Journal of Sports Sciences, 2002, 20, 951± 959

Maximal-intensity isometric and dynamic exercise

performance after eccentric muscle actions
CHRISTOPHER BYRNE1* and ROGER ESTON2
1
Military Physiology Laboratory, Defence Medical Research Institute, Defence Science and Technology Agency,
74 Loewen Road, Republic of Singapore 248843 and 2School of Sport, Health and Exercise Sciences,
University of Wales Bangor, George Building, Bangor, Gwynedd LL57 2PX, UK

Accepted 10 June 2002

A well-documented observation after eccentric exercise is a reduction in maximal voluntary force. However, little
is known about the ability to maintain maximal isometric force or generate and maintain dynamic peak power.
These aspects of muscle function were studied in seven participants (5 males, 2 females). Knee extensor iso-
metric strength and rate of fatigue were assessed by a sustained 60 s maximal voluntary contraction at 80°
and 40° knee ¯ exion, corresponding to an optimal and a shortened muscle length, respectively. Dynamic peak
power and rate of fatigue were assessed during a 30 s Wingate cycle test. Plasma creatine kinase was measured
from a ® ngertip blood sample. These variables were measured before, 1 h after and 1, 2, 3 and 7 days after
100 repetitions of the eccentric phase of the barbell squat exercise (10 sets ´ 10 reps at 80% concentric one-
repetition maximum). Eccentric exercise resulted in elevations in creatine kinase activity above baseline
(274 ± 109 U ´l-1; mean ± sxÅ ) after 1 h (506 ± 116 U ´l-1, P < 0.05) and 1 day (808 ± 117 U ´l-1, P < 0.05). Iso-
metric strength was reduced (P < 0.05) for 7 days (35% at 1 h, 5% at day 7) and the rate of fatigue was lower
(P < 0.05) for 3 days at 80° and for 1 day at 40°. Wingate peak power was reduced to a lesser extent (P < 0.05)
than isometric strength at 1 h (13%) and, although the time course of recovery was equal, the two variables
diþ ered in their pattern of recovery. Eccentrically exercised muscle was characterized by an inability to generate
high force and power, but an improved ability to maintain force and power. Such functional outcomes are
consistent with the proposition that type II ® bres are selectively recruited or damaged during eccentric exercise.

Keywords: eccentric exercise, fatigue, muscle damage, strength, Wingate test.

Introduction 1992; Cleak and Eston, 1992; Byrne et al., 2001) or 1

month (Howell et al., 1993; Sayers and Clarkson, 2001).
Exercise-induced muscle damage often results from Muscle function has primarily been assessed using
eccentric muscle actions and is synonymous with measures of isometric maximal voluntary force (Davies
impaired muscle structure (Friden et al., 1983; and White, 1981; Newham et al., 1983b; Jones et al.,
Newham et al., 1983a; Jones et al., 1986), metabolism 1989; Clarkson et al., 1992; Cleak and Eston, 1992;
(O’ Reilly et al., 1987; Asp et al., 1998) and function Howell et al., 1993; Sayers and Clarkson, 2001) and,
(Sargeant and Dolan, 1987; Clarkson et al., 1992). more recently, dynamic isokinetic force (Golden
Perhaps the most important feature of eccentric exercise- and Dudley, 1992; Eston et al., 1996; Hortobagyi et al.,
induced muscle damage is the long-lasting impairment 1998; Byrne et al., 2001; Byrne and Eston, 2002). Much
of muscle function. Strength is typically reduced less is known about the ability of eccentrically exercised
immediately after eccentric exercise, followed by a muscle to sustain isometric force (Davies and White,
linear recovery over a time course of hours (Davies and 1981; Balnave and Thompson, 1993; Behm et al., 2001)
White, 1981; Newham et al., 1983b), days (Golden or the ability to generate and maintain maximal power
and Dudley, 1992; Hortobagyi et al., 1998; Byrne and output (Sargeant and Dolan, 1987).
Eston, 2002) or even in excess of 1 week (Clarkson et al., Fatigue has been de® ned as `any exercise-induced
reduction in force generating capacity’ (Gandevia,
* Author to whom all correspondence should be addressed. e-mail: 2001). The long-lasting reduction in force-generating
[email protected] capacity that results from eccentric exercise is more

