Eur J Appl Physiol (2012) 112:3585–3595
DOI 10.1007/s00421-012-2339-3
O R I G I N A L A R T I CL E
Early-phase muscular adaptations in response to slow-speed
versus traditional resistance-training regimens
Mark D. Schuenke · Jennifer R. Herman · Roger M. Gliders ·
Fredrick C. Hagerman · Robert S. Hikida · Sharon R. Rana ·
Kerry E. Ragg · Robert S. Staron
Received: 6 December 2011 / Accepted: 27 January 2012 / Published online: 12 February 2012
© Springer-Verlag 2012
Abstract Thirty-four untrained women participated in a
6-week program to investigate slow-speed versus “normal”
speed resistance-training protocols. Subjects were divided
into: slow-speed (SS), normal-speed/traditional-strength
(TS), normal-speed/traditional muscular endurance (TE),
and non-exercising control (C) groups. Leg press, squats,
and knee extensions were performed 2 days/week for the
Wrst week and 3 days/week for the remaining 5 weeks
(»2 min rest). The SS group performed 6–10 repetitions
maximum (6–10RM) for each set with 10 s concentric
(con) and 4 s eccentric (ecc) contractions. The TS and TE
groups performed sets of 6–10RM and 20–30RM, respec-
tively, at “normal” speed (1–2 s/con and ecc contractions).
TE and SS trained at the same relative intensity (»40–60%
1RM), whereas TS trained at »80–85% 1RM. Pre- and
post-training muscle biopsies were analyzed for Wber-type
composition, cross-sectional area (CSA), and myosin heavy
Communicated by William J. Kraemer .
M. D. Schuenke (&)
Department of Anatomy, College of Osteopathic Medicine,
University of New England, Biddeford, ME 04005, USA
e-mail: mschuenke@une.edu
J. R. Herman
Department of Biomedical Sciences, Rocky Vista University
College of Osteopathic Medicine, Parker, CO 80134, USA
R. M. Gliders · S. R. Rana
School of Allied Health Sciences and Wellness,
College of Health Sciences and Professions,
Athens, OH 45701, USA
F. C. Hagerman · R. S. Hikida · K. E. Ragg · R. S. Staron
Department of Biomedical Sciences,
Heritage College of Osteopathic Medicine,
Ohio University, Athens, OH 45701, USA
chain (MHC) content. The percentage of type IIX Wbers
decreased and IIAX increased in all three training groups.
However, only TS showed an increase in percentage of type
IIA Wbers. CSA of Wber types I, IIA, and IIX increased in
TS. In SS, only the CSA of IIA and IIX Wbers increased.
These changes were supported by MHC data. No signiW-
cant changes for any parameters were found for the C
group. In conclusion, slow-speed strength training induced
a greater adaptive response compared to training with a
similar resistance at “normal” speed. However, training
with a higher intensity at “normal” speed resulted in the
greatest overall muscle Wber response in each of the vari-
ables assessed.
Keywords Human skeletal muscle · Fiber types ·
Histochemistry · Hypertrophy · Myosin heavy chains ·
Slow-speed training
Introduction
Resistance training appears to be the most eVective means
of inducing hypertrophy, but it is not clear what speciWc
protocol is best for optimizing this adaptation. Tradition-
ally, regimens using relatively high intensity [»70–85% of
one repetition maximum (1RM)] are prescribed for opti-
mizing muscle hypertrophy and strength in novice and
intermediately trained individuals (Campos et al. 2002;
Kraemer et al. 2002). However, in addition to training
intensity, other factors such as speed of contraction may
have an impact on muscular adaptations. Slowing the con-
centric (con) and/or eccentric (ecc) phases of muscle con-
traction increases the duration under tension per repetition.
For example, Shepstone et al. (2005) compared two diVer-
ent isokinetic speeds (fast = 3.66 rad/s and slow = 0.35 rad/s)
123
3586
in an 8-week study. Both groups developed signiWcant
hypertrophy and signiWcantly decreased myosin heavy
chain (MHC) IIx content. However, these changes were
signiWcantly greater in the fast-trained group, despite the
slow-contracting muscles being under tension approxi-
mately 10 times longer. Similarly, Munn et al. (2005)
reported an 11% greater increase in 1RM strength follow-
ing 6 weeks of fast (2.44 rad/s) isokinetic training com-
pared to slow (0.87 rad/s).
Although studies using isokinetic training add valuable
information to our body of resistance-training knowledge,
isokinetics are not practical for everyday exercise due to the
special equipment involved. In addition, isokinetics do not
mimic normal movement and do not result in optimal mus-
cle adaptations compared to isotonic/dynamic resistance
training (Kovaleski et al. 1995; Remaud et al. 2009). In the
Wrst known study to investigate the eVects of velocity-spe-
