(Journal of Strength & Conditioning Research) Schoenfeld BJ, Peterson MD, Ogborn D, Contreras B, Sonmez GT-[Article] Effects of Low- Versus High-Load Resistance Training on Muscle Strength and Hypertr
(Journal of Strength & Conditioning Research) Schoenfeld BJ, Peterson MD, Ogborn D, Contreras B, Sonmez GT-[Article] Effects of Low- Versus High-Load Resistance Training on Muscle Strength and Hypertr
(Journal of Strength & Conditioning Research) Schoenfeld BJ, Peterson MD, Ogborn D, Contreras B, Sonmez GT-[Article] Effects of Low- Versus High-Load Resistance Training on Muscle Strength and Hypertr
TE
EP
Dan Ogborn3
Bret Contreras4
Gul T. Sonmez1
C
C
TE
Abstract
The purpose of this study was to compare the effect of low- versus high-load resistance
training (RT) on muscular adaptations in well-trained subjects. Eighteen young men experienced
in RT were matched according to baseline strength, and then randomly assigned to 1 of 2
experimental groups: a low-load RT routine (LL) where 25-35 repetitions were performed per set
per exercise (n = 9), or a high-load RT routine (HL) where 8-12 repetitions were performed per
set per exercise (n = 9). During each session, subjects in both groups performed 3 sets of 7
different exercises representing all major muscles. Training was carried out 3 times per week on
non-consecutive days, for 8 total weeks. Both HL and LL conditions produced significant
increases in thickness of the elbow flexors (5.3 vs. 8.6%, respectively), elbow extensors (6.0 vs.
5.2%, respectively), and quadriceps femoris (9.3 vs. 9.5%, respectively), with no significant
differences noted between groups. Improvements in back squat strength were significantly
greater for HL compared to LL (19.6 vs. 8.8%, respectively) and there was a trend for greater
increases in 1RM bench press (6.5 vs. 2.0%, respectively). Upper body muscle endurance
(assessed by the bench press at 50% 1RM to failure) improved to a greater extent in LL
compared to HL (16.6% vs. -1.2%, respectively). These findings indicate that both HL and LL
training to failure can elicit significant increases in muscle hypertrophy among well-trained
young men; however, HL training is superior for maximizing strength adaptations.
C
C
EP
Keywords: Light weights, strength-endurance continuum, muscle growth, low intensity strength
training, high-intensity strength training
maximize strength is also of particular importance for many athletes, as the capacity to produce
near maximal forces is often required in sport. Given the direct correlation between muscle cross
TE
sectional area (CSA) and force production (29), training-induced hypertrophy is essential for
optimal strength adaptation assuming growth is specific to contractile elements.
Resistance training (RT) is the primary mode of exercise for enhancing muscular
adaptations. Studies show that regimented resistive exercise can promote marked increases in
EP
muscle strength and hypertrophy, with improvements seen irrespective of age and gender (23,
24). Current guidelines state that loads of 65% 1RM are necessary to elicit favorable increases
in hypertrophy, with even higher loads needed to maximize strength (25, 26, 31). It has been
C
C
postulated that heavy loading is required to fully recruit higher threshold motor units (25). Based
on this claim, it stands to reason that optimal improvements in strength and hypertrophy can only
be accomplished through complete motor unit activation via the use of heavy loads.
Some investigators have recently challenged this view that heavy loads are required to
induce muscular adaptations, and claim that recruitment of the full spectrum of motor units is
achievable with low-load training, provided that repetitions are carried out to muscular failure.
