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Strength, power, and postural control in seniors: Considerations for functional

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DOI: 10.1080/17461390802478066

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 European Journal of Sport Science, November 2008; 8(6): 325
340

REVIEW ARTICLE

Strength, power, and postural control in seniors: Considerations for

functional adaptations and for fall prevention

URS GRANACHER1, LUKAS ZAHNER1, & ALBERT GOLLHOFER2

1
Institute of Exercise and Health Sciences, University of Basel, Basel, Switzerland, and 2Institute of Sport and Sport Science,
University of Freiburg, Freiburg, Germany

Abstract
The ageing neuromuscular system is affected by structural and functional changes that lead to a general slowing down of
neuromuscular performance and an increased risk of falling. As a consequence, the process of ageing results in a reduced
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008

ability to develop maximal and explosive force, as well as in deficits in static and dynamic postural control. A decrease in the
number and size of type II fibres in particular accounts for the age-related decline in muscle mass (sarcopaenia) and strength
performance. Multiple denervation and re-innervation processes of muscle fibres seem to be responsible for the reduced
number of muscle fibres. Recently, it has been suggested that it is not the decline in motoneurons that accounts for the loss
in number of muscle fibres but the disturbed potential of fibre regeneration and re-innervation. Furthermore, an age-related
reduction in the number of satellite cells has also been associated with sarcopaenia. The ability to compensate for platform
and gait perturbations deteriorates with ageing as reflected in longer onset latencies and inefficient postural responses. All
sites within the somatosensory system are affected by ageing and therefore contribute to postural instability. However,
morphological changes of muscle spindles appear primarily to be responsible for the impaired ability to compensate for
balance threats in old age. Given these neuromuscular limitations in old age, it is important to apply adequate training
interventions that delay or even reverse the onset of these constraints. Strength training has the potential to enhance
maximal as well as explosive force production capacity. This is accomplished by neural factors, including an improved
recruitment pattern, discharge rate, and synchronization of motor units. Furthermore, an increase in number of satellite
cells most likely accounts for training-induced muscle hypertrophy. Recent studies have investigated the impact of balance
training in old age on the ability to develop maximal and explosive force. In addition, the effects of balance training on reflex
activity during gait perturbations were also examined. Increases in maximal and explosive force production capacity and an
improved ability to compensate for gait perturbations were observed. It is evident from the literature that researchers are
increasingly studying the effects of more specifically designed training programmes on performance in populations of older
adults. Thus, in the near future, strength training could be replaced by high-velocity forms of power training and balance
training by perturbation-based training programmes. It is hypothesized that this new approach is more efficient in terms of
fall prevention than the traditional approach.

Keywords: Ageing, maximal and explosive force production capacity, postural control, strength and balance training

Introduction old age, (e) the rationale for an impaired postural

Rapid increases in the numbers of senior citizens in
western industrial countries demand intense research
attention regarding the effects of ageing on neuro-
muscular performance and its functional conse-
quences. This review article describes (a) the
increased risk of falling in old age, (b) maximal and
explosive force production capacity in old age, (c) the
rationale for the reduced maximal and explosive force
production capacity in old age, (d) postural control in
control in old age, (f) the importance of strength
training in old age, (g) adaptive processes following
strength training in old age, (h) the importance of
balance training in old age, and (i) adaptive processes
following balance training in old age.
Risk of falling in old age
Around 28
35% of individuals over the age of 65
years sustain at least one fall over a one-year period

Correspondence: U. Granacher, Institute of Exercise and Health Sciences, University of Basel, Brüglingen 33, CH-4052 Basel, Switzerland.
E-mail: urs.granacher@unibas.ch