ciWc training using isotonic/dynamic rather than isokinetic
exercises, Morrissey et al. (1998) compared slow (2 s con/
2 s ecc) and fast (1 s con/1 s ecc) resistance training. Fol-
lowing 21 training sessions, there was no signiWcant diVer-
ence in the percentage of improvement between the training
groups for isotonic 1RM squat testing. However, for isoki-
netic post-testing, the fast-trained group signiWcantly
increased strength when tested at fast and slow velocities,
whereas the slow-trained group did not signiWcantly
improve strength at either test velocity. The authors sug-
gested that fast training resulted in a greater increase in fast
Wber-type area, relative to slow training, thereby giving the
fast-training group an advantage in velocity-speciWc test-
ing. However, the authors did not analyze any muscle tissue
to substantiate this hypothesis.
Subsequent low-velocity resistance training studies
have slowed down the repetitions to what has sometimes
been referred to as “super slow” resistance training (Keeler
et al. 2001; Westcott et al. 2001; Hunter et al. 2003; Neils
et al. 2005). “Super slow” resistance training typically
involves a 10 s concentric contraction followed by a 4–5 s
eccentric contraction. This type of slow repetition training
has been purported to improve aerobic capacity to a
greater extent than traditional-strength or muscular endur-
ance resistance training (Hutchins 1992). The assertion
being that a longer contraction time places demand on both
aerobic and anaerobic energy sources. To date, studies
comparing “super slow” to traditional resistance-training
protocols have produced equivocal results in strength
gains with reports of no diVerence (Neils et al. 2005),
greater gains for slow speed (Westcott et al. 2001), and
greater gains for traditional speed (Keeler et al. 2001). In a
study from our laboratory (Rana et al. 2008), 6 weeks of
traditional resistance training (1–2 s con/1–2 s ecc) pro-
duced superior strength gains compared to low-velocity
training (10 s con/4 s ecc).
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Eur J Appl Physiol (2012) 112:3585–3595
Limited data exist on the eVects of slow-speed dynamic
resistance training and actual adaptations within the mus-
cle. To our knowledge, only two studies (Gillies et al.
2006; ClaXin et al. 2011) have utilized muscle biopsies to
assess the muscular eVects of varying contraction times
during dynamic resistance training. Neither of these
investigations compared “super slow” to “normal” speed
resistance training and neither had a control group. The
purpose of the present study was to determine the muscu-
lar eVects [Wber-type composition and cross-sectional area
(CSA)] of slow-speed (10 s con/4 s ecc) compared to nor-
mal-speed (1–2 s con/1–2 s ecc) resistance training in pre-
viously untrained individuals using a combination of
histochemical and immunohistochemical methods. To our
knowledge, this is the Wrst study investigating slow- ver-
sus normal-speed strength training of varying intensities
utilizing speciWc methods to elucidate the entire range of
Wber types.
Methods
Subjects
Thirty-four healthy, untrained women (21.1 § 2.7 years)
volunteered to participate in the present study. The subject
population consisted of exclusively females, because in pre-
vious studies, it has been easy to Wnd eager, untrained female
subjects with excellent compliance to protocols. All subjects
were informed of the procedures, risks, and beneWts, and
signed an informed consent document approved by the Ohio
University Institutional Review Board before participation.
Prior to inclusion, complete medical histories and a physical
examination (including a thorough musculoskeletal screen-
ing) were performed. Although healthy, all subjects were
considered untrained and had not participated in a regular
exercise program for at least 6 months prior to the start of the
study. The women were randomly divided into four groups:
normal speed/traditional strength (TS, n = 9) (20.6 § 1.9
years, 165.6 § 4.9 cm), normal speed/traditional local mus-
cular endurance (TE, n = 8) (22.3 § 3.9 years, 161.9 §
8.3 cm), slow speed (SS, n = 10) (19.4 § 1.3 years, 168.0 §
4.2 cm), and non-exercising control (C, n = 7) (22.9 § 2.4
years, 163.6 § 4.5 cm).
Anthropometric assessments
Anthropometric measurements (total body mass, estimated
fat-free mass, and estimated percentage body fat) were
determined before and after the 6-week training period
(Table 1). Skinfold measurements were obtained from three
sites (anterior thigh, posterior brachium, and suprailiac)
prior to extraction of the pre- and post-study muscle
Eur J Appl Physiol (2012) 112:3585–3595 3587