(10). Indeed, there is evidence to suggest that highly fatiguing resistive exercise may reduce the
threshold for recruitment of high-threshold motor units (21, 50, 59). It is thus possible that a
greater number of FT motor units are called into play during low-load RT as the point of
muscular failure is reached. However, research has yet to demonstrate whether recruitment at
adaptations in low- versus high-load training programs. Results of these studies are conflicting,
with some studies finding superiority for heavier load training (11, 20, 55) and others showing
TE
no significant differences (28, 32, 37, 44, 57, 58). One fundamental issue with the current state of
the literature is that most studies have reported results from untrained subjects. It is well
established that trained individuals respond differently than those who lack training experience
(42). Early phase RT strength adaptations are predominantly related to improvements in the
EP
ability of the nervous system to efficiently activate and coordinate muscles, whereas the role of
hypertrophy for strength becomes increasingly more relevant as one gains experience (53, 54). In
addition, a "ceiling effect" makes it progressively difficult for trained individuals to increase
C
C
muscular gains over time, thereby necessitating progressive RT protocols to elicit continual
hypertrophic and strength responses (3). Moreover, there is emerging evidence that consistent
training alters the acute response to resistance exercise. Trained muscle differs not only from a
structural (30, 52) and functional (2, 22, 51, 52) perspective but alterations in anabolic
intracellular signaling in rodents (39) and humans (13) along with altered acute protein synthetic
(43, 56, 64), mitochondrial protein synthetic (64) and transcriptional responses (19) may indicate
an attenuated hypertrophic response. As such, current findings from untrained subjects cannot
necessarily be generalized to the response that might be expected among from a well-trained
population. The purpose of this study therefore was to compare the effect of low- versus highload training on muscular adaptations in resistance-trained subjects. We hypothesized that the
Subjects were pair-matched based on initial strength capacity and then randomly assigned
to a group that either performed training at a loading range of 8-12 repetitions or a group that
TE
performed 25-35 repetitions to muscle failure. All other RT variables (e.g., exercises performed,
rest, repetition tempo, etc.) were held constant. The training interventions lasted 8 weeks with
subjects performing 3 total body workouts per week. Testing was carried out pre- and post-study
for muscle strength, muscle endurance, and muscle hypertrophy of the elbow flexors (biceps
EP
brachii and brachialis), elbow extensors (triceps brachii), and quadriceps femoris.
Subjects
Subjects were a convenience sample of 24 male volunteers (age = 23.3 yrs; body mass =
C
C
82.5 kg; height = 175 cm; resistance training experience = 3.4 yrs), recruited from a university
population. Subjects were between the ages of 18-35, did not have any existing musculoskeletal
disorders, were free from consumption of anabolic steroids or any other illegal agents known to
increase muscle size for the previous year, and were experienced lifters (i.e., defined as
consistently lifting weights at least 3 times per week for a minimum of 1 year, and regularly
performing the bench press and squat). The range of lifting experience for all subjects was
between 1.5 and 9 years of consistent training.
TE
groups of the body. The exercises performed were: flat barbell press, barbell military press, wide
grip lat pulldown, seated cable row, barbell back squat, machine leg press, and machine leg
extension. The exercises were chosen based on their common inclusion in bodybuilding- and
strength-type RT programs (5, 12). Subjects were instructed to refrain from performing any
EP
additional resistance-type or high-intensity anaerobic training for the duration of the study.
Training for both routines consisted of three weekly sessions performed on nonconsecutive days for 8 weeks. Sets were carried out to the point of momentary concentric
C
C
muscular failure, i.e., the inability to perform another concentric repetition while maintaining
proper form. Cadence of repetitions was carried out in a controlled fashion, with a concentric
action of approximately one second and an eccentric action of approximately two seconds.
Subjects were afforded 90 seconds rest between sets. The load was adjusted for each exercise as
needed on successive sets, to ensure that subjects achieved failure in the target repetition range.