Journal of Sports Sciences ISSN 0264-0414 print/ISSN 1466-447X online Ó 2002 Taylor & Francis Ltd
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952 Byrne and Eston

suitably viewed as muscle weakness rather than muscle a reduction in maximal isometric strength of a similar
fatigue, since it is not an acute eþ ect that is reversed by magnitude.
rest. Gandevia (2001) stated that the maximal force- Several events that occur during and after eccentric
generating capacity of muscles starts to decline once exercise-induced muscle damage are likely to have a
exercise commences, so that fatigue really begins almost large impact on maximal dynamic performance.
at the onset of exercise and develops progressively Evidence suggests that type II ® bres are selectively
before the muscles fail to perform the required task. damaged during eccentric exercise (Friden et al., 1983;
This aspect of muscle function has been studied Jones et al., 1986) and a prolonged decrease in muscle
infrequently after eccentric exercise. Davies and White glycogen content is commonly observed (O’ Reilly et al.,
(1981) investigated the ability of the triceps surae 1987; Costill et al., 1990), which is especially pro-
muscle group to maintain isometric force after eccentric nounced in type II ® bres (Asp et al., 1998). The rapid
exercise and reported the musculature to be weaker but and marked rise and subsequent decline in force and
no more fatiguable, whereas Balnave and Thompson power output during maximal intensity exercise, as
(1993) reported that the knee extensors were weaker observed during sustained MVCs or maximal cycling,
but, surprisingly, less fatiguable after eccentric exercise. is thought to be closely related to the activation and
Both studies used an electrically elicited isometric rapid fatigue of type II ® bres (Nevill et al., 1996). As the
fatigue test consisting of trains of stimuli at 20 Hz, generation of power output by muscle shortening is
lasting 300 ms and repeated every second for 2 min. more energetically demanding and requires higher rates
Using voluntary muscle actions, Behm et al. (2001) of adenosine triphosphate (ATP) regeneration than
reported a signi® cant reduction in endurance time isometric force generation (Woledge et al., 1985), the
of the elbow ¯ exors when isometric contractions were possibility exists that more dramatic performance
maintained at 50% of the pre-exercise maximal volun- decrements will be observed for power-generating
tary contraction (MVC) force. No previous studies have ability than those commonly observed for the genera-
investigated the ability of eccentrically exercised muscle tion of isometric force. Therefore, in the second part of
to maintain maximal voluntary force. The ® rst part of this study, we examined the eþ ect of eccentric exercise
this study assessed the ability of the knee extensors to on the ability to generate maximal isometric force and
maintain force during sustained isometric MVCs of 60 s peak power (i.e. ® rst few seconds) and investigated the
duration after eccentric exercise. Evidence suggests that rate of fatigue during a 60 s isometric MVC and during
muscle fatigue during MVCs is length-dependent a 30 s maximal-intensity Wingate cycle test.
(Fitch and McComas, 1985; McKenzie and Gandevia,
1987), with muscle demonstrating a greater ability to
maintain force and appear fatigue-resistant when tested Methods
at a shortened rather than an optimal muscle length
for force production. To investigate this phenomenon Participants and design
further, MVCs were performed at an optimal muscle Seven healthy participants, ® ve males and two females
length and at a shortened length before and for up to (age 22.6 ± 4.4 years, height 1.78 ± 0.08 m, mass
7 days after eccentric exercise. 75.7 ± 11.7 kg; mean ± s), were involved in the study,
To complement the isolated knee extensor experi- all of whom were moderately active, but had not
ments and in an attempt to make the results of the study participated in any resistance training of the lower limbs
more applicable to sport (Jones, 1993), we examined in the 6 months before the study, and none had
the ability of eccentrically exercised muscle to generate any musculoskeletal defects. Each participant provided
and maintain maximal power output. It is somewhat written informed consent to take part in the study,
surprising that performance in maximal intensity sprint which had previously been approved by the School
exercise, being essential to performance in many sports, ethics committee. The muscle group studied was the
has received almost no attention after eccentric exercise. knee extensors. Each participant was evaluated for each
In highlighting the lack of research in this area, Clarkson criterion measure before, 1 h after and 1, 2, 3 and 7 days
and Newham (1995) stated that it is not known if the after a bout of eccentrically biased resistance exercise,
time course of recovery for power generation is the same chosen to induce the symptoms of exercise-induced
as that for isometric strength. Previously, only Sargeant muscle damage.
and Dolan (1987) have investigated the ability of eccen-
trically exercised muscle to generate maximal power
Isometric strength at 40° and 80° knee ¯ exion
output, reporting an immediate, signi® cant and sus-
tained reduction in short-term power output, measured Isometric strength and fatigue were measured for the
concentrically on an isokinetic cycle ergometer. This knee extensors of the non-dominant limb using a
reduction persisted for 4 days and was accompanied by Kin-Com (500H, Chattecx, Chattanooga, TN, USA)
Strength, power and fatigue after eccentric exercise 953