Table 1 Anthropometric data Strength training protocols

C TE TS SS
The exercising subjects participated in a 6-week resistance-
Body mass (kg) training program targeting the quadriceps femoris muscle
Pre 72.5 § 15.0 64.7 § 8.3 64.1 § 7.9 63.9 § 8.4 group. Workouts were performed 2 days/week for the Wrst
Post 72.1 § 14.7 64.9 § 8.9 64.6 § 8.2 63.9 § 8.5 week and 3 days/week for the remainder of the study
Body fat (%) (6 weeks/17 total training sessions). The SS and TE groups
Pre 25.6 § 4.8 24.2 § 4.2 23.4 § 3.3 21.0 § 3.1 were designed to be equal in intensity (»40–60% 1RM),
Post 25.6 § 5.2 24.3 § 4.6 22.7 § 3.2 21.1 § 2.9 whereas the TS group trained at »80–85% 1RM. Three sets
Fat mass (kg) of each exercise were performed in the Wxed order of leg
Pre 19.2 § 7.9 15.8 § 4.6 15.2 § 3.6 13.6 § 3.5a press, squat, and knee extension. During the entire training
Post 19.1 § 8.1 16.0 § 4.5 14.9 § 3.6 13.6 § 3.3a period, no variation or supplementary exercises were per-
Fat-free mass (kg) formed. The SS group performed 6–10RM for each set with
Pre 53.3 § 7.3 48.9 § 5.1 49.0 § 4.6 50.3 § 5.5 10 s con/4 s ecc contractions for each repetition, TS and TE
Post 53.0 § 6.7 49.0 § 5.0 49.7 § 4.8 50.3 § 5.7 performed 6–10RM and 20–30RM, respectively, for each
a
set with 1–2 s con/1–2 s ecc contractions. Thus, both SS
SigniWcantly diVerent from respective value in C group (p < 0.05)
and TS trained within the same range of repetitions to fail-
ure (6–10RM). A verbal count was provided to ensure that
biopsies and were used in the equation proposed by Jackson the duration of muscular contraction was exactly as pre-
et al. (1980) for body composition analysis of women. scribed. If at any point a subject was able to perform the
maximal number of repetitions (e.g. 10 reps for SS and TS;
Maximal strength and endurance tests 30 reps for TE) for all three sets of a given exercise, the
weights for that exercise were increased to ensure that sub-
Prior to beginning the study, all subjects (including the con- jects performed the appropriate number of repetitions. The
trols) participated in an orientation session for familiariza- training subjects performed repetitions until failure and
tion with the equipment and exercises. During this session, rested »2 min between sets and exercises. All subjects
proper lifting technique was demonstrated and practiced for were supervised, monitored for proper technique, and ver-
each of the three lower limb exercises (leg press, squat, and bally encouraged throughout each training session. Work-
knee extension). A goniometer was initially used to deter- outs began and ended with 10–15 min of calisthenics,
mine each subject’s correct depth set point for the leg press stretching, and low-intensity cycling or rowing.
and squat exercises. Both maximal dynamic strength
(1RM) and local muscular endurance (maximum number of Muscle biopsies
repetitions performed with 60% of 1RM) were assessed for
each of the exercises at the beginning and end of the study. Biopsies (80–160 mg) were extracted from the superWcial
Because of the exhaustive nature of the endurance test, the region of the vastus lateralis muscle, using the percutaneous
maximal strength test was always performed Wrst. After muscle biopsy technique (Bergström 1962). To ensure ade-
warming up (one set of 8–10 repetitions with 40–50% of quate sample sizes, large biopsy specimens were obtained
the predicted 1RM and one set of 3–4 repetitions with 60– using a “double-chop” method (Staron et al. 1990) com-
70% of the predicted 1RM), the load was set at 90% of the bined with suction (Evans et al. 1982). Samples were
predicted 1RM and was increased after each successful lift obtained approximately 16 cm proximal to the superior bor-
until failure (Staron et al. 1990). Periods of approximately der of the patella. Because of possible variations in Wber-
4–5 min rest were allotted between each attempt to ensure type distribution from superWcial to deep and proximal to
recovery. A test was considered valid if the subject used distal (Blomstrand and Ekblom 1982), pre- and post-biop-
proper form and completed the lift in a controlled manner sies were extracted in proximity (»1 cm) using the pre-
through the entire range of motion without assistance. Once biopsy scar and depth markings on the needle (Staron et al.
the 1RM was determined, 60% of this value was calculated 1990). The muscle samples were oriented in tragacanth
for the local muscular endurance test. After a suYcient gum, immediately frozen in isopentane cooled by liquid
recovery period (»10 min), the subjects performed as many nitrogen to ¡159°C, and stored at ¡80°C until further anal-
repetitions as possible with 60% of 1RM until failure. Spe- yses could be performed. Of the 34 subjects, 31 success-
ciWc strength data for this study have been previously fully donated both pre- and post-biopsy samples (C, n = 7;
reported (Rana et al. 2008). TS, n = 9; SS, n = 9; TE, n = 6).

