78 Faculty of Sport, University of Ljubljana, ISSN 1318-2269 Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
Nejc Šarabon TRANSCRANIAL MAGNETIC 
 STIMULATION OFFERS NEW 
 POSSIBILITIES FOR THE STUDY 
 OF MOTOR CONTROL 
 TRANSKRANIALNA MAGNETNA 
 STIMULACIJA ODPIRA NOVE 
 MOŽNOSTI ZA RAZISKOVANJE 
 KONTROLE GIBANJA
Abstract
Since its introduction in 1985, transcranial magnetic 
stimulation has been used as a tool for non-invasive 
exploration of motor control in humans. This paper 
reviews transcranial magnetic motor cortical stimu-
lation as a revolutionary research tool in neurophysi-
ology and its wider application. Historical facts on 
the development of magnetic nerve stimulation are 
first presented, followed by the basic physical and 
physiological mechanisms underlying this stimula-
tion. The core of the paper provides an overview of 
the methods and measurements of transcranial mag-
netic stimulation paradigms, such as (i) central mo-
tor conduction time, (ii) threshold and amplitude of 
the motor potentials evoked by transcranial stimula-
tion, (iii) input-output properties of the corticospinal 
tract, (iv) cortical brain mapping using transcranial 
stimulation, (v) cortical silent period, and (vi) intrac-
ortical inhibitory and facilitatory mechanisms tested 
by paired pulse paradigms. Finally, there is a dis-
cussion of the current knowledge of the application 
of transcranial magnetic stimulation in relation to 
neuromuscular potentiation, central fatigue, neural 
plasticity and sensorimotor integration. Proposals 
are put forward for potential novel applications of 
magnetic stimulation in sports science. 
Key words: motor control, brain stimulation, tran-
scranial magnetic stimulation, corticospinal excit-
ability, plasticity, central fatigue.
Faculty of Sport, University of Ljubljana, Slovenia
Corresponding author:
Faculty of Sport, University of Ljubljana
Gortanova 22, SI-1000 Ljubljana, Slovenia
Tel.: +386 1 5207787
Fax.: +386 1 520 77 50
E-mail: nejc.sarabon@sp.uni-lj.si
Izvleček
Transkranialna magnetna stimulacija se od izna-
jdbe leta 1985 uporablja kot neinvazivna metoda za 
raziskovanje kontrole gibanja pri človeku. Ta članek 
podaja pregled transkranialne stimulacije možganske 
motorične skorje, ki se kot inovativno raziskovalno 
orodje uporablja v nevrofiziologiji ter njej sorodnih 
raziskovalnih področjih. Najprej je navedenih nekaj 
zgodovinskih dejstev o razvoju magnetne stimulacije 
živcev, čemur sledi razlaga fizikalnih in fizioloških 
mehanizmov stimulacije. Opisana so anatomska 
področja in prevodne poti, ki jih s to stimulacijsko 
tehniko testiramo. Jedro članka predstavlja pregled 
najpogostejših merilnih tehnik, pri katerih uporablja-
mo transkranialno magnetno stimulacijo. Kratko so 
pojasnjeni: (i) centralni motorični prevodni čas; (ii) 
prag in amplituda motoričnih potencialov vzbujenih 
s transkranialno stimulacijo; (iii) vhodno-izhodne 
lastnosti kortikospinalnega trakta; (iv) določanje 
topografskih možganskih map z uporabo transk-
ranialne stimulacije, (v) obdobje tišine možganske 
skorje in; (vi) znotrajmožganska inhibicija in fa-
cilitacija, ki ju testiramo s tehniko parnih magnet-
nih dražljajev. Skozi živčno-mišično potenciacijo, 
centralno utrujenost, plastičnost živčnega sistema 
in senzomotorično integracijo, sklenemo uporabo 
transkranialne magnetne stimulacije na področju 
športne znanosti. Na koncu so podane nekatere sug-
estije za bodočo uporabo transkranialne magnetne 
stimulacije pri proučevanju kontrole gibanja. 
Ključne besede: kontrola gibanja, možganska stimu-
lacija, transkranialna magnetna stimulacija, ko-
rtikospinalna vzdražnost, plastičnost, centralna 
utrujenost.

Transcranial magnetic stimulation & sport science 79 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
Introduction
Motor evoked potentials (MEPs) are established as diagnostic tests and research tools, provid-
ing objective measure of function in their related neural processing systems and conducting 
tracts. In neurophysiology, peripheral muscle and nerve stimulation was followed by stimu-
lation of the central nervous system. Transcranial electrical stimulation (TES) (Merton & 
Morton, 1980) was preceded by direct stimulation of brain structures during surgery. TES, 
which uses high voltage electrical pulses of short duration applied to the scalp overlying the 
motor cortex, is an uncomfortable procedure, inappropriate for routine clinical use. In 1985 the 
technique of non-invasive transcranial magnetic stimulation (TMS) was introduced (Barker, 
Jalinous, & Freeston, 1985a), which led to a new era of research into motor control and cortical 
function. Since then, TMS has become a widely used diagnostic tool in neurophysiology, as 
attested by the volume of literature on the subject (for review see Bailey, Karhu, & Ilmoniemi, 
2001; Cantello, Tarletti, & Ciravdi, 2002; Hallett, 2000; Lisanby, Luber, Perera, & Sackeim, 
2000; Petersen, Pyndt, & Nielsen, 2003; Siebner, & Rothwell, 2003; Walsh, & Cowey, 2000; 
Weber, & Eisen, 2002). 
The purpose of this paper is to present the technical, anatomical and physiological basis 
of TMS. Measurements of the magnetically evoked motor potentials (MEPs) are described. 
Emphasis is placed on research applications of TMS, such as brain mapping, sensorimotor 
integration, intra-cortical excitatory and inhibitory mechanisms and neuroplasticity. Potential 
new applications in sport science are proposed. 
Basic principles and neurophysiology of TMS
The basic mechanism of nerve cell activation is identical for TES and TMS. Both techniques 
cause a change in the transmembrane potential, which, if the stimulus is strong enough, results 
in the initiation of a nerve action potential (Pascual-Leone, Davey, Rothwell, Wassermann, 
& Puri, 2002). However, apart from TES, TMS, based on the principle of electromagnetic 
induction, employs a pulse of magnetic field to cause current flow in the tissue. Switching the 
electrical current on or off in a primary (stimulation) coil will cause currents to be induced in 
a secondary coil placed near neural tissue, although the primary coil is not itself in electrical 
contact with target neural structures. The independence of electrical resistance of the skull 
enables non-invasive brain stimulation by TMS. For peripheral, spinal cord and transcranial 
magnetic stimulation, different shapes (circular, double circular, double cone), and sizes (Ø=50 
to 100 mm) of magnetic coils are available. The main difference between the double-circular 
or figure-of-eight coil and the standard circular coil is that the former has a more focal area 
of effect. 
When using TMS, it is necessary to be aware of the complexity of the structures and processes 
that are probed. Motor function in humans is served by several distinct, yet interconnected, 
neurofunctional components. For example, when voluntary movement of the hand is to be 
executed, numerous parallel, as well as sequential, integrative sensorimotor processes are 
involved. In the same way as perceptual skills reflect the ability of sensory systems to detect, 
analyse and differentiate certain physical stimuli, dexterity indicates the ability of the motor 
system to plan, co-ordinate and execute movements. In fact, these two systems are inseparable, 
operating in a complementary manner. All the three main categories of movements (reflexive, 