ISSN 1746-1391 print/ISSN 1536-7290 online # 2008 European College of Sport Science
DOI: 10.1080/17461390802478066
 326 U. Granacher et al.
(Blake et al., 1988; Campbell, Reinken, Allan, &
Martinez, 1981) and this increases to 32
42% in
adults over the age of 75 years (Downton &
Andrews, 1991; Tinetti, Speechley, & Ginter,
1988). About 20% of falls require medical attention;
15% result in joint dislocations, soft tissue bruises,
and contusions, while 5% result in fractures, with
femoral neck fractures occurring at a rate of 1
2%
(Kannus et al., 1999). These fall-related injuries
place a high financial burden on the public health
care system. In the United States, the estimated
direct medical costs for fatal and non-fatal fall-
related injuries totalled US$19.2 billion in 2000
(Stevens, Corso, Finkelstein, & Miller, 2006). Be-
sides the financial costs, fall-related injuries affect
the elderly in terms of a poor quality of life due to
restricted mobility and functional decline. Numer-
ous epidemiological studies have identified a multi-
tude of risk factors for falling (Lord & Dayhew,
2001). Reduced muscle strength as well as deficits in
static and dynamic postural control are important
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008
risk factors for falls in old age (Hausdorff et al., 2001;
Pijnappels, van der Burg, Reeves, & van Dieen,
2008).
Maximal and explosive force production
capacity in old age
Biologic ageing results in an unavoidable decrease in
maximal and explosive force production capacity
(Skelton, Greig, Davies, & Young, 1994). Between
the ages of 30 and 80 years, the average reported
decreases in maximal strength range from 20 to 40%
(Frontera, Hughes, Lutz, & Evans, 1991; Merletti,
Farina, Gazzoni, & Schieroni, 2002), depending on
study design (cross-sectional vs. longitudinal), the
age groups investigated (old vs. very elderly), health
status (e.g. free of chronic disease vs. chronic
disease), and fitness (sedentary vs. active). In terms
of age-related effects on the limb muscles, it is
reported that declines in maximal strength affect
the extensor muscles of the lower extremities most
severely, particularly the ankle and knee extensors
(Frontera et al., 1991; Lynch et al., 1999). This is
presumably due to reduced physical activity in old
age, which primarily affects the lower extremities.
The impact of neuromuscular ageing differs not
only in terms of the muscle groups and types of
contraction studied but also in relation to the
initiation of force. Recent data indicate that age-
related decreases in explosive force production
capacity exceed those in maximal strength, with the
most severe losses occurring between the seventh
and ninth decade of life (McNeil, Vandervoort, &
Rice, 2007). This decline has been illustrated in
concentric types of muscle actions, in jumps in the
stretch
shortening cycle, as well as in explosive
isometric contractions (Bosco & Komi, 1980; Häk-
kinen, Pastinen, Karsikas, & Linnamo, 1995; Iz-
quierdo, Aguado, Gonzalez, Lopez, & Häkkinen,
1999). McNeil et al. (2007) reported a 25% decrease
in power of the dorsiflexors (isotonic contractions)
between the third and seventh decade of life. This
reduction was doubled in the next two decades, so
that men in their ninth decade of life produced 60%
less power than young men (mean age 26 years).
From a functional or therapeutic point of view, it
is of interest to determine whether maximal force
capacity and/or explosive power have an impact on
activities of daily living (ADL), postural control, and
the occurrence of falls. Significant correlations
between explosive force production capacity of the
leg extensors and the performance of ADL, such as
stair-climb time, habitual and maximal gait velocity,
and time to complete the 6-min walk test, have been
reported (Bean et al., 2002; Puthoff & Nielsen,
2007). Manini et al. (2007) provided sex-specific
knee extensor strength cut-points to identify well-
functioning older adults (age 70
79 years) at high
and low risk of future mobility problems. It was
reported that women and men with cut-points of
1.01 and 1.13 N ×m ×kg 1, respectively, were at high
risk and those with values of 1.34 and 1.71 N ×m ×
kg 1, respectively, were at low risk of severe limita-
tions to their mobility (Manini et al., 2007). In
another study, Izquierdo et al. (1999) investigated
the association between rate of force development of
the leg extensors and postural control in a cohort of
young (mean age 21 years), middle-aged (mean age
40 years), and older men (mean age 71 years). The
authors observed that the decreased ability to
develop force rapidly in elderly men is associated
with a reduced neuromuscular capacity in control-
ling postural sway. In terms of the high incidence of
falling in old age, Pijnappels et al. (2008) identified
lower limb strength and rate of moment generation
as limiting factors for balance recovery after tripping.
These authors reported that maximum isometric
push-off force in a leg press was the best measure to
differentiate elderly fallers from non-fallers. Re-
cently, asymmetry in lower limb muscle strength
has been detected as a risk factor for falls in elderly
women (mean age 74 years) (Skelton, Kennedy, &
Rutherford, 2002). It is argued that asymmetry
between the limbs most likely impairs the mainte-
nance of the body’s centre of gravity over the small
base of support (Portegijs et al., 2006).
Rationale for the reduced maximal and
explosive force production capacity in old age
It is well established that there is an association
between the magnitude of skeletal muscle mass and
the force a muscle can generate (Kirkendall &

Garrett, 1998). Thus, a decline in skeletal muscle
mass with ageing, also known as ‘‘senile sarcopae-
nia’’ (Roubenoff, 2000), must contribute to the age-
related strength loss. In fact, studies have established
a 30
50% decrease in skeletal muscle mass in both
men and women between the ages of 40 and 80 years
(Akima et al., 2001; Lexell, Taylor, & Sjostrom,
1988). Janssen and colleagues (Janssen, Heymsfield,
Wang, & Ross, 2000) applied whole-body magnetic
resonance imaging in a heterogeneous (ethnically
mixed) sample of 468 men and women aged 18
88
years. The authors observed a reduction in skeletal
muscle mass relative to body mass starting in the
third decade. However, a noticeable decrease in
absolute skeletal muscle mass was not identified
until the end of the fifth decade. The loss in whole-
body muscle mass was independent of change in
stature and was greater in men than in women, both
in absolute terms and relative to body mass.
Furthermore, the decline in skeletal muscle mass
with age was greater in the lower body in both men
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and women. Given that the muscles of the lower
extremities are required for most ADL (e.g. walking,
stair climbing), the predominant loss of muscle mass
in the lower body can most likely be explained by an
age-related reduction in physical activity.
What are the factors that contribute to sarcopaenia?
The reduced muscle mass with ageing is caused by
(a) a reduction of the volume of individual fibres, (b)
a reduction of the total number of fibres, (c) a
combination of the two (Faulkner, Larkin, Claflin, &
Brooks, 2007), and (d) alterations in the internal
arrangement of muscle fibres, known as ‘‘muscle
architecture’’. The latter was investigated by Narici
and colleagues (Narici, Maganaris, Reeves, & Ca-
podaglio, 2003), who analysed gastrocnemius med-
ialis muscle fascicle length as well as the angle at
which the fascicles insert into the tendinous sheath
(aponeurosis), also known as pennation angle, in a
cohort of young (age 27
42 years) and older (age
70
81 years) physically active men. It was reported
that gastrocnemius muscle fascicles were shorter by
10% in elderly compared with young adults. In
addition, pennation angle was 13% less in the
elderly. These results are most likely caused by an
age-related loss of sarcomeres in series. According to
Narici et al. (2003), this loss impairs muscle function
in old age by affecting the length
tension, the force

velocity, and the power
velocity relationships of the
associated muscle. However, ageing results not only
in a loss in sarcomeres in series but also in a decline
in sarcomeres in parallel. In fact, a reduction in
muscle mass before 50 years of age is primarily
attributable to a loss in the cross-sectional area of
individual fibres (sarcomeres in parallel) due to a
Exercise and fall prevention in seniors 327
sedentary lifestyle (Lexell & Taylor, 1991). Later in
life, additional loss in muscle mass is caused by a
reduction in the number of muscle fibres. Lexell et
al. (1988) observed a 50% decrease in the number of
muscle fibres in the vastus lateralis in men aged 50