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3588 Eur J Appl Physiol (2012) 112:3585–3595

Muscle Wber typing
The frozen biopsy specimens were thawed to ¡24°C and
serially sectioned (12 m) for histochemical and (6 m)
immunohistochemical analyses. Routine myoWbrillar
adenosine triphosphatase (mATPase) histochemistry was
performed using preincubation pH values of 4.3, 4.6, and
10.0 (Fig. 1) (Guth and Samaha 1969; Brooke and Kaiser
1970). Immunohistochemical methods were also utilized
to assist in Wber-type delineation. Serial cross sections
were incubated in one of the following mouse IgG mono-
clonal antibodies (mAb) (University of Iowa Develop-
mental Studies Hybridoma Bank): A4.951 (anti-MHCI),
N2.261 (anti-MHCI + anti-MHCIIa), and A4.74 (anti-
MHCIIa + anti-MHCIIx) (Fig. 2). Sections were then pre-
pared using VECTASTAIN® Elite ABC (avidin–biotin
peroxidase complex) kits (Vector Laboratories, Burlin-
game, CA). Both the mATPase histochemistry and MHC
immunohistochemistry were combined to determine the
overall Wber-type composition. A direct comparison of

Fig. 1 mATPase histochemical Wber typing. Representative staining patterns of Wbers at pH 4.3 (a), pH 4.6 (b), pH 10.0 (c). Table represents gen-
eral schematic indicating relative staining intensities

Fig. 2 MHC immunohistochemical Wber typing. Representative immunostaining patterns of Wbers with A4.591 (a), N2.261 (b), A4.74 (c). Table
represents general schematic indicating relative staining intensities

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Eur J Appl Physiol (2012) 112:3585–3595
both preparations was made for each biopsy using the six
serial sections (3 for histochemistry and 3 for immunohis-
tochemistry). Six Wber types (I, IC, IIC, IIA, IIAX, and
IIX) were distinguished based on their respective staining
intensities (see Figs. 1, 2). Human fast Wber types IIAX
and IIX (Smerdu et al. 1994; Ennion et al. 1995) have
been previously referred to as IIAB and IIB, respectively
(e.g. Campos et al. 2002). Although the two methods for
Wber typing were in agreement for the identiWcation of the
majority of Wbers, the combination of methods was espe-
cially important for the appropriate classiWcation of type
IIAX Wbers.
Fiber CSA measurements
The NIH Imaging software program (version 1.62) was uti-
lized to obtain CSA measurements from approximately
»300 Wbers per muscle sample with at least 50 Wbers per
each major Wber type (I, IIA, IIX). All area measurements
were made by one investigator on samples prepared for
mATPase histochemistry following preincubation at pH
4.6. With this preparation, Wber borders were distinct and
there were no indications of Wber shrinkage. In addition,
Wber-type area percentages for the major Wber types (I, IIA,
and IIX) were determined by combining Wber-type CSA
and Wber-type percentage data using the formulae:
I = I + IC, IIA = IIC + IIA + ½IIAX, IIX = IIX + ½IIAX
(Fry et al. 1994; Staron et al. 2000).
MHC analysis
Myosin heavy chain analyses were performed on the biopsy
samples using sodium dodecyl sulfate (SDS)-polyacryl-
amide electrophoretic techniques (Fig. 3). The protocol for
analyzing the specimens was based on the procedures of
Perrie and Bumford (1986) with modiWcations used for sin-
gle Wber analysis (Staron 1991; Staron and Hikida 1992).
BrieXy, six to ten serial cross sections (12 m thick) from
each biopsy were placed into 0.5 ml of a lysing buVer con-
taining 10% (wt/vol) glycerol, 5% (vol/vol) 2-mercap-
toethanol, and 2.3% (wt/vol) SDS in 62.5 mM tris
(hydroxymethyl) aminomethane HCl buVer (pH 6.8), and
were heated for 10 min at 60°C. Small amounts of the
Fig. 3 Myosin heavy chain (MHC) content as determined by SDS-PAGE analysis of muscle biopsy samples obtained pre- and post-study from a
representative subject in each group (TS, SS, C, TE). Note the decrease in MHCIIx at the end of the study in all three training groups
3589
extracts (7.5 l) were loaded on 4–8% gradient SDS-poly-
acrylamide gels with 4% stacking gels (Bar and Pette
1988), run overnight (20–24 h) at 120 V, and stained with
Coomassie Blue. MHC isoforms were identiWed according
to their apparent molecular masses compared with those of
marker proteins and migration patterns from single Wber
analyses. Relative MHC isoform content was subsequently
determined using a laser densitometer. These data were
then compared with calculated percent area occupied by the
major Wber types.
Statistical analyses
The statistical package SigmaStat (version 2.03) was utilized
for all analyses. Descriptive statistics were used to derive
mean values § standard deviations (SD) for all variables. A
two-way analysis of variance (ANOVA) with repeated mea-
sures (2 £ 4 design: 2 time points by 4 groups) was used to
determine diVerences within group (pre-post) and between
groups (C, TS, SS, TE) for each variable. Tukey’s Honest
SigniWcant DiVerence (HSD) test was used for post hoc anal-
ysis where signiWcant diVerences were detected for main
eVects. A one-way ANOVA followed by Tukey’s HSD were
used to assess percent change in CSA. For all analyses,
diVerences were considered signiWcant at p · 0.05.
Results
Anthropometric assessment
There were no signiWcant within-group diVerences detected
between pre- and post-training values for any of the anthro-
pometric variables (Table 1). SigniWcant main eVects
between groups were detected for fat mass (FM) (p = 0.02)
and percent body fat (% BF) (p = 0.014). For FM, post hoc
analysis revealed a signiWcant diVerence between SS and C
within both time points (pre, p = 0.024; post, p = 0.027).
However, post hoc analysis revealed only a non-signiWcant
trend for SS % BF to diVer from C % BF within both time
points (pre, p = 0.061; post, p = 0.069). There were no
other signiWcant between-group diVerences detected for
any other anthropometric measurement.
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3590 Eur J Appl Physiol (2012) 112:3585–3595