isokinetic dynamometer. Participants performed iso- was analysed for lactate using an Accusport analyser
metric MVCs of the knee extensors at 40° and 80° knee (Boehringer Mannheim, Lewes, UK). The participants
¯ exion, corresponding to a shortened and optimal performed a 5 min warm-up on a cycle ergometer at
muscle length for torque generation, respectively. 100 W, which included a ¯ at-out sprint at 3 min for 5 s,
Newham et al. (1991) have previously reported that 80° followed by 5 min of rest. The participants were then
knee ¯ exion corresponds to the optimal angle for torque transferred to the test ergometer (Monark 814 E). Seat
generation in this muscle group. The 40° joint angle was height was adjusted for comfort and was recorded,
chosen to represent short muscle length, as it is a similar toe clips were secured, the resistive load was attached
relative position to the muscle length used by McKenzie and a restraining harness was positioned to prevent the
and Gandevia (1987), who reported less fatigue in the participants rising from the seat. The resistive load
elbow ¯ exors at short versus optimal muscle length. The corresponded to 10% and 8% of body mass for males
testing positions were obtained by entering full knee and females, respectively. The participants began pedal-
extension (0°) as a reference value into the Kin-Com ling at 50± 60 rev ´min-1 with the external load sup-
visual display. The reproducibility of this method was ported. Upon the command `3, 2, 1, GO’ , the load was
checked on each test occasion by noting the Kin-Com applied abruptly, the participants pedalled ¯ at out for
angle display when the lever arm was at true 90° 30 s, and power output data, corrected for the moment
(determined by spirit level). The pre-test angle display of inertia of the ¯ ywheel (Lakomy, 1986), was logged by
at true 90° was used as the criterion. If any diþ erence computer for 30 s. The participants then performed a
was noted, the process was repeated until the criterion warm-down, which consisted of 2 min of cycling at 100
was achieved. Three submaximal and one maximal W. A post-exercise blood lactate sample was taken at
practice repetitions acted as warm-up at each test 3 min. Peak power was quanti® ed as the highest power
position. Three MVCs of 3 s duration, presented in a output in the ® rst 5 s of the test. Regression analysis was
mixed order, were performed at each joint angle with a performed on the 30 data points and the regression
60 s rest between repetitions. The highest peak torque coeý cient (b) was used to quantify fatigue.
from the three muscle actions was used as the criterion The rationale for using the regression coeý cient to
score for short and optimal muscle length, respectively. quantify fatigue in MVC and Wingate tests is that the
calculation takes into account every data point rather
than a simple percentage decline based on the highest
Isometric fatigue at 40° and 80° knee ¯ exion
and lowest data points. Balnave and Thompson (1993)
Fatigue was assessed by having the participants perform also used this method, which allows for a direct com-
a 60 s MVC at both 40° and 80° knee ¯ exion. A target parison between the two studies. In addition, there
of 100% of the current MVC was entered into the appears to be no standard or widely accepted method of
Kin-Com visual display, which gave real-time feedback quantifying fatigue in the muscle function literature.
of the participants’ torque production and the target
torque. The test started when the participants achieved
Creatine kinase measurement
the 100% MVC target torque. Tests at 40° and 80°
were presented in a mixed order and were separated by Plasma creatine kinase activity was determined from a
20 min of recovery. Torque was recorded each second ® ngertip blood sample. A warm ® ngertip was cleaned
during the 60 s MVC with a sterile alcohol swab and allowed to dry. Capillary
Fatigue was quanti® ed as the slope of a linear regres- puncture was made with a softclix lancet and a sample of
sion line, ® tted to the 60 data points. The slope of the whole fresh blood (30 ml) was pipetted from a capillary
line or regression coeý cient (b) represents the rate of tube onto the test strip and analysed for creatine
change in y with a unit change in x. In this study, for kinase activity using a colorometric assay procedure
every unit change in x (time), y (torque) changed by b. (Re¯ otron, Boehringer Mannheim, Lewes, UK). This
Since torque declined over time during the fatigue test, system uses a plasma separation principle, which is
b always took a negative value. As such, the less negative incorporated in the reagent carrier on the test strip.
the value, the less fatiguable the knee extensors were.
Muscle-damaging exercise
The Wingate test
Each participant performed 10 sets of 10 repetitions
The Wingate test is a maximal-intensity exercise test of the eccentric phase of the barbell squat exercise.
used to assess power output of the legs. The sensitivity The load on the barbell corresponded to 80% of the
and reliability of the test have previously been examined participant’ s concentric one-repetition maximum. They
(Bar-Or, 1987). Before testing, a ® ngertip blood sample started the movement with the barbell resting on their
was taken using a softclix lancet and capillary tube and shoulders, body erect, legs fully extended (knee = 180°)
954 Byrne and Eston