All routines were directly supervised by the research team, which included a National Strength
and Conditioning Association certified strength and conditioning specialist and certified personal
trainers, to ensure proper performance of the respective routines. Attempts were made to
progressively increase the loads lifted each week within the confines of maintaining the target
repetition range. Prior to training, the LL group underwent 30-repetition maximum (RM) testing
To avoid potential dietary confounding of results, subjects were advised to maintain their
customary nutritional regimen and to avoid taking any supplements other than that provided in
TE
the course of the study. Self-reported food records were collected twice during the study: one
week before the first training session (i.e. baseline) and during the final week of the training
protocol. A 3-day dietary recall booklet was provided to subjects to assess potential differences
in total energy and macronutrient intakes between groups. Subjects were shown how to properly
EP
fill out the booklet, and were instructed to record all food items and their respective portion sizes
consumed for the designated period of interest. The Interactive Healthy Eating Index (Center for
Nutrition Policy and Promotion, United States Department of Agriculture;
C
C
http://www.usda.gov/cnpp) was used to analyze food records. Each item of food was
individually entered into the program, and the program provided relevant information as to total
energy consumption, as well as amount of energy derived from proteins, fats, and carbohydrates
over the three reference days. To facilitate recovery, subjects were provided with a supplement
on training days containing 24g protein and 1g carbohydrate (Iso100 Hydrolyzed Whey Protein
Isolate, Dymatize Nutrition, Farmers Branch, TX). The supplement was consumed within one
hour post-exercise, as this time frame has been purported to help potentiate increases in muscle
protein synthesis following a bout of RT (4).
Measurements
Medical Imaging, Ltd, Jiang Su Province, China). The technician, who was not blinded to group
assignment, applied a water-soluble transmission gel (Aquasonic 100 Ultrasound Transmission
TE
gel, Parker Laboratories Inc., Fairfield, NJ) to each measurement site, and a 5 MHz ultrasound
probe was placed perpendicular to the tissue interface without depressing the skin. When the
quality of the image was deemed to be satisfactory, the technician saved the image to hard drive
and obtained MT dimensions by measuring the distance from the subcutaneous adipose tissue-
EP
muscle interface to the muscle-bone interface as per the protocol by Abe et al. (1). Measurements
were taken on the right side of the body at three sites: (1) elbow flexors (combination of biceps
brachii and brachialis) (2) elbow extensors (triceps brachii) and (3) quadriceps femoris
C
C
(combination of rectus femoris and vastus intermedius). For the anterior and posterior arm,
measurements were taken 60% distal between the lateral epicondyle of the humerus and the
acromion process of the scapula at the midline of the arm (measured from the cubital fossa); for
the quadriceps femoris, measurements were taken 50% between the lateral condyle of the femur
and greater trochanter for the quadriceps femoris at the midline of the thigh (measured from the
distal aspect of the patella). Sites were measured with a vinyl measuring tape and then marked
with a felt pen to ensure precision from session to session. During upper extremity
measurements, participants remained seated with their arms relaxed in an extended position;
measurements for the quadriceps were obtained while standing with legs in a relaxed, extended
position. Ultrasound has been validated as a good predictor of gross muscle hypertrophy in these
thickness return to baseline within 48 hours following a RT session (38). To further ensure
accuracy of measurements, at least 2 images were obtained for each site. If measurements were
TE
within 10% of one another the figures were averaged to obtain a final value. If measurements
were more than 10% of one another, a third image was obtained and the closest of the measures
were then averaged. The test-retest intraclass correlation coefficient (ICC) from our lab for
EP
thickness measurement of the elbow flexors, elbow extensors, and quadriceps femoris, are 0.976,
Muscle Strength: Upper and lower body strength was assessed by 1RM testing in the
bench press (1RMBP) followed by the parallel back squat (1RMBS) exercises. Subjects reported
C
C
to the lab having refrained from any exercise other than activities of daily living for at least 48
hours prior to baseline testing and at least 48 hours prior to testing at the conclusion of the study.
Repetition maximum testing was consistent with recognized guidelines as established by the
National Strength and Conditioning Association (5). In brief, subjects performed a general
warm-up prior to testing that consisted of light cardiovascular exercise lasting approximately 510 minutes. A specific warm-up set of the given exercise of 5 repetitions was performed at ~50%
1RM followed by one to two sets of 2-3 repetitions at a load corresponding to ~60-80% 1RM.
Subjects then performed sets of 1 repetition of increasing weight for 1RM determination. Three
to 5 minutes rest was provided between each successive attempt. All 1RM determinations were
made within 5 attempts. Subjects were required to reach parallel in the 1RMBS for the attempt to
place using free weights. Two fitness professionals supervised all testing sessions and an attempt
was only deemed successful when a consensus was reached between the two. The test-retest ICC
TE
from our lab for the 1RMBP and 1RMBS are 0.91 and 0.87, respectively.