Table 2 Muscle Wber-type percentages

Group Muscle Wber-type percentages n

I IC IIC IIA IIAX IIX

C
Pre 39.3 § 11.6 0.5 § 0.4 0.8 § 0.6 29.2 § 4.0 9.7 § 4.0 20.5 § 9.7 696 § 213
Post 42.7 § 8.7 1.5 § 2.2 1.1 § 1.0 27.8 § 7.8 7.5 § 1.4 19.4 § 9.2 1,109 § 535
TS
Pre 36.4 § 14.4 0.9 § 1.0 0.8 § 0.9 25.9 § 8.6 9.6 § 2.6 26.4 § 11.1 1,318 § 471
a a,b a
Post 37.8 § 12.8 1.4 § 1.3 2.0 § 2.2 33.5 § 9.7 13.7 § 2.6 11.6 § 7.2 1,250 § 337
SS
Pre 38.7 § 9.6 1.0 § 1.6 1.3 § 2.4 29.1 § 6.5 9.0 § 3.0 20.9 § 8.4 1,044 § 443
Post 41.4 § 12.7 2.2 § 3.1 5.1 § 3.9a,c 27.4 § 11.8 12.9 § 3.7a,b 11.0 § 7.0a 1,131 § 405
TE
Pre 41.6 § 9.7 0.1 § 0.2 0.7 § 1.3 26.9 § 4.4 9.3 § 2.0 21.4 § 8.5 1,234 § 443
Post 43.4 § 9.6 1.1 § 1.5 2.8 § 3.7 27.8 § 5.2 14.0 § 4.2a,b 10.9 § 5.0a 783 § 219
n = mean number of Wbers utilized per biopsy sample
a
SigniWcantly diVerent from pre-training value, within group (p < 0.01)
b
SigniWcantly diVerent from post-training value, from C group (p < 0.01)
c
SigniWcantly diVerent from post-training value, from TS and C groups (p < 0.05)

Fiber-type analysis eral values will be summarized here in addition to Wber

The combination of both mATPase histochemistry and
MHC immunohistochemistry was used in the overall deter-
mination of Wber-type composition using 1,105 § 383
Wbers per sample (Table 2; Figs. 1, 2). For both type IIX
and IIAX Wbers, a signiWcant training eVect and a signiW-
cant group by time interaction were detected. Post hoc anal-
yses revealed signiWcant within-group decreases in the
percentage of type IIX Wbers and increases in the percent-
age of type IIAX Wbers from pre- to post-training for all
three training groups (Table 2). No signiWcant between-
group diVerences were detected. In addition, the TS group
demonstrated a signiWcant within-group increase in the per-
cent type IIA Wbers from pre- to post-training (Table 2).
There was also a signiWcant diVerence detected within the
minor hybrid type IIC population. A signiWcant training
eVect and group by time interaction were found for the
percentage of type IIC Wbers. Post hoc analyses showed a
signiWcant within-group increase in this population from pre-
to post-training for the SS group. The overall training eVect
may also have been partially driven by a non-signiWcant
trend within the TE group to increase the percentage of type
IIC Wbers from pre- to post-training (p = 0.055).
Fiber CSA
Some studies have reported Wber hypertrophy data in terms
of change in general mean Wber area (m2) without specify-
ing Wber type. For the purpose of comparison, similar gen-
type-speciWc results. A two-way ANOVA with repeated
measures showed a signiWcant training eVect and a signiW-
cant group by time interaction for group mean Wber CSA.
Post hoc analyses isolated the signiWcant within-group
diVerences to the TS and SS groups. The mean Wber CSA in
the TS group increased by 38.8 § 21.7% (3,265 § 543 vs.
4,508 § 1,002 m2, p < 0.001) from pre- to post-training,
whereas SS increased by 10.6 § 8.7% (3,615 § 505 vs.
3,997 § 665 m2, p = 0.026). Tukey’s HSD test showed no
signiWcance between-group diVerences for either time
point. There were no within-group diVerences for the TE or
C groups.
For speciWc Wber types, there was a signiWcant training
eVect for type I Wber CSA and signiWcant group £ time
interaction, which post hoc testing isolated to the TS group
(Table 3). A one-way ANOVA performed on the percent
change in CSA by Wber-type between groups revealed a
signiWcant between-group diVerence. Tukey’s HSD
revealed the 26.6 § 22.7% increase in TS type I CSA to be
greater compared with all other groups: C (¡1.2 § 7.3%,
p = 0.007), TE (1.0 § 15.2%, p = 0.019), and SS (6.5 §
10.1%, p = 0.047). A two-way ANOVA with repeated
measures found a signiWcant training eVect for type IIA
Wber CSA and a signiWcant group £ time interaction.
Post hoc analysis showed that both TS and SS had
within-group diVerences from pre- to post-training (increased
type IIA CSA) (Table 3). A one-way ANOVA on the
percentage change in type IIA CSA revealed a signiW-
cant between-group diVerence. Tukey’s HSD showed the