and the toes pointing forward. The movement consisted sent a change in the length± tension relationship of the
of an eccentric action of the knee extensors to lower muscles due to a non-uniform distribution of sarcomere
the barbell to a knee angle of approximately 90°. The length change during eccentric exercise (for a review,
barbell was then raised to the starting position by see Morgan and Allen, 1999). Strength was reduced
two individuals acting as spotters. Two minutes of 1 h after exercise and remained so for 7 days (Fig. 1).
rest separated each set. Since all participants were Strength was approximately 35% lower 1 h after
unaccustomed to this form of exercise, the load was exercise (P < 0.05), 28% lower after 1 day (P < 0.05),
moderate to ensure correct technique and the move- 20% lower after 2 days (P < 0.05), 15% lower after 3
ment was slow and careful to avoid injury. The volume days (P < 0.05) and 5% lower after 7 days (P < 0.05).
of exercise was high and the amplitude of the stretch
large, since these parameters have been shown to induce
muscle damage (Newham et al., 1988). Furthermore, Isometric fatigue at 40° and 80° knee ¯ exion
this form of exercise has previously been shown to pro-
Signi® cant main eþ ects were revealed for time (F5,30 =
duce long-lasting decrements in muscle function (Byrne
23.2, P < 0.001) and angle (F1,6 = 17.0, P < 0.01) on the
and Eston, 2002).
rate of fatigue. The results also revealed a signi® cant
interaction of time and angle (F5,30 = 6.6, P < 0.001).
Statistical analysis Before damaging exercise, the knee extensors were
signi® cantly less fatiguable (P < 0.05) at 40° (b =
Isometric strength and fatigue data were analysed by -2.39 ± 0.26) than at 80° (b = -5.50 ± 0.72). After
two separate, two-factor (time and angle) repeated- eccentric exercise, the knee extensors became less
measures analyses of variance. A comparison of iso- fatiguable; however, the magnitude and time course of
metric strength and Wingate peak power was analysed this eþ ect was dependent on joint angle (see Fig. 2). At
by a two-factor (time and mode) repeated-measures 80°, the knee extensors were less fatiguable (P < 0.05)
analysis of variance. Wingate fatigue, creatine kinase after 1 h, 1 day and 3 days. At 40°, the knee extensors
activity and post-Wingate blood lactate concentration were less fatiguable (P < 0.05) after 1 day only.
were analysed using separate single-factor repeated- Figure 3 shows the response of one participant to the
measures analyses of variance. The assumption of isometric fatigue tests before eccentric exercise and 1
sphericity was tested by the Mauchly test of sphericity. day after. Before eccentric exercise, there was a greater
Any violations of this assumption were corrected initial isometric force generation and a greater rate of
using the Greenhouse-Geisser adjustment to raise the fatigue at the optimal versus shortened muscle length,
critical value of F, as indicated by (GG). To determine the as evidenced by the steeper slope of the regression line
regression coeý cient (b) to quantify fatigue, individual at 80° (b = -7.68) than at 40° (b = -1.84) knee ¯ exion
regression analyses were conducted on performances
during the isometric 60 s MVC tests and the Wingate
30 s test. Statistical signi® cance was set at P < 0.05.
Post-hoc tests were performed with paired-sample t-tests
using the Bonferroni correction technique.

