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Training-Induced Changes in Neural Function
Per Aagaard
Department of Neurophysiology, Institute of Medical Physiology, Panum Institute and Team Danmark
Testcentre, Sports Medicine Research Unit, Bispebjerg Hospital, University of Copenhagen, Denmark

AAGAARD, P. Training-induced changes in neural functions. Exerc. Sport Sci. Rev., Vol. 31, No. 2, pp. 61– 67, 2003.
Adaptive changes can occur in the nervous system in response to training. Electromyography studies have indicated adaptation
mechanisms that may contribute to an increased efferent neuronal outflow with training, including increases in maximal firing
frequency, increased excitability and decreased presynaptic inhibition of spinal motor neurons, and downregulation of inhibitory
pathways. Keywords: CNS, spinal, human, motor neurons, muscle

INTRODUCTION the pickup volume of the recording electrodes. This overall
interference signal is modified by a multitude of intracellular
Adaptive alterations can be induced in the neuromuscular and extracellular factors, which all exert a significant influ-
system in response to specific types of training. Thus, in- ence on the pattern of spatial and temporal summation of the
creases in maximal contraction force and power as well as single action potentials. From a physiological perspective, the
maximal rate of force development (RFD) will occur not only EMG interference signal is a complex outcome of motor unit
because of alterations in muscle morphology and architecture recruitment and firing frequency (rate coding) that also re-
(2), but also as a result of changes in the nervous system flects changes in the net summation pattern of motor unit
(1,4,12). potentials, as occurs with motor unit synchronization. Nu-
Evidence of the adaptive change in neural function with merous studies have reported increased EMG amplitude after
training has been provided through the use of electromyo- resistance training. The training-induced increase in EMG
graphy (EMG). Although consistent data can be obtained by that has been observed in highly trained strength athletes
EMG recording (7), inherent methodological constraints indicates that neural plasticity also exists in subjects with
may sometimes exist for the measurement of surface EMG highly optimized neural function.
during voluntary muscle contraction. To overcome some of Substantial cancellation of the EMG interference signal
these problems, EMG normalization procedures, single motor can occur due to out-of-phase summation of motor unit
unit recording techniques, and measurements of evoked re- action potentials (MUAPs), and it has been suggested, there-
flex responses (Hoffmann reflex, V-wave) have been increas- fore, that the EMG interference amplitude does not provide
ingly used to examine the change in neural function induced a true estimate of the total amount of motor unit activity (6).
by training. For example, increased motor unit synchronization will cause
the EMG signal amplitude to increase (16) attributable to the
elevated incidence of in-phase MUAP summation. Conse-
CHANGES IN EFFERENT NEURAL DRIVE ASSESSED quently, the increase in EMG interference amplitude ob-
BY EMG served after resistance training could indicate changes in
motor unit recruitment, firing frequency, and MUAP syn-
The interference EMG (Fig. 1) comprises the composite chronization. In addition, not all studies have been able to
sum of all the muscle fiber action potentials present within demonstrate elevated EMG activity after resistance training.
The inability to detect longitudinal EMG changes could be,
at least in part, due to changes in skin and muscle tissue
Address for correspondence: Per Aagaard, Ph.D., Dept. of Neurophysiology, Institute properties (subcutaneous fat layer, muscle fiber pennation
of Medical Physiology 16.5.5, Panum Institute, Blegdamsvej 3, 2200 Kbh-N, Copen-
hagen, Denmark (E-mail:
angle) or could arise from changes in electrode positions
Accepted for publication: October 3, 2002. between testing sessions. As discussed below, however, it is
possible to reduce some of these limitations by employing
Exercise and Sport Sciences Reviews
intramuscular EMG recordings, EMG normalization proce-
Copyright © 2003 by the American College of Sports Medicine dures, and measurements of evoked reflex responses.