80 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
rhythmic and voluntary) are complexly organized within the brain and spinal cord. Planning 
and setting of voluntary movement are organized in the cerebral cortex, with different areas 
communicating with subcortical areas, the cerebellum and the spinal cord. The initiation of a 
voluntary movement is driven by pyramidal cells in the primary motor cortex. The output of 
the motor system through muscle activity however depends on parallel processes and states 
of other serial components of the corticospinal tract. TMS similarly activates only a restricted 
part of the motor system, with the evoked MEPs being influenced by other functional elements 
of the sensorimotor system. 
Pyramidal cells are not independent units that generate rudimentary neural activity autono-
mously, but rather are subject to excitatory and inhibitory input transmitted from other nerve 
cells. Different types of brain stimulation of the motor system are capable of activating py-
ramidal cells in different ways. Patton and Amassian (1954) were the first to measure direct 
and indirect volleys (D and I waves, respectively) following transcranial electrical stimulation 
using epidural electrodes (Figure 1). Lower intensities of stimulation evoked short latency D 
waves that were ascribed to direct activation of pyramidal cells at the axon hillock (Day et al., 
1987; 1989). Increasing the intensity of stimulation evoked I waves, which were attributed to 
indirect or transsynaptic activation of the same corticospinal neurons. A time gap of 1.5 to 2 
ms between D and I1 as well as between successive I2, I3, and I4 waves was observed.
Figure 1: Comparison of TMS (left) and TES (right). Magnetic stimulation with a figure-of-eight 
coil oriented in a posterior-to-anterior direction activates pyramidal neurons predominantly 
transsynaptically. This causes multiple descending indirect volleys (I
1
-I
4
) of the cervical spinal 
cord and longer latencies of the MEPs detected by the EMG electrodes. Conversely, TES, or 
lateral-to-medial orientation of the TMS, causes direct activation of the pyramidal cells at 
the axon hillock. The MEP latency is therefore shorter as a result of a predominantly direct 
descending volley (DW). 
Later studies (Amassian, Quirk, & Stewart, 1990; Burke et al., 1993; Di Lazzaro et al., 1999; 
Edgley, Eyre, Lemon, & Miller, 1990; 1997) have shown that TMS acts on the corticospinal 
neurons in a very similar way to high intensity electrical brain stimulation. Threshold in-

Transcranial magnetic stimulation & sport science 81 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
tensity TMS transsynaptically activates the motor cortex preferentially through excitatory 
cortical interneurons. Increasing the magnetic stimulation results in the appearance of D 
waves, which cause an approximate 2 ms shortening of the MEP latency. The orientation of 
the magnetic coil also affects the nature of activation and quantitative parameters of MEP 
(Brasil-Neto, Cohen, Panizza, Nilsson, Roth, & Hallett, 1992b; Mills, Boniface, & Schubert, 
1992; Pascual-Leone, Cohen, Brasil-Neto, & Hallett, 1994b). Using a figure-of-eight coil, the 
lateral-to-medial orientation preferentially evokes D waves, while the posterior-to-anterior coil 
orientation causes transsynaptic activation. Furthermore, TMS activates other local intrabrain 
circuits that could be tested by other stimulation paradigms such as paired pulse stimulation, 
which is described in a later section. 
The spinal motor neurons of the cord are the “final common pathway” of the motor sys-
tem, with which the higher centres and pyramidal cells make direct or indirect connections 
via multiple descending tracts (Bernhard & Bohm, 1954; Landgren, Philips, & Porter, 1962; 
Kuypers, 1981). Anatomical studies (Kuypers, 1981; Schoen, 1964) have demonstrated that 
the majority of corticospinal fibres terminate in the ventral horn, where the spinal motor 
neurons are located. Although the estimation of conduction velocities of different sections 
of the corticospinal pathway is possible by the application of either TMS at different levels 
(Gandevia, Petersen, Butler, & Taylor, 1999; Nielsen, Petersen, Deutschl, & Bellegaard, 1993; 
Ugawa et al., 1991) or by measuring descending volleys evoked by TMS at different levels of 
the spinal cord (Burke, Hicks, Gandevia, Stephen, Woodforth, & Crawford, 1993; Di Lazzaro 
et al., 1998; Rothwell et al., 1994), this estimation is too imprecise to draw firm conclusions 
about the activated pathway. 
Supportive evidence for the monosynaptic nature of the TMS-activated corticospinal projec-
tions comes from single motor unit studies considering their firing probability. Although 
reports on short rising times in post-stimulus time histograms are in favour of monosynaptic 
projections, no general agreement has been reached on this issue. The publications available 
(Burke, Gracies, Mazevet, Meunier, & Pierrot-Deseilligny, 1992, 1994; Burke, Gracies, Meu-
nier, & Pierrot-Deseilligny, 1992; Nicolas, Marchand-Pauvert, Burke, & Pierrot-Deseilligny, 
2001; Pierrot-Deseilligny, 1996; Pierrot-Deseilligny, & Marchand-Pauvert, 2002) provide neu-
rophysiological evidence for the possible existence of a cervical propriospinal system, which 
is postulated to play a mediating role in the transmission of descending motor commands to 
cervical motor neurons. 
TMS is much more than just a tool for testing the continuity of the corticospinal system. Since 
its introduction, it has been developed and many different diagnostic applications have been 
established. In the following section of this paper methods and measurements that employ 
motor cortex single- and double-pulse TMS paradigms will be presented. Consideration of 
other types of magnetic stimulation (high-frequency train TMS, and single TMS of other 
neuroanatomical regions) would make this review too large.
Methods and measurements 
Central motor conduction time
The majority of today’s clinical neurophysiology laboratories routinely apply TMS as part of 
their diagnostic procedure. Clinicians are mainly interested in central and peripheral con-