Results

Isometric strength at 40° and 80° knee ¯ exion

Absolute values for isometric peak torque at 40° and 80°
were 302 ± 19 N ´m (mean ± sxÅ ) and 495 ± 33 N ´ m,
respectively. There was a highly signi® cant main eþ ect
for time (F5,30 = 50.8, P < 0.001) but no diþ erence in
strength loss between joint angles (F1,6 = 2.5, P > 0.05).
Although the extent of strength loss was clearly length-
dependent 1 h after exercise (see Fig. 1), the time by
angle interaction was not signi® cant (F5,30 = 1.0, Fig. 1. Changes in isometric strength at 40° (short; solid
P > 0.05). A disproportionate loss of relative strength bars) and 80° (optimal; open bars) knee ¯ exion after eccentric
at short versus optimal muscle length, either acute or exercise-induced muscle damage. Values are means (± sxÅ )
long-lasting, has previously been observed (Saxton expressed as a percentage of pre-exercise strength. * Sig-
and Donnelly, 1996; Byrne et al., 2001) and may repre- ni® cantly diþ erent (P < 0.05) from pre-exercise.
Strength, power and fatigue after eccentric exercise 955

(Fig. 3a). One day after eccentric exercise, the initial iso- common observation during the last 15 s of the tests
metric force was reduced at both 40° and 80° (Fig. 3b). after eccentric exercise. This is clearly evident in Fig. 3b,
The rate of fatigue was still greater at 80° (b = -4.58) where force intermittently dropped to zero as the
than at 40° (b = -1.40); however, at both 40° and 80°, participant could no longer continue the exercise. The
the knee extensors were less fatiguable than before participant’ s ability to quickly overcome this fatigue
eccentric exercise, which was most evident at 80°. Thus, and generate force suggests that central fatigue was
after eccentric exercise, the knee extensors became responsible.
weaker but less fatiguable ± that is, there was an
improved ability to maintain force. Task failure was a
Isometric strength at 80° knee ¯ exion versus Wingate peak
power
A signi® cant main eþ ect was shown for time (F5,30 =
20.4, P < 0.01) and, although the main eþ ect for mode
was not signi® cant (F1,6 = 2.6, P > 0.05), the interaction
of time and mode was (F5,30 = 4.8, P < 0.05). Figure 4
illustrates the eþ ect of eccentric exercise on the ability to
generate isometric force and Wingate peak power. The
interaction of time and mode was the result of initial
diþ erences (1 h after exercise) of the eþ ect of eccentric
exercise on isometric force (30% reduction) and
Wingate peak power (13% reduction) and subsequent
diþ erences in the pattern of recovery. Isometric force
followed a linear recovery pattern at day 1 (26%) and
day 2 (19%), whereas peak power suþ ered further
decrements at day 1 (18%) and day 2 (16%) before
starting to recover.
Fatigue during the Wingate test was reduced at all
Fig. 2. Fatiguability of the knee extensors during 60 s MVCs
instants in time, but only signi® cantly so 1 day after
at 40° (short; solid bars) and 80° (optimal; open bars) knee
¯ exion before and after eccentric exercise-induced muscle
exercise (F5,30 = 5.3, P < 0.05). Values for Wingate
damage. Fatiguability is expressed as the regression coeý cient fatigue were -12.75 ± 2.26 before eccentric exercise,
(b); the less negative the coeý cient, the less fatiguable -10.0 ± 2.03 at 1 h after, -8.33 ± 2.01 at 1 day after,
the muscles. * Signi® cantly diþ erent (P < 0.05) from 40°. -8.24 ± 1.88 at 2 days after, -8.97 ± 1.72 at 3 days
** Signi® cantly diþ erent (P < 0.05) from pre-exercise. after and -10.60 ± 1.95 at 7 days after exercise. Figure 5

Fig. 3. (a) Pre-eccentric exercise data for one participant showing the rate of fatigue during the 60 s MVC fatigue test at 40°
(solid circles) and 80° (open circles) knee ¯ exion. Regression lines ® tted to the data represent b = -1.84 at 40° and b = -7.68 at
80°. (b) Data for the same participant 1 day after eccentric exercise-induced muscle damage, showing the rate of fatigue
during the 60 s fatigue test at 40° (solid circles) and 80° (open circles) knee ¯ exion. Regression lines ® tted to the data represent
b = -1.40 at 40° and b = -4.58 at 80°.
956 Byrne and Eston