Importantly. When contractile force is less than (12). CHANGES IN MOTONEURON FIRING FREQUENCY continues to increase at stimulation rates higher than that needed to achieve maximum tetanic tension (Fig. In fact. This phenomenon has been referred to as the 62 Exercise and Sport Sciences Reviews www.B. In these specific contraction conditions. and intact of force development. interval). This pattern was less frequent after intense resistance training. 90°–10° ⫽ range of motion. It Motor unit firing rates have been recorded at much higher is possible. as demonstrated using constant-frequency stimula- firing frequency has a strong influence on the contractile rate tion of single motor units.acsm-essr. whole isolated muscle. Data from (1). top curve). When rectified and low-pass filtered EMG signals were analyzed in untrained . it can be temporarily elevated by maximal force generation (MVC). Resistance training with heavy loads has consistently been shown to increase maximal eccentric and slow concentric contraction strength (quadriceps femoris muscle. A. where the concentric and eccentric EMG signals became more similar. which are characterized by high levels of contractile force generation.Figure 1. After a regimen of prolonged resistance training with heavy loads. bottom curve). that supramaximal firing rates in the frequencies than that needed to achieve full tetanic fusion in initial phase of a muscle contraction serve to maximize the force. therefore. the rate of force development human muscle. firing rates of 100 –200 Hz can be ob. Knee joint moment (Moment of Force) and interference EMG recorded in an untrained subject during maximal effort concentric and eccentric contractions of the quadriceps muscle performed in an isokinetic dynamometer (knee joint angular velocity 30°s⫺1. During eccentric contraction large EMG spikes typically were observed separated by interspike periods of low or absent activity. 0° ⫽ full extension). For example. which typically occurs the addition of an extra discharge pulse (1–5 ms interpulse 250 – 400 ms after the onset of contraction. the suppression of the EMG signal amplitudes was fully abolished (rectus femoris (RF)) or partially removed (lateral vasti (VL) and medial vasti (VM)) in parallel with a marked increase in maximal eccentric muscle strength. muscle activation appears to be suppressed in untrained subjects despite a maximal voluntary effort (EMG. 2) (9). EMG amplitudes were 20 – 40% less during maximal eccentric than concentric contraction (see B). rate of force development rather than to influence maximal served at the onset of maximal voluntary muscle contraction contraction force per se. with much lower rates (15–35 Hz) at the instant of the maximal tetanized level.

involving maximal intentional rate of force development. Copyright © have been observed after resistance training (3. makes it possible to reach a higher force and velocity during fast movements. in- catch-like property of skeletal muscle. Although the of force development and maximal firing frequency. this difference is reduced with resistance muscle before and after a period of ballistic-type resistance training. Acutely. Biomech. the frequency of muscle fiber action potentials ob- tained during maximal voluntary contraction was signifi- cantly greater in trained elderly weight lifters compared with age-matched untrained individuals. ballistic-type resistance training. i. and third induced by resistance training may effectively overrule the time intervals between successive action potentials. recorded in the tibialis anterior than young subjects. J. it is possible that the firing of discharge in the firing pattern of individual motor units following doublets at the onset of contraction and during the phase of resistance training (12).12). wire electrodes). Appl. Used with permission. a marked increase in EMG amplitude and the rate of rise in EMG can be seen in the initial contraction phase (Fig.] ticular. Volume 31 䡠 Number 2 䡠 April 2003 Training and Neural Function 63 . markedly increased the incidence of discharge doublets in the firing pattern of individual motor units (from 5% to 33%) while also increasing the rate of force development (12). An increase in mo- age-related decline in maximal discharge rate. training. Force-time curves for isolated motor units in the rat soleus unexpected perturbations in postural balance. and when activated at a supramaximal rate (RG) an increase in efferent neural drive.G. Thus. firing frequency probably represent major mechanisms re- like property. Moreover. muscle when activated at the minimum frequency needed to elicit maxi.. time to peak tension or the recruitment threshold. Duchateau and terspike interval ⬍ 10 ms) may be observed in the firing colleagues recently reported concurrent increases in the rate pattern of single motor neurons (see (12)).e. A training-induced increase in the rate of force devel- opment. Note that the rate of force de.. therefore. The maximal firing frequency of motor units can be ex- amined by use of intramuscular EMG-recording techniques (multipolar needle. the rate of force development is enhanced with mal tetanic fusion (PO). in the firing frequency of motor units (Fig. 12:285–291. Training-induced increases in the maximal frequency of muscle fiber action potentials appear to occur in both young and elderly individuals. both in highly trained athletes as well as elderly individuals who need to control Figure 2. second. so-called discharge doublets (in- of force development. The counteract the gradual decline in activation of muscle fibers number of discharge signals analyzed in each interspike period ranged between 243 and 609. This elevated incidence in discharge rising muscle force serves to enhance the initial generation of doublets and the corresponding rise in initial motoneuron muscle contraction force by taking advantage of the catch. Instantaneous motor unit firing frequency (⫾SEM) at the derly subjects demonstrate a lower maximal discharge rate onset of maximal ballistic contractions. Based on such tech- niques. Increases in firing frequency appeared to occur and the associated impairment in muscle function observed independently of motor unit size. CHANGES IN RATE OF FORCE RISE AND EMG DEVELOPMENT An increase in the rate of force development is perhaps the single most important functional benefit induced by resis- tance training. hence increasing the rate of force development. the rate of force development plays an important role in the ability to perform rapid and forceful movements. 