82 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
duction times, which can provide valuable insight into pathologically changed processes of 
neural signal propagation. 
The time between TMS of the motor cortex and the EMG response in the target muscle is the 
result of the propagation of an action potential along the whole motor pathway – from the 
cortex to the spinal segment, synaptic transmission to spinal motor neurons, along peripheral 
nerves from the spinal cord to the muscle and neuromuscular synaptic transmission. Com-
binations of electrophysiological measures enable independent estimation of the central and 
peripheral motor conduction times. To calculate the central motor conduction time, conduc-
tion in the peripheral segment of the motor pathway is estimated and then subtracted from 
the onset latency of the TMS-elicited MEP. To measure peripheral motor delays, F waves or 
direct stimulation of the efferent roots and nerves over the spinal columns are mainly used 
(Barker, Jalinous, Freeston, & Jarratt, 1985b; Merton, Morton, Hill, & Marsden, 1982; Rossini, 
Marciani, Caramia, Roma, & Zarola, 1985; Zarola et al., 1989). Calculations of the central 
conduction time from root stimulation will give slightly higher values due to the distance 
between the anterior horn synapse and the point of stimulation (Maccabee, Amassian, Eberle, 
& Rudell, 1993). 
During contractions, MEPs are both larger in amplitude and shorter in latency. The main 
shifts occur at the beginning of force gradation, with no further significant latency change 
being observed with volitional contraction above 10% of the maximal voluntary force (Ross-
ini, Caramia, & Zarola, 1987). This contraction-dependent time change shortens in the proc-
ess of biological maturation and reaches adult values during adolescence (Caramia, Desiato, 
Cicinelli, Iani, & Rossini, 1993). There is only a weak correlation between age and the central 
conduction time (Eisen, Shytbel, Murphy, & Hoirch, 1990; Mills & Nithi, 1997a). Values of 
central motor conduction times in women are slightly lower than those in men, due to the 
differences in height and arm length (Furby, Bourriez, Jacquesson, Mounier-Vehier, & Guieu, 
1992; Mills & Nithi, 1997b). Clinical studies provide evidence for prolonged central conduc-
tion times under pathological conditions such as stroke (Abbruzzese, Morena, Dall’Agata, 
Abbruzzese, & Favale, 1991), multiple sclerosis (Rossini et al., 1987) and upper motor neuron 
diseases (Barardelli, Inghilleri, Craccu, Mercuri, & Manfredi, 1991).
Threshold and amplitude of the MEP
In neurophysiology, a threshold represents the minimal stimulus required to provoke a defined 
response (Reid, Chiappa, & Cros, 2002). The cortical motor threshold to magnetic stimulation 
is usually defined as the stimulus intensity required to elicit reproducible responses of 50 to 
100 µV in about 50% of 10 to 20 consecutive trials (Rothwell et al., 1999). According to the two 
ma i n recom mend at ions , t h reshold i ntensit y c a n be ach ieved by eit her i nc rea si ng or dec rea si ng 
(Rossini et al., 1994; Mills & Nithi, 1997b) the stimulation in steps of 1 to 5% of the maximum 
output intensity. However, when MEPs are recorded from pre-activated muscle, the required 
amplitude level ranges from 200 to 300 µV. 
The TMS threshold is a complex measure that is influenced not only by the excitability of 
corticospinal cells, but also by the activity of the cortical cells that project to them, as well as 
by the excitability of spinal motor neurons, through which motor signals are conveyed to the 
periphery (Morita et al., 2000). Therefore, analytical approaches should be applied to selectively 
identify excitability levels at the intracortical, corticospinal, spinal and peripheral levels. In 
adults, the threshold is independent of age and gender. It is lowest for the hand muscles and 

Transcranial magnetic stimulation & sport science 83 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
highest for more proximal muscles (Mills & Nithi, 1997b; Wassermann, McShane, Hallett, & 
Cohen, 1992). 
The set-up for TMS conventionally employs electromyography for detecting evoked muscle 
responses. In acquired MEPs, amplitudes and rectified areas are the most common excitability 
parameters analysed (Figure 2). Either absolute or normalized values can be used. The maxi-
mal compound motor action potential, the M wave, is used for the purpose of normalization. 
Because of the variability of MEPs detected by the surface EMG, it is necessary to use averaged 
responses for final analyses. Statistical standards for reliable MEPs are available (Brasil-Neto, 
McShane, Fuhr, Hallett, & Cohen, 1992a), however, the majority of researchers have used 
fewer (4 to 10) consecutive magnetic stimuli to compose the final MEP (Stedman, Davey, & 
Ellaway, 1998; Thickbroom, Byrnes, & Mastaglia, 1999).  Although factors such as changes in 
coil positioning (Bohning, Denslow, Bohning, Walker, & George, 2003), background voluntary 
contractions, or phases of the cardiac or respiratory cycles should be considered, they are not 
of vital importance for the reproducibility of MEPs (Amassian, Cracco, & Maccabee, 1989). 
Recent studies employing the triple stimulation technique (Magistris, Rösler, Truffert, & My-
ers, 1998; Magistris, Rösler, Truffert, Landis, & Hess, 1999) have confirmed that the major 
MEP variability arises from phase cancellations. Additionally, evidence has been provided 
for the ability of a high intensity TMS to completely activate the goal muscle. This combined 
type of stimulation also has some pitfalls that prevent it simply replacing the conventional 
TMS application. 
Fig u re 2 : The avera ge response of he fi rst dors a l i nteros seu s mu scle of t he ha nd composed of 20 
consecutive magnetic stimuli. The most frequently measured parameters of the transcranially 
elicited MEPs include latency, area, and peak-to-peak amplitude.
Input-output curves
Spontaneous physiological oscillations in motor neuron excitability at both the cortical and 
spinal levels may play a role in fluctuations in the response size, while overall corticospinal 

