Bonacci, J., Green, D., Saunders, P. U., Franettovich, M., Blanch, P. and Vicenzino, B. (2011) ‘Plyometric
training as an intervention to correct altered neuromotor control during running after cycling in
triathletes: a preliminary randomized controlled trial’, Physical Therapy in Sport, 12(1): 15–21.
Brisswalter, J. and Legros, P. (1995) ‘Use of energy cost and variability in stride length to assess an optimal
running adaptation’, Perceptual and Motor Skills, 80(1): 99–104.
Burkett, L. N., Kohrt, W. M. and Buchbinder, R. (1985) ‘Effects of shoes and foot orthotics on VO2
and selected frontal plane knee kinematics’, Medicine and Science in Sports and Exercise, 17(1): 158–63.
Chapman, R. F., Laymon, A. S., Wilhite, D. P., McKenzie, J. M., Tanner, D. A. and Stager, J. M.
(2012) ‘Ground contact time as an indicator of metabolic cost in elite distance runners’, Medicine and
Science in Sports and Exercise, 44(5): 917–25.
Cohen, R., Mitchell, C., Dotan, R., Gabriel, D., Klentrou, P. and Falk, B. (2010) ‘Do neuromuscular
adaptations occur in endurance-trained boys and men?’, Applied Physiology, Nutrition, and Metabolism,
35(4): 471–9.
Costill, D. L., Thomason, H. and Roberts, E. (1973). ‘Fractional utilization of the aerobic capacity during
distance running’, Medicine and Science in Sports, 5(4): 248–52.
Craib, M. W., Mitchell, V. A., Fields, K. B., Cooper, T. R., Hopewell, R. and Morgan, D. W. (1996)
‘The association between flexibility and running economy in sub-elite male distance runners’, Medicine
and Science in Sports and Exercise, 28(6): 737–43.
Dalleau, G., Belli A., Bourdin, M. and Lacour, J. R. (1998) ‘The spring-mass model and the energy cost
of treadmill running’, European Journal of Applied Physiology and Occupational Physiology, 77(3): 257–63.

34
 Neuromuscular adaptations in running
Downloaded By: 10.3.98.104 At: 00:08 10 May 2022; For: 9780203123355, chapter3, 10.4324/9780203123355.ch3