80 years. The reduction in the number of muscle
fibres in old age is accompanied by an infiltration of
connective tissue and fat (Edstrom & Ulfhake,
2005), as well as a deterioration of satellite cells.
Renault and colleagues (Renault, Thornell, Eriks-
son, Butler-Browne, & Mouly, 2002) observed a
1.44
1.77% lower proportion of satellite cells in
older (mean age 74 years) than in younger (mean age
23 years) individuals. The reduced number of
muscle fibres and satellite cells has been attributed
to ageing-induced susceptibility to nuclear apoptosis
(i.e. death of a nucleus without death of the entire
cell) (Alway & Siu, 2008). A decline in the number
of satellite cells has a double-negative effect on the
ageing muscle. First, the regenerative potential of
human skeletal muscle is impaired, and second, the
ability of the ageing muscle to respond to adequate
training loads with hypertrophy is diminished.
The question of whether a specific fibre type is
affected by neuromuscular ageing requires clarifica-
tion. Studies indicate that type II fibres are more
vulnerable to the ageing process than type I fibres
(Figure 1) (Coggan et al., 1992; Lexell et al., 1988).
Lexell et al. (1988) observed a 26% reduction in the
size of type II fibres in the vastus lateralis from age
20 to 80 years. A biopsy study of the gastrocnemius
muscle of young (mean age 24 years) and elderly
men and women (mean age 64 years) showed a 13

31% decrease in type IIa and IIb cross-sectional area
(Coggan et al., 1992). There also appear to be
various mechanisms responsible for the reduction
in number and size of skeletal muscle fibres in old
age. A decline in physical activity appears to be
primarily responsible for the loss in muscle size,
whereas nuclear apoptosis of muscle fibres and a
disruption in neural input most likely causes the
reduction in type II muscle fibres in particular
(Alway & Siu, 2008; Doherty, 2003). The latter is
caused by the successive removal of especially large
alpha motoneurons (Terao et al., 1996b). The loss of
fast motoneurons leaves type II fibres denervated.
Some fibres become re-innervated by axonal sprout-
ing of small alpha motoneurons (Roos, Rice, &
Vandervoort, 1997). The result is an increased
innervation ratio in the remaining slow motor units
(Roos et al., 1997). Fibres that do not become re-
innervated remain denervated and are eventually
lost. Recent evidence suggests, however, that it is not
the loss in motoneurons that is responsible for
sarcopaenia, but a disturbed potential of fibre
regeneration and re-innervation (Edstrom et al.,
2007). It is argued that the age-related loss of
 328 U. Granacher et al.
Figure 1. Age-related changes in the neuromuscular system responsible for the reduced maximal and explosive force production capacity,
as well as for an impaired static and dynamic postural control.
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008
motoneurons is relatively small (10
15%) (Ulfhake
et al., 2000) and that this decline cannot fully
account for the decrease in the number of muscle
fibres. Besides the decline in number and size of
skeletal muscle fibres, there are also signs of regen-
eration in terms of an increased expression of
markers of muscle fibre regeneration (Edstrom &
Ulfhake, 2005). This implies that a number of
motoneurons are partially denervated of muscle
fibres; however, at the same time, the intact branches
of these neurons try to re-innervate nearby discon-
nected muscle fibres. Since the plasticity of motor
axons is decreased in senescence (Carlson & Faul-
kner, 1998), it is not the successive removal of
motoneurons but a failure to re-innervate vacant
fibres that eventually results in the loss of muscle
fibres (Edstrom et al., 2007).
The decline in muscle mass alone does not
account for the age-related decrease in maximal
and explosive force production capacity. In addition,
neural factors (Roos, Rice, Connelly, & Vander-
voort, 1999) also contribute to impaired strength
performance in old age. In a study by Macaluso et al.
(2002), maximum isometric torque (MVC) and
surface electromyography (EMG) of the knee ex-
tensors (vastus lateralis) and flexors (biceps femoris)
were assessed in young (mean age 23 years) and
elderly healthy active women (mean age 69 years).
The authors reported marked reductions in MVC
torque of the knee extensors and flexors in older
versus young adults that were paralleled by signifi-
cant decreases in voluntary drive to the associated
agonist muscles as well as increases in antagonist co-
activation during knee extension. First, higher levels
of antagonist co-activation contribute to strength
deficits in old age; second, they appear to serve as a
compensatory mechanism for a reduced ability to
fully activate the agonist muscle; and third, they
stiffen the muscle joint complex and thus provide
more stability (Macaluso et al., 2002). The reduced
central drive to the agonist muscle can be attributed
to a smaller number of recruited motor units, a
decreased firing rate of individual motor units, and
an impaired motor unit synchronization (Macaluso
et al., 2002). Connelly and colleagues (Connelly,
Rice, Roos, & Vandervoort, 1999) observed age-
related decreases in motor unit discharge rate for the
tibialis anterior. However, Kamen and Roy (2000)
found no differences in motor unit synchronization
between young and elderly adults in the first dorsal
interosseous muscle. On the basis of the existing
literature, it is not possible to allocate the described
neural deficits to either constraints in maximal or
explosive force production. Further studies are
necessary to clarify the issue of neural deficits.
Postural control in old age
Postural control has been defined as the control of
the body’s position in space for the purpose of
balance and orientation (Shumway-Cook & Woolla-
cott, 2001). The ability to control posture can be
described as a dynamic process across the life span.
There is evidence that young children and elderly
adults show the largest magnitudes of postural sway,
when measured on a force platform. Therefore, a U-
shaped dependency between balance and age can be
postulated (Hytonen, Pyykko, Aalto, & Starck,
1993).
Era et al. (2006) assessed static postural control on
a force platform in a randomly selected sample of
7979 individuals aged 30 years and over. They