2). At the onset of rapid cluding an elevated incidence of discharge doublets (12). As described above. which suggests that neural adaptation mechanisms. 4). Interestingly. 1996. Although el. Supramaximal activation increases motor unit velocity of unloaded shortening. as changes were not related to either with increasing age.e. the increase in maximal firing frequency performed at maximal intentional rate of force development.001). Figure 3. In par- 1996 Human Kinetics Publishers. 3). maximal fir- ing frequency has been reported to increase in response to resistance training (Fig. as all post- represent a highly beneficial type of neural adaptation to training values were greater than pretraining values (P ⬍ 0. Parallel in- velopment is greater at supramaximal rate of stimulation. particularly by increases that also elicited maximal tetanic fusion. This would toneuron firing frequency was observed following training. which are considerably less than the time it takes to reach maximal muscle force (~300 ms). [Adapted from creases in the rate of force development and EMG amplitude Nelson. Data from (12). are highly important for the training-induced increase in the rate muscle contractions in vivo. i. A. Rapid movements may involve muscle con- traction times of 50 to 200 ms. together functional consequences of such discharge doublets are not with a sixfold increase in the incidence of discharge doublets fully understood. Importantly. Bars show the mean discharge frequency recorded in the initial.

Data from (3). using single motor unit recordings in muscles of the hand and lower back. as recurrent inhibition appears to be absent in the smaller distal muscles of the hands and feet. rectus femoris (RF)) during maximal isometric contraction before (open bars) and after (closed Electromyography recordings in untrained subjects have bars) 14 wk of resistance training.01. Importantly. Notably. inhibition in muscle activation appears to be downregulated or fully removed in response to resistance training with heavy sponsible for the increase in the rate of force development loads (1). Efferent motor output during maxi- spinal motor neurons. whereas no effect was observed during concentric contractions (14). vastus medialis (VM). group pathway. as suggested by a depression in H-reflex amplitude during eccentric contraction. cells receive several types of supraspinal synaptic input that can enhance as well as depress the recurrent pathway. and has been considered to central descending pathways. * P ⬍ 0. This suggests that explosive-type resistance training (i. suggesting that the Figure 4. muscle (vastus lateralis (VL).org . Electrical stimulation of passive versus active muscle has been used to address the issue of neural activation during maximal eccentric contractions. training-induced eccentric strength typically observed with this type of changes in muscle fiber size and muscle architecture (2) training. which results in reduced recurrent inhibition. with maximal concentric contraction (1). rate of EMG rise.05. would additionally contribute to the increase in rate of force Although several mechanisms have been proposed. However. as EMG is reduced compared ⬎ pre: RFD and average EMG. this * P ⬍ 0. Post eccentric contractions (Fig.. which explains the marked increase in maximal observed with training. preferential activation of high-threshold motor units has been observed in the triceps surae muscle during submaximal eccentric contraction. Thus. the excitability of spinal motor neurons or presynaptic inhibition of Ia afferents in the soleus muscle appears to differ between submaximal ec- centric and concentric contractions at matched EMG lev- els. However. ** P ⬍ 0. Interestingly. group III muscle affer- 64 Exercise and Sport Sciences Reviews www. this evoked increase in eccentric contraction strength was seen in untrained sub- jects but not in strength-trained subjects. has been considered as a mal voluntary muscle contraction not only is regulated by limiting factor for discharge rate. Time intervals denote time relative to shown that muscle activation is suppressed during maximal contraction onset (for RFD) or onset of EMG (for all EMG parameters). apparent inhibition in maximal eccentric muscle strength erage EMG and rate of EMG rise) obtained in the quadriceps femoris can be removed by resistance training.e. Renshaw cell activity appears to be more inhibited during maximal phasic muscle contractions. actual neural regulatory pathways responsible for the sup- The enhancement of motor unit activity may involve pression of muscle activation during eccentric contraction changes in neural circuitry. the majority of studies. training involving a high rate of force development) may be optimal for evoking changes in maximal firing rate of motor units. Despite a maximal intended effort by the subjects. which was suggested to result from increased presynaptic inhibition of Ia afferents that synapse onto low- threshold motor neurons. Contractile rate of force development (RFD) and EMG (av. 1).001.01. Com- pared with steady-force contractions. Animal experiments have shown that Renshaw Ia and II muscle spindle afferents. ** P ⬍ 0. this effect may be restricted to certain mus- cles. the force achieved during maximal eccentric contraction was enhanced with superimposed elec- trical stimulation. have failed to demonstrate selective recruit- ment of high-threshold motor units during eccentric con- tractions. for example. the development. Recurrent Renshaw inhibition of remain unidentified. but also is modulated by have a regulating influence on the reciprocal Ia-inhibitory afferent inflow from group Ib Golgi organ afferents. CHANGES IN NEURAL INNERVATION DURING MAXIMAL ECCENTRIC MUSCLE CONTRACTION It has been suggested that eccentric muscle contractions require unique neural activation strategies. Furthermore.acsm-essr.