84 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
excitability may vary as a function of time. For further improvement in metric characteris-
tics of MEPs, TMS is applied at different intensities and an input-output curve is composed. 
This was first performed by Devane, Lavoie, and Capaday (1997) in a report on the stimulus-
response properties of TMS. They found the relationship to be sigmoidal and thus strongly 
non-linear (Figure 3). Additionally, it was independent of the behaviour of a single motor 
neuron. The most striking result was that the steepness, i.e. the gain of the relation, changes 
as a function of the level of motor activity of the same qualitative task. This output parameter, 
as well as the others measured in TMS input-output studies (Carroll, Riek, & Carson, 2001; 
Devane et al., 1997; Ridding & Rothwell, 1997), most likely reflects the excitability properties 
of the population of cortical interneurons, corticospinal neurons, and the motor neuron pool 
with interneuronal relays. 
Figure 3: The TMS input-output curve for the first dorsal interosseus muscle. The following 
parameters are used as measures of corticospinal excitability: MT – measured threshold, CT 
– threshold calculated from the function, XI – x intercept of the tangent, MEP
max
 – maximal 
MEP defined by the function, S
50
 – stimulus intensity at which MEP size is 50% of the MEP
max
, 
k – slope parameter of the function, k
max
 – maximal slope of the function. MEP amplitudes 
are expressed as percentages of the maximal M wave (MW
max
).  
To compose the function technically, MEP amplitudes are plotted against the stimulus inten-
sity, and the data fitted with the Boltzmann equation:
MEP(s) =    
MEP max
 (Equation 1)
                 
1 + e
m(S
50
 – S)
where MEP
max
 is the maximum MEP defined by the function; m, the slope parameter of the 
function; S
50
, the stimulus intensity at which the MEP size is 50% of the maximal MEP. Be-
sides the slope parameter, which is usually taken as a general measure of the excitability of 

Transcranial magnetic stimulation & sport science 85 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
the pathway, a peak slope, which occurs at a stimulus intensity equal to S
50
, is defined by the 
relationship:
m* MEP max
        4         
. (Equation 2)
The purpose of fitting a mathematical function to the experimentally observed relationship 
between stimulus intensity and response magnitude is to provide a succinct description of 
the input-output properties of the corticospinal pathway (Carroll et al., 2001). The sigmoidal 
shape of the input-output relation is most likely due to a combination of factors. The experi-
mentally measured curve, at least in part, reflects the well-known fact (Henneman, 1957) that 
stimuli of increasing strength recruit motor neurons with increasing motor unit potentials. 
Namely, sigmoidal input-output relations occur in systems composed of excitable elements 
with a wide distribution of spike amplitudes (Erlanger, & Gasser, 1937; Rall, 1955). Other 
relevant factors are related to modulating influences arising from the properties of the neural 
circuitry. Multiple components of the corticospinal volley and perhaps a greater tendency for 
synchronization of single motor unit discharges with increasing stimulus strength (Poliakov 
& Miles, 1994) additionally contribute to the non-linear input-output properties. Thus, all the 
parameters of the sigmoidal function are complex in origin. For example, the plateau value of 
the function is probably not the maximal response to a purely excitatory corticospinal volley. 
Inversely, it represents the balance between excitatory and inhibitory components, including 
recurrent inhibition of later-recruited motor neurons by those recruited earlier. However, 
test-retest reliability assessment of the input-output parameters of the corticospinal pathway 
has shown that TMS can be used in long-term studies (Carroll et al., 2001). Nevertheless, such 
repeatability does not apply to classical single-intensity TMS applications.
TMS brain mapping
Since the first brain mappings of the sensorimotor system (Penfield, & Boldrey, 1937; Penfield, 
& Rasmussen, 1950), several studies using transcranial electrical and magnetic stimulation, 
positron emission tomography or functional magnetic resonance imaging have confirmed the 
existence of functional representations in the brain. The motor cortex is organized in terms 
of movements rather than muscles, including complex convergence and divergence. Different 
regions of the brain are stimulated in TMS mapping of cortical motor areas; the consecutive 
motor effects are quantified and related to the stimulated scalp site (Thickbroom et al., 1999; 
Weber & Eisen, 2002). While the need for spatial resolution of the stimuli has been empha-
sised, the figure-eight coil is generally employed. The coil is systematically moved over the 
scalp using a co-ordinate system referenced to the vertex (Cohen et al., 1991; Wassermann et 
al., 1992). To achieve focal activation stimulation should be of low intensity, which however, 
increases the amplitude variability of MEP (Kiers, Cros, Chiappa, & Fang, 1993). In addition, 
there is no firm rule regarding coil positioning, although obvious discrepancies between an-
terior-posterior and lateral-medial orientations have been proved (Mills et al., 1992; Werhahn 
et al., 1994; Wilson, Day, Thickbroom, & Mastaglia, 1996). 
A clear definition of TMS will provide a useful map of sites on the scalp from which responses 
can be obtained for each muscle of interest. The two most important parameters of such maps 
are the amplitude-weighted centre of the map (centre of gravity) and the point of maximum 
response. The area of the map is more difficult to estimate because of the non-focal nature 

86 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
of the stimulation, especially when higher intensities are used (Levy, Amassian, Schmid, & 
Jungreis, 1991) or target muscles are voluntarily pre-contracted (Nielsen, 1994), although these 
conditions give higher MEP amplitude consistency. 
An alternative to the use of the standardized TMS intensity of 1.2 motor threshold or a stimula-
tion output 20% above the threshold intensity (Wassermann et al., 1992) was provided by Clas-
sen et al. (1998), who mapped the thresholds for evoking MEPs at particular scalp locations.
Cortical silent period
Inhibitory neural systems are essential for shaping and modelling of the flow of excitatory 
signals and maintaining synaptic stability. In addition to the excitatory response, TMS also 
has some inhibitory effects. When TMS is applied during voluntary muscle contraction, MEP 
is followed by a pause in the electromyographic activity, known as the cortical silent period 
(Ahonen, Jehkonen, Dastidar, Molnar, & Häkkinen, 1998; Bertasi, Bertolasi, Frasson, & Pri-
ori, 2000) (Figure 4). The early phase (0 to 50 ms) of this phenomenon is ascribed to spinal 
inhibitory mechanisms, while the later phase is mainly due to the activation of cortical in-
terneurons acting over the neural output elements of the motor cortex (Ziemann, Lonnecker, 
Steinhoff, & Paulus, 1996a). These implicated interneuronal circuits are mediated by the in-
hibitory neurotransmitter GABA (Inghillieri, Berardelli, Cruccu, & Manfredi, 1993). There 
is also evidence that the silent period is a separate process to MEP (Wilson, Thickbroom, & 
Mastaglia, 1995).
F i g u r e 4 : Th e c o r t i c a l s i l e n t p e r i o d ( C S P ) o c c u r s w h e n m a g n e t i c s t i m u l a t i o n o f t h e c o r r e s p o n d i n g 
motor cortex is applied during an ongoing voluntary contraction. The potentiated MEP is 
followed by depression of the electrical muscle activity. After 80 to 300 ms the voluntary EMG 
signal rebounds.