observed that deterioration of the postural control
mechanisms begins relatively early in life. Differ-
ences in balance performance were already apparent
among young (30- to 39-year olds) and middle-aged
adults (40- to 49-year olds) and became even more
pronounced after the age of 60 years. The differ-
ences between age groups were seen in both males
and females but in males the decline with age was
more evident (Maki, Holliday, & Fernie, 1990).
Anthropometric factors such as height and foot
length could also account for the observed gender
differences, at least to a certain extent (Maki et al.,
1990).
In terms of age-related effects on dynamic pos-
tural control, Lin and Woollacott (2002) determined
postural muscle response characteristics following
various sizes of support surface perturbations in
young (mean age 25 years), stable older (mean age
73 years), and unstable older adults (mean age 76
years). Slower onset latencies, smaller magnitudes of
postural responses, and longer maintenance of
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008
postural muscle activation were observed in response
to platform perturbations in both stable and unstable
older adults compared with the young adults.
Whereas delays in onset times and smaller ampli-
tudes of muscle responses can be classified as age-
related deteriorations in postural control, the pro-
longed muscle activation might be a compensatory
mechanism to help preserve postural stability (Lin &
Woolacott, 2002). It is interesting to note that,
unlike the stable older adults, unstable older adults
were not able to show this compensatory mechanism
in all test conditions. Using a more functional
approach, Tang and Woollacott (1998) investigated
postural responses to unexpected forward slips
during walking in young adults (mean age 25 years)
and active older adults (mean age 74 years). A
similar activation sequence of postural muscles in
response to accelerating perturbation impulses in
young and elderly individuals was observed. How-
ever, the postural responses of older adults showed
longer onset latencies, smaller magnitudes, and
longer burst durations than those of young adults.
Older adults also displayed a longer co-activation
duration for the knee agonist/antagonist pair on the
perturbed and on the non-perturbed side. Different
gait characteristics (earlier contralateral foot strike,
shortened stride length) were also observed between
the two age groups.
Inefficient balance strategies make older adults
more prone to falling than young adults. Pavol and
colleagues (Pavol, Owings, Foley, & Grabiner, 2001)
identified delayed support limb loading (
145 ms)
in response to an external perturbation (trip) as a
deficit that increases the risk of falling. In addition,
Fernie and colleagues (Fernie, Gryfe, Holliday, &
Llewellyn, 1982) studied individuals aged over 63
Exercise and fall prevention in seniors 329
years and observed that postural sway was signifi-
cantly greater for those who fell one or more times in
a year than for those who did not fall. Downton and
Andrews (1990) reported that 49% of their elderly
participants aged over 75 years suffered occasional
or frequent subjective feelings of postural instability,
which was associated with having fallen in the
previous year. In a subsequent study, postural sway
was measured with a force plate and an association
between subjective feelings of instability and objec-
tively impaired balance was detected (Downton,
Sayegh, & Andrews, 1991).
Rationale for an impaired postural control in
old age
For many years, the control of posture was attrib-
uted solely to automatic or reflex controlled muscle
activations (Horak, Macpherson, Rowell, & Shep-
herd, 1996). However, today it is believed that
attentional resources are necessary to stabilize the
body’s centre of gravity effectively over the base of
support (Woollacott & Shumway-Cook, 2002).
Therefore, it can be postulated that the control of
posture demands the complex processing and inte-
gration of sensory information provided by vision,
proprioception, and the vestibular system on a spinal
and supraspinal level (Lephart & Fu, 2000). There is
evidence that proprioceptive input in particular
contributes to the sensory control of balance and
to the successful accomplishment of many ADL in
old age (Bacsi & Colebatch, 2005). Indeed, Mion
et al. (1989) identified impaired proprioception as a
contributing factor to falls among patients in a
rehabilitative setting. Therefore, the focus here will
be on the effects of ageing on proprioceptive
receptors even though age-related impairments
have been proposed at every stage of the postural
control system (Marchetti & Whitney, 2005). [For a
detailed review on this topic, see Shaffer and
Harrison (2007).]
Different articular, muscular, and cutaneous me-
chanoreceptors, including muscle spindles, golgi
tendon organs, pacinian corpuscles, and free nerve
endings, are involved in sensory perception (Lephart
& Fu, 2000). Muscle spindles in particular contri-
bute to human erect posture regulation (Kavounou-
dias, Gilhodes, Roll, & Roll, 1999). It has been
reported that ageing causes morphological changes
to the muscle spindles in terms of an increased
spindle capsule thickness and a loss of intrafusal and
nuclear chain fibres (Liu, Eriksson, Thornell, &
Pedrosa-Domellof, 2005; Swash & Fox, 1972).
These deteriorations appear to be muscle specific
(Kararizou, Manta, Kalfakis, & Vassilopoulos, 2005)
and probably result in a desensitization of this
receptor type (Liu et al., 2005). The reduced afferent
 330 U. Granacher et al.
input via muscle spindles in old age could affect the
control of the muscle’s length and velocity of
contraction and therefore the individual’s ability to
respond adequately to balance threats. Few studies
have investigated the effects of ageing on golgi
tendon organs and articular receptors, although
Morisawa (1998) observed that the total number of
pacinian corpuscles, ruffini receptors, golgi tendon
organ-like receptors, and free nerve endings decrease
with age. The decline in the number of articular
receptors most likely impairs feedback mechanisms
controlling joint movements. In addition, a demye-
lination of sensory axons (Scaglioni, Narici, Maffiu-
letti, Pensini, & Martin, 2003), a decrease in the
number of sensory neurons (Maisonobe, Hauw,
Dani, Hori, & Walter, 1997), changes in presynaptic
inhibition (Earles, Vardaxis, & Koceja, 2001), and a
reduction in the number of inter- (Terao et al.,
1996a) as well as alpha- and gamma-motoneurons
(McNeil, Doherty, Stashuk, & Rice, 2005) contri-
bute to a decreased static and dynamic joint position
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008
sense (Meeuwsen, Sawicki, & Stelmach, 1993;
Verschueren, Brumagne, Swinnen, & Cordo,
2002). Furthermore, deteriorations in ankle plantar
flexor strength, voluntary muscle activation, and
tendon mechanical properties have recently been
associated with a decline in static postural control
(Onambele, Narici, & Maganaris, 2006). Taken
together, these deteriorations (Figure 1) result in
an impaired postural control and eventually lead to a
higher incidence of falls (Tinetti et al., 1988).
Importance of strength training in old age
Several studies (Brochu et al., 2002; Izquierdo et al.,
2003; Holviala, Sallinen, Kraemer, Alen, & Hakki-
nen, 2006) have investigated the effects of resistance
training on maximum voluntary contraction, rate of
force development, and postural control in old age
(Table I). It has been frequently observed that even
in this age group strength training results in an
increase in maximal as well as explosive force
production capacity (Newton et al., 2002). Given
that the ability to generate rapid force is, from a
functional or fall prevention point of view, more
important than the capacity to produce maximal
strength (Suetta, Magnusson, Beyer, & Kjaer,
2007), it is of paramount importance to adopt
strength training programmes that have the potential
to enhance explosive force production capacity.
Indeed, recent work has indicated that strength
training combined with modified power exercises
or even high-speed power training has a greater
impact on explosive force production capacity and
ADL in old age than traditional heavy resistance
strength training (Sayers, 2007). From a functional
and therapeutic point of view, it is important to
determine whether training-induced strength gains
can be sustained in old age. Smith and colleagues
(Smith, Winegard, Hicks, & McCartney, 2003)
measured dynamic muscle strength (one repetition
maximum, 1-RM, or the load that can be lifted only
once) of the arm flexors/extensors and the leg
extensors in three groups of older adults (mean age
72 years). Group I had been undertaking weight
training continuously for 5 years. Group II stopped
after 2 years and Group III acted as a control. After 2
years of strength training, Groups I and II made
significant gains in all muscle groups tested. After
another 3 years of training (Group I) or detraining
(Group II), muscle strength remained significantly
above the baseline value for Group I. The detraining
group also remained above baseline, whereas dy-
namic muscle strength of the control group declined
over the 5-year period. Resistance training certainly
has the potential to improve strength performance in
old age. However, results on the long-term effects of
strength training on postural control are contra-
dictory. Whereas Brochu et al. (2002) and Holviala et
al. (2006) observed improved postural control after
resistance training, Schlicht and colleagues
(Schlicht, Camaione, & Owen, 2001) did not. [For
a systematic review on this topic, see Orr, Raymond,
and Fiatarone (2008).] A recent study investigated
the acute effects of resistance training in elderly
persons on postural control (Moore, Korff, &
Kinzey, 2005). As a result of this study, therapists
and practitioners are advised to assist elderly clients
immediately after resistance training due to an acute
fatigue-induced negative effect of resistance exercises
on postural control.
Many studies have examined the effectiveness of
exercise in preventing falls. Few studies, however,
have investigated the impact of strength training
alone on fall prevention (Fiatarone et al., 1994).
However, to the authors’ knowledge, no effect of
resistance training on fall prevention has been
shown. Most studies used some combination of
resistance, balance, endurance, and flexibility exer-
cises and found a reduced incidence of falling
(Rubenstein et al., 2000).
Adaptive processes following strength training
in old age
It is well established that the ageing neuromuscular
system has the potential to respond to an adequate
training regimen with plasticity (Vandervoort,
2002). Different factors contribute to increases in
strength performance depending on the duration of
the training period and the applied training load.
During the first 2 weeks of resistance training,
strength gains occur mainly as a result of an
improved motor skill coordination, which can be
 Table I. Studies of the impact of strength and power training on neuromuscular performance in elderly individuals