Fig. 6. Creatine kinase activity after eccentric exercise-

induced muscle damage (mean ± sxÅ ). *Signi® cantly diþ erent
Fig. 4. Changes in isometric MVC strength at 80° knee (P < 0.05) from pre-exercise values.
¯ exion (solid bars) and Wingate peak power (open bars) after
eccentric exercise-induced muscle damage. Values are means
Creatine kinase activity
(± sxÅ ) expressed as a percentage of pre-exercise values. * Sig-
ni® cantly diþ erent (P < 0.05) from pre-exercise. ** Signi® cant Figure 6 shows that creatine kinase activity was elevated
diþ erence (P < 0.05) between strength and power. (F5,30 = 3.3, P < 0.05) above baseline 1 h and 1 day after
exercise.

Discussion

Evidence of muscle damage

The eccentric exercise protocol produced changes in
all muscle function variables. Isometric strength at 80°
and Wingate peak power were reduced by 30% and
13% 1 h after exercise, respectively, and only returned
to pre-exercise values 7 days later. During sustained
maximal voluntary contractions and the Wingate test,
the knee extensors were signi® cantly less fatiguable for
3 days and 1 day after eccentric exercise, respectively.
Fig. 5. Data for one participant showing the decline in power Creatine kinase activity was signi® cantly elevated 1 h
output during the 30 s Wingate test before (open circles) and after exercise, reached a peak at day 1 and then gradually
1 day after (solid circles) eccentric exercise.
returned to baseline values. The immediate and sus-
tained loss of muscle function and the increase in blood
displays the response of one participant to the Wingate levels of creatine kinase are common indirect markers of
test before and 1 day after eccentric exercise. Similar exercise-induced muscle damage (Warren et al., 1999).
to the isometric responses in Fig. 3, the most striking
diþ erence between pre- and post-exercise responses
Potential mechanisms of muscle function impairment
is the reduction in force-generating capacity at the
start of exercise. Power output could be well main- The results of the isometric and dynamic parts of the
tained, albeit at lower than pre-exercise values. Post- study suggest that an inability to generate high force
Wingate blood lactate concentration did not change and power at the start of exercise is the common under-
signi® cantly (F5,30 = 1.2, P > 0.05) after eccentric lying mechanism responsible for the observed changes
exercise. The post-Wingate blood lactate concentra- in strength, power and fatigue after eccentric exercise.
tions were 10.6 ± 0.6 mmol ´l-1 before eccentric exercise The inability to generate an initial high force and power
and 11.1 ± 1.2, 9.6 ± 1.0, 9.7 ± 1.0, 9.1 ± 0.7 and 10.6 ± will depend on events that occur at or distal to the neuro-
1.2 mmol´ l-1 1 h, 1 day, 2 days, 3 days and 7 days after muscular junction (peripheral fatigue) and on a failure
exercise, respectively. to activate the muscles voluntarily (central fatigue).
Strength, power and fatigue after eccentric exercise 957