Muscle Endurance: Upper body muscular endurance was assessed by performing bench
press using 50% of 1RM (50%BP) for as many repetitions as possible to muscular failure with
proper form. Successful performance was achieved if the subject displayed a five-point body
EP
contact position (head, upper back and buttocks firmly on the bench with both feet flat on the
floor) and executed a full lock-out. Initial testing used baseline 1RMBP and final testing used the
subjects 1RMBP at the end of the study to determine muscular endurance. The values were
C
C
expressed in terms of volume load to account for differences between absolute strength from
baseline to the studys end. Muscular endurance testing was carried out after assessment of
muscular strength to minimize effects of metabolic stress interfering with performance of the
latter.
Statistical Analyses
Descriptive statistics were used to explore the distribution, central tendency, and
variation of each measurement for both groups, with an emphasis on graphical methods such as
histograms, scatterplots, and boxplots. Descriptive statistics for strength capacity, and sitespecific muscle thickness were reported at baseline, at 8-weeks, and as change from baseline.
Paired t-tests were used to examine differences from baseline to post-intervention, within groups.
values (centered at the mean values) as predictors. This model is equivalent to an analysis of
covariance, but has the advantage of providing estimates associated with each group, adjusted for
TE
baseline characteristics that are potentially associated with the primary outcomes. Coefficient of
the LL group indicator was employed to estimate the mean post-intervention outcome (e.g.
muscle thickness at post-intervention) associated with LL, compared with HL, and the intercept
estimated the mean of post-intervention HL. Regression assumptions were checked. Independent
EP
t-tests were used to evaluate differences between groups at baseline. Values are reported as mean
(SD). Two-tailed alpha was set a priori at 0.05.
Results
C
C
A total of 18 subjects completed the study; 9 subjects in LL and 9 subjects in HL. Six
subjects dropped out prior to completion; 2 because of minor injuries sustained during training
(one in each group) and 4 for personal reasons. Overall attendance was good for those who
completed the study, with a mean participation rate of 93.7% in HL and 95.1% in LL. No
significant differences were noted between groups in any baseline measure. There were no
differences in any dietary measure either within- or between-subjects over the course of the
study. Results of all outcomes are presented in Table 1.
Place Table 1 About Here
Muscle Thickness
10
TE
increased muscle thickness from baseline to post-study by 2.7 2.2 mm (6.0%) and 2.3 3.3
mm (5.2%), respectively (p < 0.05). No significant between-group differences were noted for
absolute or relative change, nor when adjusting for baseline (see Figure 2).
Place Figure 2 About Here
EP
Ultrasound imaging of the quadriceps femoris showed that both the HL and LL groups
increased muscle thickness from baseline to post-study by 5.3 2.2 mm (9.3%) and 5.2 4.8
mm (9.5%), respectively (p < 0.05). No significant between-group differences were noted for
C
C
absolute or relative change, nor when adjusting for baseline (see Figure 3).
Place Figure 3 About Here
Maximal Strength
6.5% (p < 0.01); the LL group showed a non-significant increase of 2.0% (see Figure 4). No
significant between-group differences were noted for absolute or relative change, nor when
adjusting for baseline. However, there was a strong trend for HL producing superior results for
absolute (+4.6 kgs 5.0) and relative (+4.5%) strength.
Place Figure 4 About Here
11
TE
The LL group showed a significant increase in muscular endurance of 16.6% (p < 0.05);
whereas no differences were noted from baseline in the HL group (see Figure 6). Significant
differences were noted between groups, such that LL training produced superior results for
absolute (+230.6 52.5 kgs) and relative (+17.8%) gains. After adjustment for baseline values,
EP
the difference remained significant, such that LL produced superior results compared to HL
C
C
Discussion
To the authors knowledge, this is the first study to evaluate muscular adaptations in lowversus high-load training in well-trained individuals. The study produced several important
findings. With respect to gross measures of muscle hypertrophy, LL significantly increased
muscle thickness of the upper and lower extremities. In comparison to a traditional hypertrophy
protocol of 8-12 repetitions per set, the LL condition produced similar gains in thickness of the
elbow flexors (5.3 vs. 8.6%, respectively), elbow extensors (6.0 vs. 5.2%) and quadriceps
femoris (9.3 vs. 9.5%). These results run contrary to generally accepted hypertrophy training
guidelines, which profess that loads of at least 65% are necessary to stimulate muscle growth in
well-trained individuals (25, 26, 31).