Age Type of Exercise Exercise Strength Postural

Authors Sex (years) n training duration intensity gain control
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Fiatarone et al. (1990) m/f ]80 10 HRT 8 weeks 80% Leg extensors (1-RM): 174% Improved performance in: tandem gait
1-RM speed, 48%
Grimby et al. (1992) m ]78 9 HRT 8 weeks 100% Concentric action knee extension (308/s): 10%
isokinetic Eccentric action knee extension (308/s): 19%
Lexell et al. (1995) m/f ]70 35 HRT 11 weeks 85% Elbow flexion (1-RM): 49%
1-RM Knee extension (1-RM): 163%
Häkkinen et al. (1998a) m/f ]67 42 PT 24 weeks 50
80% Knee extension (MVC): 36
57%
HRT 1-RM Leg extension (RFD): 28
40%
Knee flexion (MVC): 14
17%
Harridge et al. (1999) m/f ]85 11 HRT 12 weeks 80% 1-RM Knee extension (1-RM): 134%
Knee extension (MVC): 17%
Yarasheski et al. (1999) m/f ]76 17 HRT 12 weeks 65
75% Knee extension (MVC): 6
22%
1-RM Knee extension (608/s): 6
7%
Häkkinen et al. (2000) m/f ]62 36 PT 24 weeks 50-80% Leg extension (1-RM): 23
29% Improved performance in: walking
HRT 1-RM Leg extension (MVC): 23
32% speed, 11%
Schlicht et al. (2001) m/f ]61 24 HRT 8 weeks 75% Leg extension, hip abduction, hip adduction, Improved performance in: walking
1-RM gluteal press, leg press, ankle extension speed, 17%; 5-repetition
(1-RM): 20
48% sit-to-stand time, -15%; no impact of
HRT on one-legged blind balance test
Fielding et al. (2002) f ]72 30 HVRT 16 weeks 70% 1-RM High-velocity resistance training:
vs. Leg extension: 35% (1-RM), 97% (power)
LVRT Knee extension: 45% (1-RM), 33% (power)
Low-velocity resistance training:
Leg extension: 33% (1-RM), 45% (power)