de Koning, J. J. (1992) ‘Adaptations in running kinematics and kinetics due to shoes and playing surfaces’,
Proceedings of the Third International Symposium on Sport Surfaces; a Symposium of the University of Calgary,
Calgary: University of Calgary.
Divert, C., Baur, H., Mornieux, G., Mayer, F. and Belli, A. (2005a) ‘Stiffness adaptations in shod running’,
Journal of Applied Biomechanics, 21(4): 311–21.
Divert, C., Mornieux, G., Baur, H., Mayer, F. and Belli, A. (2005b) ‘Mechanical comparison of barefoot
and shod running’, International Journal of Sports Medicine, 26(7): 593–8.
Elliott, M. C., Wagner, P. P. and Chiu, L. (2007) ‘Power athletes and distance training: physiological
and biomechanical rationale for change’, Sports Medicine, 37(1): 47–57.
Ferrauti, A., Bergermann, M. and Fernandez-Fernandez, J. (2010) ‘Effects of a concurrent strength and
endurance training on running performance and running economy in recreational marathon runners’,
Journal of Strength and Conditioning Research, 24(10): 2770–8.
Fletcher, J. R., Esau, S. P. and Macintosh, B. R. (2010) ‘Changes in tendon stiffness and running economy
in highly trained distance runners’, European Journal of Applied Physiology, 110(5): 1037–46.
Häkkinen, K., Komi, P. V. and Alen, M. (1985) ‘Effect of explosive type strength training on isometric
force- and relaxation-time, electromyographic and muscle fibre characteristics of leg extensor muscles’,
Acta Physiologica Scandinavica, 125(4): 587–600.
Hayes, P. R., Bowen, S. J. and Davies, E. J. (2004) ‘The relationships between local muscular endurance
and kinematic changes during a run to exhaustion at vVO2max’, Journal of Strength and Conditioning
Research, 18(4): 898–903.
Hayes, P. R., French, D. N. and Thomas, K. (2011) ‘The effect of muscular endurance on running
economy’, Journal of Strength and Conditioning Research, 25(9): 2464–9.
Heise, G., Shinohara, M. and Binks, L. (2008) ‘Biarticular leg muscles and links to running economy’,
International Journal of Sports Medicine, 29(8): 688–91.
Jones, A. M. (2002) ‘Running economy is negatively related to sit-and-reach test performance in
international-standard distance runners’, International Journal of Sports Medicine, 23(1): 40–3.
Kram, R. and Taylor, C. R. (1990) ‘Energetics of running: a new perspective’, Nature, 346(6281): 265–7.
Lieberman, D. E. (2012) ‘What we can learn about running from barefoot running: an evolutionary
medical perspective’, Exercise and Sport Sciences Reviews, 40(2): 63–72.
McCann, D. J. and Higginson, B. K. (2008) ‘Training to maximize economy of motion in running gait’,
Current Sports Medicine Reports, 7(3): 158–62.
Markovic, G. and Mikulic, P. (2010) ‘Neuro-musculoskeletal and performance adaptations to lower-
extremity plyometric training’, Sports Medicine, 40(10): 859–95.
Mikkola, J., Vesterinen, V., Taipale, R., Capostagno, B., Häkkinen, K. and Nummela, A. (2011) ‘Effect
of resistance training regimens on treadmill running and neuromuscular performance in recreational
endurance runners’, Journal of Sports Sciences, 29(13): 1359–71.
Paavolainen, L., Häkkinen, K., Hamalainen, I., Nummela, A. and Rusko, H. (1999a) ‘Explosive-strength
training improves 5-km running time by improving running economy and muscle power’, Journal of
Applied Physiology, 86(5): 1527–33.
Paavolainen, L., Nummela, A., Rusko, H. and Häkkinen, K. (1999b) ‘Neuromuscular characteristics and
fatigue during 10 km running’, International Journal of Sports Medicine, 20(8): 516–21.
Paavolainen, L. M., Nummela, A. T. and Rusko, H. K. (1999c) ‘Neuromuscular characteristics and
muscle power as determinants of 5-km running performance’, Medicine and Science in Sports and Exercise,
31(1): 124–30.
Perl, D. P., Daoud, A. I. and Lieberman, D. E. (2012) ‘Effects of footwear and strike type on running
economy’, Medicine and Science in Sports and Exercise, 44(7): 1335–43.
Robbins, S. E. and Hanna, A. M. (1987) ‘Running-related injury prevention through barefoot adaptations’,
Medicine and Science in Sports and Exercise, 19(2): 148–56.
Rodgers, M. M. and Leveau, B. F. (1982) ‘Effectiveness of foot orthotic devices used to modify pronation
in runners’, The Journal of Orthopaedic and Sports Physical Therapy, 4(2): 86–90.
Ross, A., Leveritt, M. and Riek, S. (2001) ‘Neural influences on sprint running: training adaptations and
acute responses’, Sports Medicine, 31(6): 409–25.
Saunders, P. U., Pyne, D. B., Telford, R. D. and Hawley, J. A. (2004a) ‘Reliability and variability of
running economy in elite distance runners’, Medicine and Science in Sports and Exercise, 36(11): 1972–6.
Saunders, P. U., Pyne, D. B., Telford, R. D. and Hawley, J. A. (2004b) ‘Factors affecting running
economy in trained distance runners’, Sports Medicine, 34(7): 465–85.

35
 William A. Braun
Downloaded By: 10.3.98.104 At: 00:08 10 May 2022; For: 9780203123355, chapter3, 10.4324/9780203123355.ch3

Saunders, P. U., Telford, R. D., Pyne, D. B., Peltola, E. M., Cunningham, R. B., Gore, C. J. and
Hawley, J. A. (2006) ‘Short-term plyometric training improves running economy in highly trained
middle and long distance runners’, Journal of Strength and Conditioning Research, 20(4): 947–54.
Taipale, R. S., Mikkola, J., Nummela, A., Vesterinen, V., Capostagno, B., Walker, S., Gitonga, D.,
Kraemer, W. J. and Häkkinen, K. (2010) ‘Strength training in endurance runners’, International Journal
of Sports Medicine, 31(7): 468–76.
Tanaka, H. and Swensen, T. (1998) ‘Impact of resistance training on endurance performance. A new
form of cross-training?’, Sports Medicine, 25(3): 191–200.
Thayer, R., Collins, J., Noble, E. G. and Taylor, A. W. (2000) ‘A decade of aerobic endurance training:
histological evidence for fibre type transformation’, The Journal of Sports Medicine and Physical Fitness,
40(4): 284–9.
Turner, A. M., Owings, M. and Schwane, J. A. (2003) ‘Improvement in running economy after 6 weeks
of plyometric training’, Journal of Strength and Conditioning Research, 17(1): 60–7.
Young, W. B. (2006) ‘Transfer of strength and power training to sports performance’, International Journal
of Sports Physiology and Performance, 1(2): 74–83.

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