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Table 3 Muscle Wber cross-sectional area Percent Wber-type area and MHC analyses
Group Fiber cross-sectional area (m2)
Analysis of calculated data on the percent Wber area occu-
Type I Type IIA Type IIX pied by each major Wber type (I, IIA, IIX) showed a signiW-
C cant overall training eVect for type IIX percent Wber area
Pre 3,923 § 1,002 4,428 § 1,168 3,221 § 983
and a signiWcant group £ time interaction. Follow-up testing
Post 3,854 § 935 4,284 § 793 3,405 § 1,046
revealed signiWcant decreases in the percent area occupied
by type IIX for both TS and SS from pre- to post-training
TS
(Table 4). In the TE group, the decrease in percent IIX Wber
Pre 3,342 § 379 3,693 § 663 2,695 § 666
a a area was non-signiWcant (p = 0.067). In addition, a two-
Post 4,218 § 834 4,839 § 735 3,696 § 825a
way ANOVA with repeated measures showed a signiWcant
SS
training eVect and group £ time interaction for the type IIA
Pre 3,837 § 554 3,879 § 920 2,843 § 583
percent Wber area. Tukey’s HSD showed that TS signiW-
Post 4,052 § 443 4,358 § 1152b 3,376 § 754b
cantly increased type IIA percent Wber area from pre- to
TE
post-training (Table 4). As expected (Fry et al. 1994), these
Pre 3,726 § 313 3,918 § 560 3,021 § 946
data correlated with the MHC isoform analyses. A two-way
Post 3,725 § 332 4,268 § 625 3,287 § 779
ANOVA with repeated measures on relative MHCIIx
a
SigniWcantly diVerent from pre-training value within group (p < 0.001) isoform content found a signiWcant training eVect and
b
SigniWcantly diVerent from pre-training value within group (p < 0.05) group £ time interaction, which post hoc analysis revealed
to be signiWcant diVerences within the TS, SS, and TE
groups. From pre- to post-training, MHCIIx decreased in
32.9 § 20.4% increase in TS type IIA CSA to be greater all three training groups (TS, SS, and TE), and MHCIIa
compared to all other groups: C (¡1.3 § 15.0%, p < 0.001), increased in the TS group (Fig. 3; Table 4).
TE (9.5 § 12.0%, p = 0.037), and SS (12.3 § 11.3%,
p = 0.04). Results of the two-way ANOVA with repeated
measures showed a signiWcant training eVect and group £ Discussion
time interaction for type IIX Wber CSA. Tukey’s post hoc
revealed that both the TS and SS groups had within-group Anthropometric assessment
diVerences from pre- to post-training (increased type IIX
CSA); (Table 3). A one-way ANOVA on percent change in This investigation revealed no signiWcant changes in body
type IIX CSA revealed signiWcant between-group diVer- composition following 6 weeks of lower extremity resis-
ences (p = 0.036), which post hoc testing isolated between tance training. Although the SS group diVered from the
TS and C. The 41.1 § 32.7% increase in TS type IIX CSA controls in the present study with regard to fat mass, this
was greater compared to 6.7 § 21.0% in C (p = 0.038). diVerence was consistent and evident for the pre- and

Table 4 Relative percentage of MHC isoform content and percent Wber-type area
Group Relative % MHC isoform content Muscle Wber-type % area

MHCI MHCIIa MHCIIx I IIA IIX

C
Pre 40.7 § 16.4 36.6 § 11.3 22.7 § 13.3 39.7 § 12.0 39.1 § 8.3 21.2 § 10.1
Post 44.8 § 8.6 34.1 § 12.1 21.0 § 14.4 43.5 § 5.3 36.4 § 9.2 20.2 § 9.6
TS
Pre 37.9 § 16.2 34.4 § 14.4 27.7 § 12.6 38.4 § 15.7 35.9 § 10.7 25.7 § 10.2
Post 38.4 § 14.7 43.7 § 11.6a 17.9 § 9.7a 37.5 § 12.9 46.7 § 9.8a 15.7 § 7.1a
SS
Pre 42.0 § 13.9 35.8 § 8.4 22.2 § 9.5 42.4 § 12.4 37.2 § 6.6 20.4 § 8.7
a
Post 47.2 § 16.3 38.9 § 13.3 13.9 § 8.0 44.1 § 14.6 41.3 § 11.3 14.6 § 6.6a
TE
Pre 42.1 § 12.6 33.7 § 8.5 24.2 § 9.2 43.4 § 10.0 35.3 § 6.3 21.3 § 7.5
Post 45.9 § 12.2 40.0 § 9.1 14.1 § 6.7a 43.5 § 10.3 41.6 § 7.4 14.9 § 4.7
a
SigniWcantly diVerent from pre-training value within group (p < 0.5)