Transcranial magnetic stimulation & sport science 87 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
A cortical silent period can last up to 300 ms and is basically divided into three phases: (i) the 
early phase that comprises approximately the first 50 ms after the MEP; (ii) the intermedi-
ate phase between 50 and 75 ms, and (iii) the late phase, more than 75 ms after the response. 
Increasing the intensity of TMS does not prolong the duration of the silent period after a 
plateau has been reached (Inghillieri et al., 1993). It has also been shown that the strength of 
the background contraction does not affect the silent period (Roick, von Giesen, & Benecke, 
1993; Uozumi, Ito, Tsuji, & Murai, 1992), hence, proprioceptive mechanisms are unlikely to 
be involved in the generation of this phenomenon.
Paired TMS 
Different combinations of two consecutive magnetic stimuli can provide a better understand-
ing of the neural processes that are part of voluntary movement control. By modifying the 
intensities of the two stimuli and the time delay between them, cortical and corticospinal 
excitatory and inhibitory mechanisms can be probed. The following techniques are described 
in the literature (i) the paired pulse technique, using two equal suprathreshold stimuli at in-
terstimulus intervals (ISIs) of 10 to 250 ms (Claus, Weis, Jahnke, Plewe, & Bruhnholzl, 1992; 
Valls-Sole, Pascual-Leone, Wassemann, & Hallett, 1992), (ii) the paired pulse technique, which 
employs a subthreshold conditioning stimulus followed by the suprathreshold test stimulus 
at 1 to 20 ms ISIs (Kujirai et al., 1993; Rothwell et al., 1991a; Ziemann, Rothwell, & Ridding, 
1996b), (iii) the paired pulse technique, using an initial suprathreshold stimulus followed by 
a subthreshold stimulus at 0.5 to 6 ms ISIs (Tergau, Ziemann, Hildebrandt, & Paulus, 1997; 
Ziemann, Tergau, Wischer, Hildebrandt, & Paulus, 1998) and finally; (iv) the paired pulse 
techniques, in which the first and second stimuli are applied through different stimulating 
coils at separate stimulation sites (Ferbert et al., 1992; Ugawa, Uesaka, Terao, Hanajima, & 
Kanazawa, 1995). 
The general aim of all these protocols is to evaluate the effects of the conditioning stimulus 
on the size of the MEP elicited by the test stimulus as a function of ISI duration and stimulus 
intensity. If the test TMS is preceded by suprathreshold conditioning, facilitation of the control 
responses occurs at 10 to 40 ms ISIs, but inhibition at 50 to 200 ms ISIs (Roick et al., 1993; 
Triggs et al., 1993). This technique employs stimuli of equal intensity in the range from 120 to 
160% of the resting motor threshold. Another method is to use a stimulus of only 80% of the 
threshold intensity to condition the test stimulus that follows, resulting in maximum inhibi-
tion at 1 to 5 ms ISIs and significant faci litation of MEPs at delays of 8 to 25 ms (Fig ure 5). Such 
interactions, which are universal for all muscles (Chen et al., 1998), are however dependent on 
coi l or ient at ion, st i mu lu s i ntensit y, a nd back g rou nd mu scle ac t iv it y. Fi na l ly, ver y shor t ISIs a re 
used in paradigms that combine the suprathreshold test stimulus followed by a conditioning 
subthreshold stimulus (Ziemann et al., 1998). The behaviour of MEPs under such conditions 
shows three facilitatory peaks at ISIs of 1.1 to 1.5 ms, 2.3 to 2.9 ms and 4.1 to 4.4 ms. 
It is again obvious that the interaction between the effects of two stimuli could take place 
at the cortical, subcortical or spinal level. However, there is evidence that they are primarily 
caused by intracortical mechanisms. In all three listed stimulation techniques, researchers 
used complementary protocols to determine the exact sites and mode of interactions. They (i) 
used different intensities of TMS, (ii) systematically changed the coil orientation and replaced 
conditioning TMS with transcranial electrical stimulation, (iii) studied the effects of TMS on 

88 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
spinal excitability, and (iv) intraoperatively recorded descending volleys with epidural elec-
trodes, and used neuroactive drugs. For the technique using two suprathreshold magnetic 
stimuli it is proposed that facilitation followed by a long-lasting (150-300 ms) inhibition of 
pyramidal cells is derived from supraspinal sites. Intracortical inhibition and facilitation, 
tested by the test stimulus preceded by a subthreshold stimulus, are also confirmed to be of 
cortical origin. Moreover, there is evidence that these neural characteristics are due to separate 
mechanisms, contradicting the view that facilitation is merely a rebound following inhibition. 
Finally, the interaction of the subthreshold stimulus that follows the suprathreshold stimulus 
is very likely to test the excitability of motor cortex circuits responsible for the generation of 
I waves.
The main and most commonly used measures of TMS have been presented. Undoubtedly, it 
is important to be aware of potential side effects of the technique. Single-pulse (<1 Hz) and 
paired-pulse TMS have been proved safe for use in normal subjects. However, it is interest-
Figure 5: An example of the paired-pulse technique using subthreshold (80% MT) conditioning 
followed by the suprathreshold (130% MT) test stimulus. Averages of 20 consecutive MEPs 
are constructed for the control conditions, intracortical facilitation (ICF), and intracortical 
inhibition (ICI). The delays between the stimulation artefacts (left side of the traces) show the 
duration of the time interval between the two transcranial magnetic stimuli. ISIs of 1 to 5 ms 
result in ICI, while ISIs of 8 to 20 ms produce ICF. ICI is normally more prominent than ICF, 
as is also seen in this representative subject.