R.. Used with permission. Trends Neurosci. Yoshitake. thereby reducing the magnitude of excitatory inflow to examine the importance of neural mechanisms for the to motor neurons. 8:363–381. a training-induced re. and C. [Adapted from Moritani. contraction. and if such observed during active eccentric versus concentric con. in turn eliciting a reflex response (H) with a latency of 30 –35 ms for the human soleus muscle. Also. The use of electromyography in applied physiology. Capaday.] ents. and Stein. tractions (see above). The duction in presynaptic inhibition of Ia muscle spindle Hoffmann (H) reflex may be useful for the assessment of afferents could contribute to an elevated excitatory inflow motoneuron excitability in vivo. It has motor neurons as well as the possible alteration in presynap- been suggested that the removal of neural inhibition and tic Ia afferent inhibition during maximal eccentric contrac- the corresponding increase in maximal eccentric muscle tion. possibly by central descending path. Golgi Ib afferents Future studies are needed to address the training-induced activate interneurons in the spinal cord. Kinesiol. ticity with training (5). although it also reflects the Volume 31 䡠 Number 2 䡠 April 2003 Training and Neural Function 65 .B. difference is muscle specific. Electromyogr. that in eccentric contraction the spinal RESPONSES inflow from Golgi Ib afferents and joint afferents induce elevated presynaptic inhibition of muscle spindle Ia affer. Evoked spinal motoneuron responses examined by use of the Hoffmann (H) reflex.. A variant of the H-reflex. For example.Figure 5. 1988. The possibility ex. Not only have reduced H-reflex responses been tion of both submaximal and maximal intensity. indicating an elevated descending motor drive from supraspinal centers. 1998. therefore. Consequently. Electrical stimulation of Ia afferent nerve fibers (“sensory”) excites spinal motor neurons (␣). so-called V-waves. Elevated V-wave and H-reflex responses have been observed following resistance training. Copyright © 1988 Elsevier Applied Science Publishing. and Y. Copyright © 1998 Elsevier Science. increased excitability of spinal motor neurons and/or decreased presynaptic inhibition of muscle spindle Ia afferents. which are also change in excitability and postsynaptic inhibition of spinal influenced by descending corticospinal pathways. can be recorded using supramaximal stimulation intensities during ongoing maximal muscle contraction. J. The electrical pulse also gives rise to action potentials in the motoneuron axones (“motor”) resulting in a direct M-wave with a latency of 3– 4 ms. 11:328 –332. T. All to spinal motor neurons during maximal eccentric muscle of these pathways are expected to exhibit adaptive plas. The modulation of human reflexes during functional motor tasks. training-induced increase in maximal muscle strength. the H-reflex also appears to be markedly suppressed during passive lengthening compared with shortening of the soleus muscle. Few studies have measured responses in spinal motoneuron ents. CHANGES IN EVOKED SPINAL MOTONEURON ists. recruitment order and firing rate of individual motor units in terneuron activity. additional studies using intramuscular EMG tech- strength observed following resistance training could be niques are needed to examine further whether the specific caused by a downregulation of such spinal inhibitory in. and by recurrent inhibition from Renshaw cells. fact differ between eccentric and concentric muscle contrac- ways (1).