123
3592
post-training assessments. These Wndings are in agreement
with numerous other studies with regard to body composition
following short-term resistance training (e.g., Campos et al.
2002). Of the limited research available on slow-speed
resistance training, no signiWcant alterations in body com-
position have been reported (Keeler et al. 2001; Neils et al.
2005).
Fiber-type composition and MHC content
All three resistance-training protocols in the present study
resulted in a signiWcant decrease in the percentage of both
type IIX Wbers and MHCIIx with a concomitant increase in
the percentage of hybrid type IIAX Wbers. In addition, the
signiWcant increase in the percentage of type IIA Wbers (and
concomitant increase in % MHCIIa) following normal-
speed, high-intensity training (TS group) suggests addi-
tional transitions of IIX ! IIAX ! IIA. These Wndings are
consistent with resistance-training programs of varying
intensities and speeds (e.g., Staron et al. 1994; Kraemer
et al. 1995; Green et al. 1998; Campos et al. 2002). How-
ever, to the authors’ knowledge, the present study is the
Wrst to demonstrate these Wber-type transitions following a
“super slow” dynamic resistance-training regimen. The
most comparable study investigating the muscular eVects of
slow-speed dynamic resistance training is that of Gillies
et al. (2006) who demonstrated that training with increased
time under tension (6 s con/2 s ecc contractions; 6–8RM;
9 weeks), somewhat similar to the SS group, resulted in a
signiWcant decrease in MHCIIx and a concomitant increase
in MHCIIa content. However, that study did not report a
change in Wber-type proportions, because the pre-training
samples contained almost no (<0.05%) type IIX Wbers and
type IIAX were not delineated (Gillies et al. 2006). Further-
more, there was no control group and all of the subjects
were pre-strength trained.
Fiber CSA
In the present investigation, resistance training at both nor-
mal speed/high intensity (»80–85% 1RM) and slow speed/
low intensity (»45–60% 1RM) caused signiWcant increases
in the overall Wber CSA from pre- to post-training (disre-
garding Wber types). No changes in mean Wber CSA were
found after training at normal speed, low intensity (»45–
60% 1RM). Although high-intensity training had an overall
increase of 38.8% in Wber CSA compared to a 10.6%
increase for slow-speed training, these changes were not
statistically diVerent from each other. However, diVerences
in the adaptive responses between slow-speed and high-
intensity training were more apparent when speciWc Wber
types were considered. Slow-speed resistance training
increased the CSA of the fast Wbers IIA and IIX, whereas
123
Eur J Appl Physiol (2012) 112:3585–3595
high-intensity training signiWcantly increased the CSA of
all three major Wber types (I, IIA, and IIX). In addition, the
percent increase in CSA of types I and IIA of the high-
intensity group was signiWcantly greater than that of the
slow-speed group. The diVering degrees of change observed in
Wber size for the three training groups in the present study
paralleled the changes in strength for these same subjects
reported by Rana et al. (2008).
The present Wndings are somewhat similar to the results
of Gillies et al. (2006) who reported an increase in the CSA
of types I and IIA following resistance training with a long
concentric phase. However, the present Wndings contradict
other recent articles on low-velocity training. ClaXin et al.
(2011) have suggested both high- (leg press 111 § 31°/s)
and low-velocity (leg press 31 § 3°/s) dynamic resistance
training to elicit a similar hypertrophic response in fast
Wbers. These Wndings are questionable, because changes in
CSA were estimated by averaging Wve measurements taken
along the length of permeabilized single Wber segments (20
Wbers total), and Wbers were divided into types I and II
based solely on shortening velocity (subtypes were not
delineated). Using MRI, Tanimoto and Ishii (2006) also
reported similar hypertrophy in whole muscle CSA follow-
ing both low-intensity (»50% 1RM) slow-movement (3 s
con/3 s ecc) and high-intensity (»80% 1RM) normal-speed
resistance training. However, it is not possible to assess
Wber types or Wber transitions using MRI, so a direct com-
parison to the results of the present study cannot be done.
It is has been generally accepted that training load (i.e.,
% 1RM) has the most signiWcant impact on strength
and muscle hypertrophy (Deschenes and Kraemer 2002;
Kraemer et al. 2002; Kraemer and Ratamess 2004). A number
of investigations have reported hypertrophy in both men
and women in response to high-intensity resistance-training
programs that were comparable to TS in the present study
(e.g., Staron et al. 1990, 1991; Kraemer et al. 1995;
Campos et al. 2002). Unlike TS, the present Wndings indi-
cate no signiWcant increase of myoWber CSA (overall or for
speciWc Wber types) in response to low intensity/high repetition
training; these Wndings are in agreement with previous Wndings
(Campos et al. 2002; Tanimoto and Ishii 2006).
Interestingly, Burd et al. (2010) suggest that low-load/
high volume resistance training stimulates muscle protein
synthesis to a greater extent than high-load/low volume
training. Those Wnding (Burd et al. 2010) are in discord