Transcranial magnetic stimulation & sport science 89 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
ing that early reports on safety measures are from studies in humans (Bridgers & Delaney, 
1989). Most of the research using animal models, which was conducted thereafter (Ravnborg, 
Knudsen, & Blinkenberg, 1990; Counter, 1993; Fleischmann, et al., 1996), examined the his-
tological changes induced by prolonged exposure to TMS. Their only consistent finding was 
increased catecholaminergic activity following TMS in rats which had received direct deep 
brain stimulation because of their small head size. However, evidence on the depth of brain 
stimulation in human (Rudiak & Marg, 1994) argues against the possibility that this type of 
response could also occur in humans. 
After exposure to prolonged single-pulse TMS, subjects have shown a transient increase in the 
auditory threshold (Pascual-Leone et al., 1992), hence the use of earplugs during TMS testing 
is recommended. Apart from its use in healthy subjects, single-pulse TMS has been reported 
to induce seizures in patients with large cerebral infarcts (Hömberg & Netz, 1989) as well as 
in epileptics (Classen et al., 1995; Düzel, Hufnagel, Helmstaedter, & Elger, 1996). 
In summary, TMS does not appear to cause long-term adverse neurological, hormonal, motor, 
sensory or cognitive effects in healthy subjects. Current concepts even propose that Insti-
tutional Review Boards consider well-designed studies using single- and paired-pulse TMS 
protocol s a s bei ng of m i n i ma l r isk to ch i ld ren (Gi lber t et a l ., 20 0 4 ; L i n & Pa sc u a l-L eone, 20 02). 
For further insight into TMS safety measures, including those of high-frequency repetitive 
TMS, the reader is referred to review articles by Anand and Hotson (2002), Bridgers (1991) 
and Wassermann (2000). 
Possibilities for the use of TMS in sport science 
TMS lends itself to many different research areas of particular interest to the sport science 
disciplines. Kinesiology-related research issues for which TMS has already been, or could 
potentially be applied, include: (i) post-exercise facilitation, (ii) central fatigue, (iii) sensori-
motor integration and co-ordination, and (iv) neuronal plasticity. In the final section, some 
light is shed on applied studies that consider neurocontrol mechanisms in situations akin to 
those in sporting activities. Further proposals for the use of TMS in sport science research 
are put forward. 
When a subject performs a weak voluntary contraction of a muscle, the corticospinal pathway 
to that muscle is facilitated (Rothwell, Thompson, Day, Boyd, & Marsden, 1991b). As already 
mentioned, most facilitatory effects occur at low contraction forces (Lim & Yiannikas, 1992; 
Stedman, Davey, & Ellaway, 1998). Additionally, higher MEP potentiation has been reported 
for precision movements as compared to general grip tasks (Datta, Harrison, & Stephens, 
1989), presumably because of the larger involvement of pyramidal tract neurons in such tasks 
(Muir, & Lemon, 1983). A spread of facilitatory effects has also been shown during a voluntary 
contraction of neighbouring ipsilateral or homonymous contralateral muscles (Chiappa et al., 
1991; Hufnagel, Jaeger, & Elger, 1990), but other reports give opposing results (Samii, Canos, 
Ikoma, Mercuri, & Hallett, 1997).
Besides the immediate influence of the voluntary motor activity, there are also prolonged post-
exercise MEP potentiation effects (Brasil-Neto, Cohen, & Hallett, 1994; Samii, Wassermann, 
Ikoma, Mercuri, & Hallett, 1996). These authors found that 10-second activation could lead 
to post-exercise facilitation, which decayed to the baseline over 2 to 4 minutes. Researchers 

90 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
have postulated that these effects are due to intracortical plastic changes, as they were not 
present after transcranial electrical stimulation. It has been shown (Balbi, Perretti, Sannino, 
Marcantonio, & Santoro, 2002) that the intensity of muscle contraction does not play a sig-
nificant role in eliciting post-exercise facilitation. However, the maximum increase in MEP 
was present at the shortest and strongest muscle contractions. 
Acute potentiation manoeuvres are incorporated into sport training and competition rou-
tines. These short-term conditioning procedures include: (i) explosive strength exercises are 
performed prior to speed and agility training, (ii) joint stability exercises are combined with 
foot work co-ordination, (iii) pliometrics is carried out just before sprint starts, and (iv) in re-
sistance training, maximal and submaximal loads are combined in the same training session. 
Although peripheral electrophysiological and mechanical measures have been well evidenced 
(Enoka 1994), there are few studies that explain the acute response of the CNS to such active 
manipulations. Therefore, TMS could be effectively used to elucidate the precise nature of such 
procedures and possibly help in their optimisation.  
If a subject performs many consecutive muscle contractions or sustains a constant muscle 
contraction for a longer time, this will result in depression of MEP responses (Sacco, Thick-
broom, Byrnes, & Mastaglia, 2000) (Figure 6). During prolonged muscle activity a reduction 
in voluntary activation can significantly contribute to muscle fatigue, a phenomenon termed 
Figure 6: Mean relative changes (10 subjects) in evoked responses (+SEM) for the biceps brachii 
muscle after a 60-second maximum voluntary contraction. The pre-exercise control range 
is shaded. The motor evoked potentials (MEPs) elicited by TMS are significantly depressed 
from the third minute onwards (filled circles, P<.05). Maximal muscle potentials evoked by 
the peripheral nerve point stimulation (MW) remain unchanged. (Adapted after Sacco et al., 
2000).