A failure to activate intact force-generating structures could explain the observed inability of eccentrically
because of excitation± contraction coupling impairment exercised muscle to generate an initial high force and
and disruption of force-generating or transmitting power and, therefore, appear less fatiguable.
structures are considered to be the two main peripheral
mechanisms responsible for the loss of maximal volun-
Central fatigue
tary force after eccentric exercise (for reviews, see
Morgan and Allen, 1999; Warren et al., 2001). Although The results revealed an immediate and long-lasting
full voluntary activation has been demonstrated during reduction in the ability to generate isometric force and
brief isometric MVCs after eccentric exercise (Ruther- dynamic peak power. One hour after eccentric exercise,
ford et al., 1986; Saxton and Donnelly, 1996), it is the ability to generate isometric force was aþ ected
doubtful whether this is true for sustained isometric and to a greater extent (69.8 ± 2.6% of pre-exercise values)
dynamic muscle actions. Bigland-Ritchie et al. (1978) than Wingate peak power (87.3 ± 1.7%). The two
reported that central fatigue consistently accounted for parameters then followed diþ erent temporal patterns of
up to 30% of the force loss during sustained 60 s MVCs recovery; force recovered linearly, whereas peak power
in fresh muscle and with well-motivated participants. declined further at days 1 and 2 before starting to
Whether the central contribution to fatigue increased recover. Wingate peak power eventually recovered with
after eccentric exercise could not be determined; how- a similar time course to that of isometric strength.
ever, the increased incidence of task failure (see Fig. 3b) This bimodal pattern of muscle function recovery is in
would suggest that this is the case. The relative con- line with the results of previous studies (Sargeant and
tribution of central and peripheral mechanisms to the Dolan, 1987; MacIntyre et al., 1996) and has been
changes in muscle function observed in the present attributed to the in¯ ammatory response that accom-
study could not be determined, although a discussion of panies muscle damage (Sayers and Clarkson, 2001). In
potential mechanisms is warranted. addition to selective damage of type II ® bres, reductions
in central drive resulting from voluntary conscious or
subconscious re¯ ex mechanisms could also explain the
Selective type II ® bre damage
reductions in peak power and the temporal pattern of
The knee extensors were signi® cantly less fatiguable recovery. The reductions in peak power following the
during MVCs and Wingate tests for 3 days and 1 day initial (1 h) decrement occurred during the typical
after eccentric exercise, respectively. This is in line with arrival and peak (24± 48 h) of delayed-onset muscle
the ® ndings of Balnave and Thompson (1993), who soreness (Clarkson et al., 1992), which also accom-
raised the question: How can a muscle be less fatiguable panies muscle damage. Voluntary eþ ort may have been
after a bout of prolonged eccentric exercise than it was reduced at these times because of painful sensations or,
before? Research into fatigue mechanisms during alternatively, aþ erent signals from the active muscles
maximal exercise and selective ® bre changes that occur and tendons may have produced re¯ ex inhibition. An
in eccentrically exercised muscle help to answer this additional factor to consider is that measures of motor
question. The loss of force during isometric muscle control, such as force and position sense, are impaired
actions (Soderlund et al., 1992) and power output after eccentric exercise (Saxton et al., 1995; Brockett
during maximal cycling (Nevill et al., 1996) has been et al., 1997) and are more likely to aþ ect a complex
attributed to declining rates of phosphocreatine and dynamic motor task such as cycling to a greater extent
glycogen utilization in type II muscle ® bres. Evidence than a simple static task such as knee extension.
suggests that type II ® bres are selectively damaged
during eccentric exercise (Friden et al., 1983; Jones
Metabolic dysfunction
et al., 1986) and this is possibly due to selective type II
recruitment (Nardone et al., 1989; Enoka, 1996; Asp The role of metabolic dysfunction after eccentric
et al., 1998). If type II ® bre function is preferentially exercise is unclear, but defects are likely to aþ ect force-
aþ ected by eccentric exercise, for example because of generating capacity. The rate of energy release through
excitation± contraction coupling impairment, damage anaerobic sources is of critical importance for the
to the force-generating or transmitting structures, or development and maintenance of high force and power
selective glycogen depletion, then the contribution of output during maximal-intensity exercise (Nevill et al.,
this ® bre type to the post-eccentric exercise fatigue 1996). Reductions in high-energy phosphate have been
responses will be diminished. As such, the fatigue reported to occur 24 h after eccentric exercise in rats
responses will be without the marked rise and rapid (van der Meulen et al., 1992) and delayed muscle glyco-
decline in force and power that is associated with the gen replenishment after damaging exercise in humans
activation and rapid fatigue of type II ® bres and will, is well documented (O’ Reilly et al., 1987; Costill et al.,
therefore, appear fatigue-resistant. Such a mechanism 1990; Asp et al., 1998). It is possible that reduced
958 Byrne and Eston

concentrations of ATP and phosphocreatine, as well as exposed to prior eccentric exercise. Journal of Physiology,
glycogen depletion, in the days after eccentric exercise 509, 305± 313.
compromised the ability to generate and maintain force Balnave, C.D. and Thompson, M.W. (1993). Eþ ect of training
and power. However, evidence from recent animal on eccentric exercise-induced muscle damage. Jour nal of
(Warren et al., 2000) and human (Rawson et al., Applied Physiology, 75, 1545± 1551.
Bar-Or, O. (1987). The Wingate anaerobic test: an update
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