with the present data in which neither low intensity proto-
col (TE or SS) resulted in as much hypertrophy as the TS
group. The present data indicate that SS training provides a
mild hypertrophic stimulus, relative to the TE protocol, but
the SS protocol does not provide an adequate enough
hypertrophic stimulus to pose it as an alternative to TS
training. Protein synthesis following a high repetition/light
load workout, even when done to failure, is not necessarily
Eur J Appl Physiol (2012) 112:3585–3595
predictive of protein accretion. Fiber hypertrophy relies on
a disproportionate protein synthesis relative to protein
breakdown over an extended period (reviewed in Spiering
et al. 2008). The study of Burd et al. (2010) did not report
rate of protein breakdown and data were collected on sin-
gle bout of exercise. Furthermore, in the present study, the
low velocity of contraction performed by SS necessitated
lifting a lower % 1RM compared to TS. Although it may
be intuitive that lifting at a slow speed provides a greater
challenge than lifting the same load faster, two papers have
summarized the impact of velocity on the number of repe-
titions that can be performed (HatWeld et al. 2006; Sakam-
oto and Sinclair 2006). Both studies demonstrated slower
velocities result in fewer repetitions, and thus, total train-
ing volume (kg £ repetitions £ sets) will be greater for
subjects who lift at a volitional velocity compared to a
slow velocity. Finally, based on the size principle
(reviewed in Cormie et al. 2011), training fast motor units
(e.g., those needed for TS-like training) is optimized at
near-maximal forces (Siegel et al. 2002) and faster veloci-
ties (Coyle et al. 1981). However, training with low veloc-
ity (e.g., SS in the current study; in Burd et al. 2010; in
HatWeld et al. 2006) utilizes asynchronous Wring of smaller
and/or slower motor units, and therefore is unlikely to
result in signiWcant training adaptations. Thus, a heavy
load stimulus appears to be vital for optimally stimulating
early-phase protein accretion of muscle Wbers in a resis-
tance-training program.
The present investigation made no measurements of
molecular-signaling markers, hormone or growth factor
levels, metabolites, or oxygen concentrations in response
to the diVerent training protocols, and thus, the authors
can only speculate as to the possible conditions induced
by SS training which may have resulted in a greater
hypertrophic response compared with that of TE. Hunter
et al. (2003) examined the metabolic eVects of an acute
bout of slow-speed versus traditional-strength training
and found that traditional training resulted in greater oxy-
gen consumption and mean heart rate compared to slow-
speed training. Post-exercise lactate levels were almost
twice as high following traditional-strength training com-
pared with slow speed, and the net energy expenditure of
the traditional group was 48% greater than the slow-
speed group (Hunter et al. 2003). Although this report
would indicate that slow-speed training does not provide
as great a metabolic challenge as training with a protocol
similar to TS in the present study, information is still
lacking as to the relative eVects between various speeds
of low-intensity training, such as between SS and TE.
This information would be helpful to understand the fac-
tors that may be responsible for the greater degree of
strength gains and Wber hypertrophy experienced by SS
compared with TE.
3593
Conclusions
The present study compared the muscular responses of
high- and low-intensity/normal-speed to low-intensity/
slow-speed training protocols in previously untrained, col-
lege-aged women. To our knowledge, no previous study of
slow-speed resistance training has analyzed muscle biop-
sies to determine the full extent of potential muscular adap-
tation in previously untrained individuals. The combination
of both histochemical and immunohistochemical Wber typ-
ing methods was particularly important in the delineation of
Wbers in transition between type IIX and IIA (hybrid type
IIAX). The primary Wnding of this study was that 6 weeks
of high-intensity/normal-speed resistance training opti-
mized muscular adaptations compared to low-intensity
resistance training of varying speeds. The diVering degrees
of change observed for the three training groups paralleled
the changes in both relative and absolute strength reported
on these same subjects (Rana et al. 2008). Although train-
ing at a high intensity (»80–85% 1RM) yielded greater
increases in Wber CSA compared with training at low inten-
sities (»45–60% 1RM), low-intensity/slow-speed training
resulted in a small, but signiWcant hypertrophic response
compared to low-intensity/normal-speed training. Although
this study recapitulated that high-intensity/normal-speed
resistance training is ideal for hypertrophic and strength
gains, low-intensity/slow-speed training may prove beneW-
cial to individuals for whom resistance training with a
heavy load is not advised.
Acknowledgments We wish to thank all those individuals who
assisted in supervising the training, and especially to the subjects who
volunteered and worked so hard throughout the study.
ConXict of interest The authors have no conXicts of interest to report.
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