Exercise and fall prevention in seniors

Knee extension: 41% (1-RM), 25% (power)
Scaglioni et al. (2002) m ]65 14 MRT  16 weeks 50-80% Plantar flexion (MVC): 18% No impact of HRT on H-reflex loop
HRT 1-RM Plantar flexion (1-RM): 24%
Hruda et al. (2003) m ]75 25 PT 10 weeks Concentric action knee extension (1808/s): Improved performance in: 8 foot up and
60% go test, 31%; 30-s chair stand test,
Eccentric action knee extension (1808/s): 66%; walk time test, 33%
44%
Granacher et al. (2004, 2006) m ]60 60 HRT 12 weeks 80% Leg extension (MVC): 27% No impact of HRT on ability to compensate
1-RM Leg extension (RFD): 32
56% for gait perturbations (trip)
Holviala et al. (2006) f ]60 48 HVRT 21 weeks Leg extension (1-RM): 27% Improved performance in: walking
Leg extension (MVC): 20% speed and balance performance (PB0.001)
Leg extension (RFD): 18%
Bottaro et al. (2007) m ]60 24 PT vs. 10 weeks 60% Power training: Power training:
MRT 1-RM Leg extension: 27% (1-RM), 31% (power) Improved performance in: 8 foot up and
Arm extension: 28% (1-RM), 37% (power) go test, 15%; 30-s chair stand test, 43%
Moderate resistance training: Moderate resistance training:
Leg extension: 27% (1-RM), 8% (power) Improved performance in: 8 foot up and
Arm extension: 25% (1-RM), 13% (power) go test, 1%; 30-s chair stand test, 6%

331
 332 U. Granacher et al.

Abbreviations: mmale, f female, n number of participants, MRT moderate resistance training, HRT heavy resistance training, PTpower training, HVRT high-velocity resistance training,
classified as a learning effect. After 3
4 weeks of
strength training, predominantly neural adaptive
mechanisms account for the improved strength

LVRT low-velocity resistance training, MVCmaximum voluntary contraction, RFDrate of force development, 1-RMone-repetition maximum, CMJcountermovement jump.
performance (Sale, 2003). Häkkinen (2003) postu-
lated that an increased activation of the prime
movers (improved recruitment pattern, discharge
Postural
control

rate, and synchronization of motor units), an im-

proved co-activation of the synergists, and a reduced
co-activation of the antagonist muscles are respon-
sible for the training-induced strength gains during
this phase of training. If adequate progressive train-
ing loads are applied over a training period lasting 6
weeks or more, muscular factors contribute to the
improved strength performance. Using imaging
techniques such as computer tomography, an en-
largement of muscle cross-sectional area by 5
17%
has been observed in the elderly following resistance
strength training lasting 3 months (Ferri et al., 2003;
Häkkinen & Häkkinen, 1995). Type I, IIa, and IIb
fibre hypertrophy has been reported in response to
Leg extension (MVC): 18%

Leg extension (MVC): 28%

Strength

Leg extension (RFD): 21%

Leg extension (RFD): 51%

resistance training loads (Trappe et al., 2000; Wi-

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gain

drick, Stelzer, Shoepe, & Garner, 2002). Further-

CMJ (height): 10%

CMJ (height): 18%

more, a 10-week progressive strength training

programme resulted in type II subtype transforma-
Age 60
65 years:

Age 80
89 years:

tion from type IIb to IIab to IIa in a cohort of older

men (mean age 61 years) (Häkkinen et al., 1998b).
These results were confirmed by Sharman et al.
(2001), who observed a shift in the expression of
myosin heavy chain (MHC) isoforms form MHC IIb
to MHC IIa (vastus lateralis) after 6 months of high-
intensity
Exercise

75
80%
1-RM

intensity resistance training for the lower extremities

in elderly men and women (mean age 65 years).
What are the underlying mechanisms for training-
induced muscle hypertrophy in old age? Muscle
12 weeks
duration

fibres are post-mitotic cells. This implies that no

Exercise

further cell division takes place after embryonic

differentiation is complete. Therefore, training-in-
duced improvements in muscle cross-sectional area
Type of
training

primarily occur when new nuclei are added to the

PT

muscle fibres. Satellite cells (muscle stem cells)

provide the myonuclei for post-natal growth of
65

muscle fibres. These reserve cells are believed to

n

remain mitotically inactive, but are mobilized by

60
65;
(years)

80
89

increased mechanical loading or damage, playing a

Age

role in both adaptation and repair of muscle (Gold-

spink & Harridge, 2003). Satellite cell activation has
Sex

been documented in young men 4 and 8 days after a

f

single bout of maximal exercise (Crameri et al.,

2004). The number of satellite cells was increased in
young men by 19% following 30 days of resistance
Caserotti et al. (in press)

training and by 31% at 90 days of training. Com-

Table I (Continued)

pared with pre-training values, the number of

satellite cells remained significantly elevated at 3,
10, and 60 days but not at 90 days of detraining
(Kadi et al., 2004). Kadi and Thornell (2000)
Authors