thus allowing more of the evoked H-reflex volley to a suppressed stretch-reflex response in antagonist-agonist reach the muscle fibers as manifested by an increase in muscle pairs during postural balance tasks. magnitude of presynaptic inhibition of Ia afferent synapses also be recognized that training-induced changes in postsyn- (see (15)).org . which increases the outflow of efferent motor impulses jects. Elevated H- of spinal motor neurons or reduced presynaptic inhibition of reflex responses have been obtained during submaximal and Ia afferents would contribute to the observed increase in maximal voluntary muscle contraction after hopping training V-wave amplitude.. It has been observed. it would seem desirable to achieve impulses. (13) and resistance training (4). Resistance training can induce adaptive alterations in nervous system function. could con- stimulation of the peripheral nerve containing Ia afferents tribute to an increase in evoked V-wave and H-reflex and motor axons (Fig. which will benefit certain individuals and athletes (i. At high activate motor units during maximal voluntary contraction. suggesting that the the spinal cord toward the muscle fibers due to the volley of relative contribution of the above mechanisms may differ H-reflex impulses. A supramaximal intensity elic. the responses. From will produce an increased cancellation of the antidromic a functional perspective. neural adaptation mechanisms play important roles for the training-induced increase in maximal eccentric strength and contractile rate of force development (RFD). Hence.13). It is difficult to elicit V-wave changes in certain (ii) action potentials that propagate orthodromically from hand muscles with resistance training.11).Resistance training aimed at maximizing neural components will induce gains in muscle strength with no or only minor increases in muscle and body mass. superimposed during ongoing voluntary muscle contraction. amplitude of the H-reflex varies nonmonotonically with In cross-sectional studies.Figure 6. achieving maximum amplitude at an in.acsm-essr. Considerable training-induced plasticity appears to exist the H-reflex response reappears (denoted a V-wave) as the for the excitatory and inhibitory pathways in the spinal cord. the synaptic inhibition of soleus Ia afferents (from reciprocal H-reflex response recorded at rest remained unchanged with inhibitory pathways) is absent or greatly reduced during max. and thinnest arrows indicate a low-to-moderate influence. to avoid the V-wave amplitude (4. antidromic impulses in the motor axons now collide with Upregulation and downregulation of the H-reflex have both efferent nerve impulses generated by the voluntary motor been demonstrated in monkeys and rats exposed to long-term effort (11). Training that results in both improved neural function and gains in muscle mass will benefit not only explosive-type athletes but also aged individuals. training (4. which likely reflects altered states in the reciprocal in the axons. respectively. intensities of stimulation. distance runners. have demonstrated markedly elevated V-wave amplitudes in termediate stimulus intensity. which was interpreted as an increased ability to activating all motor axons in the peripheral nerve. These findings indicate that resting H-reflex imal plantarflexor contraction in young subjects. Excitability of the soleus H-reflex results in an increased motor neuron recruitment and firing decreased after short-term balance training by human sub- rate. any increase in descending motor drive inhibitory pathways of the tibialis and soleus muscles. however. triathletes. An increased central descending motor drive conditioning paradigms. 5) (see (4).e. resistance training appears to induce increased due to collision between (i) action potentials that travel V-wave and H-reflex amplitudes during maximal muscle con- antidromically in the motor axon toward the spinal cord and traction (4). Likewise. In the resting muscle. cyclists). thinner arrows a moderate influence. as for the frail elderly this will provide an effective mean to improve everyday physical function. the H-reflex is elicited by electrical aptic inhibition. the hand and lower limb muscles compared with untrained its a maximal direct M response (Mmax) by orthodromically subjects. In brief. an increased excitability occurrence of reflex-mediated joint oscillation. elite weight lifters and sprinters stimulus intensity. the H-reflex response is abolished Furthermore. It should measurements may not adequately reflect the state of the 66 Exercise and Sport Sciences Reviews www. such as via Golgi Ib afferents. When supramaximal nerve stimulation is between various muscles in the body. In particular. Thick arrows indicate a strong influence. that pre. along with changes in the morphology and architecture of the trained muscles. Notably.