12
The majority of prior studies standardized the number of sets such that the greater number of
repetitions performed during low-load training resulted in a higher total amount of work for this
TE
condition. With the exception of Schuenke et al. (55), all studies that did not equate volume
reported similar increases in muscle growth between high- and low-load training (28, 32, 37, 44,
57, 58). Conversely, the two studies that did equate total intra-session work showed a
hypertrophic advantage for high-load exercise (11, 20). The LL group in our study performed
EP
approximately three times the total volume (sets x repetitions) compared to the HL group. Given
that compelling evidence exists for a dose-response relationship between hypertrophy and RT
volume at least up to a certain threshold (27), it can be hypothesized that the differences in total
C
C
Another intriguing possibility is that fiber-type specific responses may have played a
role in mediating muscle protein accretion. It is generally accepted that type II fibers display an
approximately 50% greater capacity for growth compared to type I fibers. However, the superior
hypertrophic capacity type II fibers may be more a consequence of the models in which they
have been studied as opposed to an intrinsic property of the fiber itself (40). Specifically,
research to date has been biased towards RT intensities >60% 1RM, with a paucity of studies
investigating lower intensities. Given the fatigue-resistant nature of type I fibers, it seems logical
to conclude that the increased time-under-load associated with low-load training is necessary to
fully stimulate these fibers. This hypothesis is supported by Netreba et al (35), who found that
13
It should be noted that none of the subjects in our study reported training with more than 15
repetitions per set as part of their normal resistance-training programs. Thus, it is possible that
TE
the type I fibers of subjects were underdeveloped in comparison to the type II fibers as a result of
training methodologies. The type I fibers therefore may have had a greater potential for growth
compared to the type II fibers, and the extended duration of the LL sets conceivably provided a
EP
novel stimulus to promote greater growth in the endurance-oriented type I fibers. This hypothesis
It is also plausible that differing inter-muscle activation strategies in multi-joint or multimuscle movements could account for the comparable hypertrophy under certain circumstances.
C
C
For the squat exercise specifically, it has been shown that increased training intensity (%-1RM)
is associated with greater contribution from joints other than the knee (hip, ankle), whereas the
knee, and therefore quadriceps, have a constant relative mechanical effort in response to
increased training loads (7, 17). Given these findings, it is conceivable that varying training
intensities in the squat may represent equivalent stress to the quadriceps muscle group, thereby
providing a comparable training stimulus for muscle growth. Conversely, others have
demonstrated that EMG activity is greater in the quadriceps with higher than lower training loads
during single joint exercises (14). Consequently it is not possible to reconcile these findings, as
differences in experimental methodology confound the issue. Further research is required to
clarify the role of inter-muscle differences in activation with single- and multi-joint exercises
14
increases in both 1RMBP (6.5 vs. 2.0%, respectively) and 1RMBS (19.6 vs. 8.8%, respectively).
The observation of increased improvement in strength with HL despite equivalent hypertrophy is
TE
consistent with other comparisons of high- versus low-load training (32, 46). Multiple metaanalyses have identified that peak gains in strength occur with training above 60%-1RM in both
trained and untrained individuals, although the optimal intensity is higher in the trained (41, 42,
49). The disparate strength adaptations despite equivalent hypertrophic changes between HL and
EP
LL are not unexpected given that muscle hypertrophy accounts for approximately 19% of the
change in muscle strength with chronic resistance exercise in untrained individuals (15). Even in
the untrained state, muscle size is estimated to explain at most 50% of the variability in
C
C
maximum muscle strength (6), suggesting that other mechanisms contribute to alterations in
strength with training. Neural adaptations can contribute to increased strength with RT, including
but not limited to increased muscle activation, increased motor unit firing rates, increased
frequency of doublet firing, enhanced motor unit synchronization, and/or alterations in agonist-
antagonist co-activation ratios (18), however, the contribution of these mechanisms to the present
dataset is not known. Regardless of potential mechanisms, it can be inferred that muscle strength
is increased with both low- and high-load training but high-load training is superior for maximal
strength development.