investigated an association between a training-

induced increase in cross-sectional area of muscle

fibres and a gain in number of myonuclei in the
individual fibres, suggesting that satellite cell pro-
liferation is responsible for muscle hypertrophy. In
elderly males and females (mean age 76 years), it has
been shown that proliferation of satellite cells (27%)
can also be induced in response to 12 weeks of
resistance training for the lower extremities (Mackey
et al., 2007). These results indicate that long-term
resistance training generates similar rates of satellite
cell proliferation in young and elderly individuals.
Importance of balance training in old age
Various studies have examined the impact of stan-
dardized balance or sensorimotor training on
strength performance and postural control in old
age (Steadman, Donaldson, & Kalra, 2003; Gran-
acher, Gollhofer, & Strass, 2006; Granacher, Gru-
ber, Strass, & Gollhofer, 2007) (Table II). Steadman
et al. (2003) reported that 6 weeks of balance
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training improved performance in clinical balance
and mobility tests. Granacher et al. (2007) showed
that 12 weeks of balance training improved maximal
and explosive force production capacity in a cohort
of 40 healthy, elderly males aged 60
80 years. To the
authors’ knowledge, there are few published of the
effects of balance training on neuromuscular para-
meters in the elderly (Hu & Woollacott, 1994;
Mynark & Koceja, 2002; Williams, Burke, McCle-
naghan, Hirth, & Huber, 1997). Hu and Woolacott
(1994) investigated the impact of 2 weeks of multi-
sensory balance training on the temporal and spatial
organization of postural responses. Mynark and
Koceja (2002) examined the capability of elderly
individuals to down-train the soleus H-reflex in
response to a balance perturbation. Williams et al.
(1997) scrutinized the effects of long-term balance
training on the Achilles tendon reflex. The results of
these studies support the contention that balance
training results in a shortened onset latency of the
postural muscles (Hu & Wollacott, 1994) as well as
in a down-training of the H-reflex (Mynark &
Koceja, 2002) and the Achilles tendon reflex (Wil-
liams et al., 1997).
In a recent study, Granacher et al. (2006) exam-
ined the effects of 12 weeks of balance or heavy
resistance strength training on the ability to com-
pensate for gait perturbations in elderly men. Bal-
ance training resulted in a decrease in onset latency
and an enhanced reflex activity in the prime mover
compensating for the decelerating perturbation im-
pulse. No significant changes were observed in the
heavy resistance strength group or the control group.
These results imply that balance training could have
a fall prevention effect. In fact, some studies have
shown that balance training has the potential to
Exercise and fall prevention in seniors 333
reduce the incidence of falls in old age (Lord et al.,
2003; Province et al., 1995).
Recently, more specifically designed balance train-
ing programmes, so-called ‘‘perturbation-based
training regimes’’, have begun to receive attention
(Maki & McIlroy, 2005). This approach is based on
the assumption that neural control of volitional limb
movements differs in some fundamental ways to
reactions that are evoked by postural perturbation
(Maki & McIlroy, 1997). Thus, Maki et al. (2008)
argue that the most effective training programmes
should involve the use of perturbations. Recently,
Sakai and colleagues (Sakai, Shiba, Sato, & Taka-
hira, 2008) were able to show that a short-term
perturbation-based training programme on a tread-
mill significantly decreased postural sway in a cohort
of 45 community dwelling elderly individuals (mean
age 71 years).
Adaptive processes following balance training
in old age
The effects of balance training on strength perfor-
mance and reflex activity have specifically been
investigated in young populations (Gruber & Goll-
hofer, 2004; Taube et al., 2007). It is suggested that
neural mechanisms responsible for the increase in
explosive force production capacity due to training
could occur within the muscle (intramuscular adap-
tation) or within muscle groups (intermuscular
adaptation) (Gruber et al., 2007).
Gruber and Gollhofer (2004) investigated the
impact of 4 weeks of balance training on the
explosive strength qualities of the leg extensor
muscles during maximum isometric actions in a
cohort of young participants (mean age 28 years).
Balance training resulted in an increase in explosive
force production capacity that was paralleled by an
enhanced neuromuscular activation of the same
muscles. The authors suggested that an extra-facil-
itatory drive from the afferent system in terms of a
reduced presynaptic inhibition of Ia afferents could
account for the improved muscle activation at the
beginning of force production.
Taube et al. (2007) investigated cortical and spinal
adaptations in young individuals (mean age 25 years)
following balance training by means of H-reflex
stimulation, transcranial magnetic stimulation
(TMS), and conditioning of the H-reflex by TMS.
After 4 weeks of balance training, the authors
observed an improved postural stability. During the
initial training phase, cortical excitability was high.
After training, the authors reported a decrease in
motor-evoked potentials during stance perturbation
on a treadmill. This could be taken to imply that
balance training induced changes in the regulation of
human erect posture in terms of a shift from cortical
 334
Table II. Studies of the impact of balance training on neuromuscular performance in elderly individuals

Age Type of Exercise Sessions per Strength Postural

Authors Sex (years) n training duration week gain control

U. Granacher et al.
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Judge et al. (1993) f ]62 21 GBT 24 weeks 3 Knee extension: PB0.001 (1-RM) Improved performance in: single leg
HRT Leg extension: P B0.001 (1-RM) stance,
P 0.023; forward lean test, PB0.05
Wolfson et al. (1996) m/f ]65 110 BT vs. 12 weeks 3 Summed lower extremity isokinetic Performance in single leg stance:
HRT vs. strength: balance training, n.s.; heavy Balance training, P B0.02; heavy resistance
BT resistance training, n.s.; BTHRT, PB0.02
HRT training, PB0.02; BTHRT: P B0.02 Performance in sensory organization test:
Balance training, P B0.02; heavy resistance
training, n.s.; BTHRT, n.s.
Wolf et al. (1997) m/f ]70 72 BT vs. 15 weeks 1 Balance training:
TC Improved performance in: perturbation
impulse,
P B0.0001; postural sway, P B0.007
Tai Chi training:
Performance in: perturbation impulse, n.s.;
postural sway, n.s.
Williams et al. (1997) m/f ]64 16 BT 10 weeks 2 Resulting plantar flexion strength during Decrease in reflex activity in
Achilles tendon reflex: PB0.05 gastrocnemius: PB0.05
Mynark and Koceja (2002) m/f ]65 20 PTT 2 days Improved performance in: postural sway,
P B0.05;
oleus H-reflex down-training,
P B0.05
Wu et al. (2002) m/f
55 39 TC 3 years 3 Concentric action knee extension: Decrease in postural sway in eyes open/
(608/s), P B0.013 closed
eccentric action knee extension: condition in antero-posterior and
(608/s;1208/s), P B0.013 medio-lateral direction (PB0.05)
Concentric action knee flexion:
(608/s), P 0.713
eccentric action knee flexion:
(608/s;1208/s), P B0.713
Steadman et al. (2003) m/f ]60 199 BT 24 weeks 2 Improved performance in: postural sway,
P B0.22;
Berg balance score, PB0.0001;
10-min timed walk test, PB0.001
Tsang and Hui-Chan (2004) m/f ]63 49 TC 8 weeks 6 Improved performance in: sensory
organization test,
P B0.001; limits of stability test,
P B0.013
Downloaded By: [Granacher, Urs] At: 07:24 19 November 2008

Table II (Continued)