stimulation on eccentric and concentric torque-velocity relation- hagen. 534: ical limitations associated with conventional surface EMG 613– 623. 8:363–381. J. Also of impor- J. Volume 31 䡠 Number 2 䡠 April 2003 Training and Neural Function 67 . 2001. 4. J. increased mo. and M. Aagaard. Acta Physiol. Copen.L. Jesper L. Andersen. E. Yao. 3. Eur. Neurophysiol. and K.spinal circuitry during activity. Simonsen. Changes in the excitability of soleus Thanks to coworkers and colleagues who have contributed in indispensable muscle short latency stretch reflexes during human hopping after 4 wk ways: Erik B. Psychiat. 15.G. A. Neurol. Aagaard. J. W. Kinesiol. 11:328 –332. M. 8. Electromyogr. Physiol. 2000. 30:59 – 63. Appl. toneuron excitability and decreased presynaptic inhibition.J. P. as well as physiology. 1996. Duchateau. 2001. Supramaximal activation increases motor unit velocity of increased levels of central descending motor drive. 2. neural adaptation mechanisms are Sports Sci. A. Wagner. Physiol. Physiol. J. S. The trains. Y.. Dyhre. Capaday. T. Nelson. Halkjær-Kristensen. Zehr. E. and P. Seger. Clin. for continuous support and to Roger M. Biomech. Appl. Bispebjerg Hospital. The use of electromyography in applied downregulation of inhibitory neural pathways. In particular. 2002.B. for constructive input during preparation of the manuscript. A. and P. Dyhre-Poulsen.. S. 9. Sports Medicine Research Unit.B. Andersen. Experimental simulation of cat electromyo- gram: evidence for algebraic summation of motor-unit action-potential and rate of force development observed with training. Andersen. Enoka.L. system as well as in the morphology of the trained muscles 5. 1971. Sica. 1990. Van Cutsem.. 513:295–305. Chelli.P. Further unloaded shortening. E. young versus elderly individuals.. Magnusson.P. Day. Thorstensson. Magnusson. J. 1988. J. Increased rate of force development and neural drive following resistance training.E. P. Physiol. Neural control of motor output: can training change it? Exerc. Neural inhibition during maximal contractions. The modulation of human reflexes during involvement and functional significance of these neural fac. Simonsen. 83:441– 452. 16.M. 1996. Trends Neurosci. S.M. J. Bawa. Poul Dyhre-Poulsen. Resistance training elicits adaptive changes in the nervous 2002. to strength training: changes in muscle architecture. J. Magnusson. Simonsen. research is needed to obtain knowledge about the relative 10. normalizing the evoked H-reflex and V-wave re. Halkjær. 89:2249 –2257. Fuglevand. Dyhre-Poulsen. and Hanne Overgaard. Appl. and A. Eur. Yoshitake. (Fig. Physiol. Andersen. 13.P. Frigo. and G. Potentiation of “late” to examine the relative importance of these mechanisms in responses evoked in muscles during effort. G. P. Upton. Magnusson. Leffers. 140:17–22. Considerations for use of the Hoffmann reflex in exercise References studies. Andersen. J. for increased contractile strength of human pennate muscle in response sponses to the maximal M-wave minimizes the methodolog. Aagaard. J.M. S. P. A mechanism tance. Neurophysiol. Effects of electrical Michael Kjær. Kamen.P. Changes in single motor unit behavior contribute to the increase in contraction speed Acknowledgments after dynamic training in humans. 12. Neural adaptation to resistance training: Changes in evoked V-wave and H-reflex responses. increase in motor neuronal output in response to resistance 7. 6). S. Appl. 2000. Simonsen. Magnusson. orado. 13:366 –384.L. electromyogram. 2002... Westing. important for the increases in maximal eccentric strength 6. J. ships during knee extension in man. J. and C. 1998. 12:285–291. Scand. M. Voigt. E. Å. Aagaard. and P. Physiol.. and C. 2002. R.E. and Y.L. F. and 11. Rev. and R. Peter of hopping training. Hainaut. J. J. J. increases EMG amplitude and decreases force steadiness of simulated Kristensen. Appl. J.B. 93:1318 –1326. Simonsen. S. University of Col.Poulsen. Physiol.J. 1998.B. A. Benny Larsson. J.. 92:2309 –2318.H. 34:699 –711. E... Motor-unit synchronization 1. J. Enoka. 78:522–532. Physiology and interpretation of the training may involve increased firing rates.P. A. recordings. 1998. Hulliger. Neurosurg. S.. P. 86:2144 –2158.R. Moritani.P..J. Caldwell. and R. tasks and not solely in the resting muscle.B. Neurophysiol. Stein.. McComas. CONCLUSIONS Dyhre-Poulsen. functional motor tasks. Also thanks to Professor 14. Appl. 86:455– 468. J. advocating that evoked reflex eccentric and concentric quadriceps contraction: effects of resistance measurements should be performed in functional contraction training. J. tors with specific types of physical activity and training. P.