Adaptations in muscular endurance favored the LL condition, with a mean volume load
increase in 50%BP to failure of 16.6% compared to a small non-significant decrease of -1.2% for
15
consequence of strength training regardless of loading intensity. In addition, the longer timeunder-load that occurs with high-repetition, low-load training (8) differentially and favorably
TE
affects mitochondrial protein synthesis that may enhance cellular energetics, culminating in
improved fatigue resistance (9). It should be noted that the greater increase in bench press
strength for the HL group resulted in their lifting slightly higher mean loads (~2 kg) compared to
LL. However, given the large magnitude of difference in this outcome favoring LL, the
EP
additional load lifted by HL likely had minimal effect on measures of muscle endurance.
The present study had several limitations that must be considered when extrapolating
conclusions based on the results. First, the study period lasted only 8 weeks. While this duration
C
C
was sufficient to produce significant increases in muscular strength and hypertrophy, it is not
clear whether results between groups would have diverged over a longer term.
Second, muscle thickness was measured only at the middle portion of the muscle.
Although this region is generally considered to be indicative of overall growth of a given muscle,
there is evidence that hypertrophy manifests in a regional-specific manner, with greater gains
sometimes seen at the proximal and/or distal aspects (60, 61). This may be related to exercisespecific intramuscular activation and/or tissue oxygenation saturation (33, 60, 61). Thus, we
cannot rule out the possibility that greater changes in proximal or distal muscle thickness
occurred in one protocol versus the other.
16
dietary analysis revealed no differences in total daily protein intake between groups. Moreover,
there is no research suggesting that the provision of protein differentially affects a given loading
TE
EP
of muscle, recuperative abilities, and other factors could alter muscular adaptations to lowand/or high-load protocols in these individuals.
Practical Applications
C
C
In conclusion, our results provide compelling evidence that low-load training can be an
effective method to increase muscle hypertrophy of the extremities in well-trained men. The
gains in muscle size from low-load training were equal to that achieved with training in a
repetition range normally recommended for maximizing muscle hypertrophy. Provided that
maximal hypertrophy is the primary outcome goal irrespective of strength increases, these
findings suggest that a new paradigm should be considered for hypertrophy training
recommendations, with low-load training promoted as a viable option. On the other hand, if
maximizing strength gains is of primary importance, then heavier loading should be employed at
the exclusion of lower load training. Given the preservation of the strength-endurance continuum
regarding muscle strength and endurance adaptations alongside equivalent muscle hypertrophy,
17
phenotype. It therefore can be hypothesized that combining low- and high-load sets would be
optimal for maximizing muscle growth. These findings suggest a potential benefit to
TE
EP
Acknowledgements: This study was supported by a grant from Dymatize Nutrition. The
authors gratefully acknowledge the contributions of Robert Harris, Andre Mitchell, Ramon
Belliard, Azuka Utti, Francis Ansah, Romaine Fearon, and James Jackson in their indispensable
role as research assistants in this study.
References
1. Abe, T, DeHoyos, DV, Pollock, ML, and Garzarella, L. Time course for strength and muscle
thickness changes following upper and lower body resistance training in men and women. Eur. J.
Appl. Physiol. 81: 174-180, 2000.
C
C
2. Alway, SE, MacDougall, JD, Sale, DG, Sutton, JR, and McComas, AJ. Functional and
structural adaptations in skeletal muscle of trained athletes. J. Appl. Physiol. (1985) 64: 11141120, 1988.
3. American College of Sports Medicine. American College of Sports Medicine position stand.
Progression models in resistance training for healthy adults. Med. Sci. Sports Exerc. 41: 687-708,
2009.