Age Type of Exercise Sessions per Strength Postural

Authors Sex (years) n training duration week gain control

Tsang and Hui-Chan (2005)
Granacher et al. (2006)
Gatts and Woollacott
(2006, 2007)
Granacher and Gollhofer
(2006), Granacher
et al. (2007)
m/f ]60 48 TC 3 years 1 Concentric action knee extension: Performance in postural sway:
(308/s), P B0.004 Antero-posterior, PB0.411; medio-lateral,
Eccentric action knee extension: P B0.235
(308/s), P B0.049 Performance in perturbed single leg stance:
Concentric action knee flexion: Antero-posterior, PB0.000
(308/s), P B0.021
eccentric action knee flexion:
(308/s), P B0.007
m ]60 60 BT 12 weeks 3 Improved ability to compensate for
decelerating gait perturbations: P B0.05
m/f ]68 22 TC 3 weeks 5 Improved ability to compensate for
accelerating gait perturbations: PB0.004
Improved performance in: FRT:
p50.0001; right leg stance, P50.0007;
left leg
stance, P 50.0001; timed up and go test,
P 50.0001
m ]60 60 BT 12 weeks 3 Leg extension: P B0.01 (MVC) Improved performance in: FRT, p50.01;
Leg extension: P B0.01 (RFD) tandem walk test forward, P 50.01;
tandem

Exercise and fall prevention in seniors

walk test backwards, P50.01
Li et al. (2007) m/f
65 47 TC 48 weeks 1 Ankle extension: n.s. (heel rise score) Performance in: reaction time, n.s.; single
leg stance, n.s.; timed tandem walk
test, n.s.
Madureira et al. (2007) f
65 66 BT 48 weeks 1 Improved performance in: timed up
and go test, P B0.001; Berg balance
score, PB0.001
Nagy et al. (2007) m/f ]77 19 BT 8 weeks 2 Postural sway: longer sway path in
medio-lateral direction, PB0.05
Improved performance in: timed up
and go test, P B0.05
Rochat et al. (2008) m/f ]69 47 GBT 10 weeks 1 Improved performance in: gait speed,
P B0.03; stride length, PB0.56

Abbreviations: BTbalance training, TCTai Chi training, PTT perturbation training, GBTgait and balance training, n.s. non significant, FRTfunctional reach test.

335
 336 U. Granacher et al.
to subcortical areas. Thus, supraspinal rather than
spinal mechanisms seem to be responsible for the
training-induced postural improvement.
Little information is available about adaptive
mechanisms following balance training in old age.
Granacher et al. (2007) demonstrated an increased
rapid force production capacity of the leg extensors
following 12 weeks of balance training in a cohort of
elderly men (mean age 66 years). However, this was
not accompanied by enhanced neuromuscular acti-
vation of the same muscles. Therefore, it is suggested
that an improved intermuscular coordination ac-
counts for the enhanced explosive force production
capacity in elderly men following balance training,
whereas changes in the recruitment and firing rate of
motor units (intramuscular coordination) result in
the increased explosive force production in younger
individuals (Gruber & Gollhofer, 2004). This line of
argument is reinforced by the results of a study in
which the impact of balance training on elderly
individuals (mean age 80 years) was investigated in
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single-joint isokinetic movements of the lower ex-
tremities (Judge, Whipple, & Wolfson, 1994). The
authors did not observe any training-induced
changes in summed peak moment of hip, knee,
and ankle flexion/extension. Thus, the results of
Judge et al. (1994) and Granacher et al. (2007)
indicate that, in seniors, balance training mainly
affects strength performance in functional multi-
joint movements in terms of an improved intermus-
cular coordination.
In a recent study, Granacher et al. (2006) exam-
ined the effects of balance training on reflex activity
during gait perturbations in elderly men (mean age
66 years). Twelve weeks of progressive balance
training resulted in shorter onset latencies as well
as an increase in reflex activity in muscles compen-
sating for the decelerating perturbation impulse.
Gatts and Woollacott, (2006) investigated the neural
mechanisms underlying balance improvement fol-
lowing Tai Chi training in a cohort of elderly
balance-impaired seniors (mean age 77 years).
Three weeks of Tai Chi training had an impact on
the ability to compensate for accelerating perturba-
tion impulses while walking at a self-selected speed
in terms of shorter onset latencies and reduced
muscle co-contraction. Different sites within the
central nervous system could be responsible for the
observed results (e.g. proprioceptive, visual and
vestibular system). A detailed discussion of all these
sites is beyond the scope of this review. Therefore,
we will focus focus on possible adaptive processes
following balance training in the proprioceptive
system.
Granacher et al. (2006) suggest an enhanced
sensitivity of muscle spindles as the most likely
mechanism responsible for the improved dynamic
postural control. Indeed, there is evidence that
training induces metabolic changes in intrafusal
muscle fibres. Furthermore, it has been shown
previously that changes in sensitivity of the gamma
spindle drive are possible in old age following short-
term balance training (Mynark & Koceja, 2002).
However, further research is needed to determine
the exact neuromuscular sites responsible for the
improved strength performance and postural control
in old age following balance training.
Conclusion
In summary, new evidence has been gathered in
terms of the underlying neuromuscular mechanisms
responsible for the decline in strength performance
and postural control in old age. Recently, a decrease
in the number of satellite cells has been associated
with sarcopaenia
a desensitization of muscle
spindles with an impaired postural control. Resis-
tance training has the potential to increase the
number of satellite cells. Thus, muscle hypertrophy
can be induced even in elderly individuals. Balance
training seems to be able to enhance the sensitivity of
muscle spindles. This might lead to an improved
afferent sensory input, which again influences the
efferent muscular output in balance-threatening
circumstances. Therefore, strength and balance
training have the potential to attenuate or even
reverse deteriorations in old age. Interestingly, there
is now a growing body of scientific literature that
favours specifically designed strength or balance
training programmes. Therefore, it is suggested
that heavy resistance strength training and balance
training (traditional approach) or power training and
perturbation training (new approach) should be
applied in training programmes to successfully
counteract the impact of neuromuscular ageing and
the occurrence of falls. Further research is required
to confirm the adequacy of the new intervention
approach.
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