Transcranial magnetic stimulation & sport science 91 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
central fatigue (Gandevia, Allen, & McKinzie, 1995; Gandevia, Allen, Butler, & Taylor, 1996; 
Löscher, Cresswell, & Thorstensson, 1996). In the past fifteen years some investigators have 
used TMS in relation to corticospinal mechanisms of fatigue. Central fatigue and motor cor-
tical excitability have been studied during different types of muscle contractions (Löscher & 
Nordlund, 2002). During fatiguing isometric contractions there is an increase in MEP size and 
a concomitant increase in the duration of the silent period (McKay, Stokic, Sherwood, Vrbova, 
& Dimitrijevic, 1996; Sacco, Thickbroom, Thompson, & Mastaglia, 1997; Taylor, Butler, Allen, 
& Gandevia, 1996), suggesting a simultaneous increase in excitability levels of inhibitory and 
facilitatory neural structures. Thus, the relationship ratio between the two systems determines 
the actual state of the corticospinal tract and in turn modulation of MEPs during and after 
muscle activity. It seems that plastic changes at cortical sites are responsible for the changes 
in MEP, since there are no significant changes at spinal and cortical levels.
There are fewer reports on TMS and fatigue in sports-specific motor activities. The first study to 
show the possible use of TMS in sports and various kinds of everyday exercise was undertaken 
by Hollge et al. (1997). They evaluated changes in MEP size and central motor conduction time 
after various, predominantly aerobic (climbing stairs and jogging), and anaerobic (press-ups, 
dumb-bell holding, and 400 m run) exercises. Exhausting strength exercises resulted in a sig-
nificant reduction in MEP amplitudes, while no significant change was elicited by aerobic exer-
cises. In addition, Tergau et al. (2000) investigated the modulation of intracortical mechanisms 
(ICI and ICF) following pull-up exercise until exhaustion. After exercise, ICF was significantly 
reduced, while ICI remained unchanged. ICF changes were limited to the muscles performing 
the task and tended to normalize within 8 min. Both the baseline ICF level and ICF reduction 
after exercise were significantly correlated (r = 0.63 and r = 0.73 respectively) with the overall 
lifting work accomplished by the subject before complete exhaustion. Another study (Fulton, 
Strutton, McGregor, & Davey, 2002) examined fatigue-induced change in the corticospinal 
drive to back muscles in elite rowers compared to an untrained group. Each subject completed 
two exercise protocols on a rowing ergometer (i) light rowing at submaximal intensity for 
10 min and (ii) intense rowing at maximal output for 1 min. The non-rowers showed a brief 
facilitation of MEPs 2 min after light and intense exercise that was only present in the elite 
rowers after intense exercise. In the period 4 to 16 min after the light exercise the mean MEP 
amplitude was less depressed (relative to pre-exercise values) in elite rowers than in controls 
(mean 79.4 and 60.9% respectively). The authors propose that the differences are due to the 
energy requirements being closer to maximum capacity in non-rowers, therefore resulting 
in more fatigue. This notion was supported by the lack of any difference between the groups 
following intense exercise during which both groups worked at their own maximum. 
To understand mechanisms responsible for neuromuscular fatigue, TMS should be further 
employed to applied sports research protocols. Different TMS techniques may be useful tools 
in training supervision as well as in the detection of overtraining situations. Although the 
above-mentioned studies suggest that aerobic exercises are not capable of causing MEP modu-
lation, it should be pointed out that only low to moderate protocols were used. It would be 
interesting to investigate the effects of extreme situations, such as marathon or triathlon racing. 
Moreover, issues related to the arrangement of the amount and types of training during dif-
ferent training periods need to be examined. In an attempt to evoke the highest possible body 
adaptations, athletes and their trainers sometimes exceed the ability of the human body to 
regenerate. Accumulation of such catabolic loads can lead to the overtraining syndrome. Until 

92 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
the present, biochemical (Maso, Lac, Filaire, Michaux, & Robert, 2004; Uusitalo et al., 2004) 
and psychological (Maso et al., 2004) methods have been mainly employed to better diagnose 
and understand overextending and pathological central fatigue (for reviews see Armstrong, & 
VanHeest, 2002; Pearce, 2002; Urhausen & Kindermann, 2002; Petibois, Cazorla, Poortmans, 
& Deleris, 2003). TMS could be tested for its potential use in the detection and prevention of 
overloading and the resulting overtraining. As the overtraining syndrome is postulated to be 
of CNS origin, correlates with MEP parameters could be expected.
A considerable amount of work has been done in the field of neuronal plasticity, with tran-
scranial stimulation being used as a diagnostic tool (for review see Cohen et al., 1998; Sieb-
ner & Rothwell, 2003). As a still loosely defined term, neural plasticity describes the ability 
of the brain to change. Cortical plasticity encompasses a wide variety of phenomena and 
mechanisms, including modifications of cortical properties such as the strength of internal 
connections or representational patterns, or neuronal modifications, either morphological or 
functional (Donoghue et al., 1996). The mechanisms of such reorganization can be studied at 
a synaptic, cellular or regional level (Buonamano & Merzenich, 1998). The latter encompasses 
changes in the response of larger cell assemblies following prolonged input changes induced 
by different types of manipulations. In fact, all the measures of TMS can be used in the as-
sessment of such modifications. By employing TMS, neuroplasticity has been documented as 
a result of (i) short-term deafferentation (Ridding & Rothwell, 1995; Brasil-Neto et al., 1993); 
(ii) long-term deafferentation (Hall, Flament, Fraser, & Lemon, 1990); (iii) implicit and explicit 
learning (Pascual-Leone, Grafman, & Hallett, 1994a); (iv) use of the dependent plasticity of 
motor cortical representations (Classen et al., 1998) and; (v) some other pathophysiological 
conditions (Hallett, 2001). 
The acquisition of new motor skills is also associated with plastic changes in the controlling 
neural system. Learning to activate a synergistic combination of muscles in a new way will 
cause changes in cortical representations after a period of physical training (Cohen, Gerloff, 
I koma , & Ha l let t , 19 95). Neu r a l pla st ic it y i s t hu s a n u nderly i ng mecha n i sm for motor lea r n i ng. 
An understanding of the patterns, mechanisms and functional relevance of cortical plasticity 
will hopefully lead to the design of effective strategies to enhance plasticity when beneficial, 
and to downregulate it when it is maladaptive (Pons, 1998).
Animal studies have suggested that the repetitive execution of simple or well-learned move-
ments has little impact on the organisation of the motor cortex (Plautz, Milliken, & Nudo, 
2000). Comparisons between pre-training and post-training maps of M1 movement repre-
sentations revealed no task-related changes in the cortical area devoted to representation of 
individual distal forelimb movements. They concluded that repetitive motor activity alone does 
not produce functional reorganization of cortical maps. Instead, they propose that motor skill 
acquisition, or motor learning, is a prerequisite for driving representational plasticity in M1. 
Another study (Pearce, Thickbroom, Byrnes, & Mastaglia, 2000) investigated the character-
istics of the corticospinal tract measured on the FDI in a group of elite racquet players. The 
major findings of the study were an increase in the corticomotor excitability of the playing 
hand and changes in the topography of the cortical motor map for the playing hand, which 
were not seen in a control group.
An even more specifically sports training oriented study (Carroll, Riek, & Carson, 2002) used 
TMS to test central vs. spinal neural adaptations to resistance training in humans. The results 

Transcranial magnetic stimulation & sport science 93 Kinesiologia Slovenica, 10, 2, 78–104 (2004)
of their experiment demonstrate that resistance training alters functional properties of the 
corticospinal pathway in humans, but failed to show substantial cortical effects. Additionally, 
their data suggest that the reorganisation of the corticospinal motor pathway that occurs in 
response to the repetitive execution of simple movements against a large resistance is inde-
pendent of that which occurs during motor learning. This idea was further evidenced by Lun-
dbye, Marstrand, and Nielsen (2003), who also demonstrated different neural adaptations as a 
response to strength and visual-motor skill task training respectively. TMS stimulus-response 
curves showed increased corticospinal excitability after 4 weeks of hand skill practice. 
There are several other interesting research issues of physical long-term training in which 
TMS could be used for diagnostic purposes. Athletes from various sport disciplines could be 
compared for different neurocontrol parameters that would reveal mechanisms typical for 
specifically trained groups. It could be assumed that disciplines that are based on different 
motor abilities (strength, power, speed, precision, postural stability, etc.) require specific motor 
control that may be reflected in input-output properties of the central and peripheral nervous 
systems. Using the model of Pearce et al. (2000) it is possible to analyse other specific elite ath-
letes in sports such as (i) soccer vs. basketball (upper and lower extremity skilled movements), 
(ii) ballet vs. sprint (posture maintenance and pliometric activity), (iii) dart throw vs. shot put 
(high precision at low force and high force with minor precision requirements). 
Apart from plastic changes that take place during and after motor skill acquisition, many 
studies have been carried out concerning short-term modulations of corticospinal excitability 
during motor task preparation and execution (Johansson, Lemon, & Westling, 1994; Mills, & 
Kamiskidis, 1996). However, slow precision hand movements, such as reaching and grasping, 
have been studied in greatest depth (Gangitano, Mottaghy, & Pascual-Leone, 2001; Liepert, 
Dettmers, Terborg, & Weiller, 2001). During these manipulative and explorative motor acts 
corticospinal excitability continuously changes through different phases of the movement; 
thereby being additionally modulated by precision requirements, grip force, sensory feedback, 
etc. (Lemon, Johansson, & Westling, 1995). In general, it seems that many of these regulations 
of cortical excitability are derived from sensory information flow. The role of sensory input has 
been studied in great detail through voluntary motor control as well as at a more elementary 
level. The afferent flow has been demonstrated to play a major role in motor control (Wing, 
Haggard, & Flanagan, 1996), which represents a whole new sphere of scientific activity in 
which TMS has been widely applied.  
In contrast to the on-line controlled prehension movement s , TMS ha s a l so been u sed to u n r avel 
the neural control of pre-programmed fast ballistic movements (Mills & Kimiskidis, 1996). 
Previously, peripheral features of such rapid movements had been described as comprising: 
a three burst pattern, pre-movement depression, intramuscular synchronization of motor 
units and a consecutively high level of rate of force development, etc. (for review see Zehr & 
Sale, 1994). In addition to the output characteristics of the explosive movement, the central 
neural activity responsible for the preparation and execution of such motor tasks can now be 
measured (MacKinnon & Rothwell, 2000). 
TMS has also been proved to be a useful method for the study of the involvement of the mo-
tor cortex during human locomotion and other cyclic movement activities. Petersen et al. 
(2001) demonstrated suppression of ongoing EMG activity during human walking that was 
provoked by subthreshold TMS. Christensen, Andersen, Sinkjer, and Nielsen (2001) showed 

94 Transcranial magnetic stimulation & sport science Kinesiologia Slovenica, 10, 2, 78–104 (2004) 
strong facilitation of the TMS-elicited MEPs in the inactive tibialis anterior muscle during 
the stance phase. Capaday et al. (1999) compared MEP input-output curves measured (i) dur-
ing voluntary tonic activity of either the soleus or tibialis anterior muscle and (ii) during the 
early part of the swing phase and the stance phase of walking respectively. Their data suggest 
that during walking the corticospinal tract is more closely linked with the segmental motor 
circuits controlling the flexor (tibialis anterior) than with those controlling the extensor (so-
leus). Similar reports on the modulation of transmission in the corticospinal pathways to the 
soleu s have a lso been proved du r i ng bic ycl i ng (P y ndt & Nielsen, 20 03). Si m i la rly, t y pic a l c ycl ic 
movement patterns found in sports could be analysed. Additionally, different techniques of 
running, cross country skiing, bicycling, etc. could be compared, which could reveal either a 
general modulation type or specificity for each individual movement activity.  
It is clear that none of the described areas of TMS application could be excluded as irrelevant 
to sports science. It is believed that TMS could be further fruitfully implemented in kinesiol-
ogy and its subdisciplines, helping to answer questions specific to sports. Basic knowledge of 
post-exercise corticospinal facilitation could be used to pose new questions and to explore the 
effec t s of d i fferent t ra i n i ng met hod s t houg ht to provoke neu ra l ad apt at ions . Resea rch cou ld be 
carried out to determine whether the proposed early central adaptations to activation training 
really occur. Furthermore, long-term neural adaptations to strength and power training could 
be assessed more analytically by the use of TMS and tools for peripheral neuromuscular excit-
ability testing. Composing the input-output curve makes it possible to employ longitudinal 
studies that are needed to study specific adaptations over a longer time period. Manipulations 
of sensor i motor i nteg rat ion mecha n isms seem ver y prom isi ng for opt i m isi ng neu ra l plast icit y 
processes, with TMS being an indispensable tool for the evaluation of such manipulations. 
TMS and measures of cortical excitability might also be of assistance in the detection of other 
problems deriving from sports practice and theory. In short, there are many theoretical expla-
nations of training states and effects that need to be tested. In our opinion, TMS could also be 
effectively employed in research into areas such as overtraining, endurance and fatigue, and 
the effects of sport psychological techniques. 
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Acknowledgements
The author wishes to express his gratitude to Prof. Milan R. Dimitrijević, MD, DSc, for his 
guidance and continued support.