4. Aragon, AA, and Schoenfeld, BJ. Nutrient timing revisited: is there a post-exercise anabolic
window? J. Int. Soc. Sports Nutr. 10: 5-2783-10-5, 2013.
5. Baechle, TR, Earle, RW. Essentials of strength training and conditioning. In: Anonymous
Champaign, IL: Human Kinetics, 2008.
6. Bamman, MM, Newcomer, BR, Larson-Meyer, DE, Weinsier, RL, and Hunter, GR.
Evaluation of the strength-size relationship in vivo using various muscle size indices. Med. Sci.
Sports Exerc. 32: 1307-1313, 2000.
18
9. Burd, NA, Andrews, RJ, West, DW, Little, JP, Cochran, AJ, Hector, AJ, Cashaback, JG,
Gibala, MJ, Potvin, JR, Baker, SK, and Phillips, SM. Muscle time under tension during
resistance exercise stimulates differential muscle protein sub-fractional synthetic responses in
men. J. Physiol. 590: 351-362, 2012.
TE
10. Burd, NA, Mitchell, CJ, Churchward-Venne, TA, and Phillips, SM. Bigger weights may not
beget bigger muscles: evidence from acute muscle protein synthetic responses after resistance
exercise. Appl. Physiol. Nutr. Metab. 37: 551-554, 2012.
EP
11. Campos, GER, Luecke, TJ, Wendeln, HK, Toma, K, Hagerman, FC, Murray, TF, Ragg, KE,
Ratamess, NA, Kraemer, WJ, and Staron, RS. Muscular adaptations in response to three different
resistance-training regimens: specificity of repetition maximum training zones. Eur. J. Appl.
Physiol. 88: 50-60, 2002.
12. Coburn, JW, Malek, MH. NSCA's essentials of personal training. In: Anonymous
Champaign, IL: Human Kinetics, 2011.
C
C
13. Coffey, VG, Zhong, Z, Shield, A, Canny, BJ, Chibalin, AV, Zierath, JR, and Hawley, JA.
Early signaling responses to divergent exercise stimuli in skeletal muscle from well-trained
humans. FASEB J. 20: 190-192, 2006.
14. Cook, SB, Murphy, BG, and Labarbera, KE. Neuromuscular function after a bout of lowload blood flow-restricted exercise. Med. Sci. Sports Exerc. 45: 67-74, 2013.
15. Erskine, RM, Fletcher, G, and Folland, JP. The contribution of muscle hypertrophy to
strength changes following resistance training. Eur. J. Appl. Physiol. 114: 1239-1249, 2014.
16. Fitzgerald, SJ, Barlow, CE, Kampert, JB, Morrow, JR, Jackson, AW, and Blair, SN.
Muscular Fitness and All-Cause Mortality: Prospective Observations. Journal of Physical
Activity and Health 1: 7-18, 2004.
17. Flanagan, SP, and Salem, GJ. Lower extremity joint kinetic responses to external resistance
variations. J. Appl. Biomech. 24: 58-68, 2008.
18. Gabriel, DA, Kamen, G, and Frost, G. Neural adaptations to resistive exercise: mechanisms
and recommendations for training practices. Sports Med. 36: 133-149, 2006.
19
21. Houtman, CJ, Stegeman, DF, Van Dijk, JP, and Zwarts, MJ. Changes in muscle fiber
conduction velocity indicate recruitment of distinct motor unit populations. J. Appl. Physiol. 95:
1045-1054, 2003.
TE
22. Huczel, HA, and Clarke, DH. A comparison of strength and muscle endurance in strengthtrained and untrained women. Eur. J. Appl. Physiol. Occup. Physiol. 64: 467-470, 1992.
EP
23. Ivey, FM, Roth, SM, Ferrell, RE, Tracy, BL, Lemmer, JT, Hurlbut, DE, Martel, GF, Siegel,
EL, Fozard, JL, Jeffrey Metter, E, Fleg, JL, and Hurley, BF. Effects of age, gender, and
myostatin genotype on the hypertrophic response to heavy resistance strength training. J.
Gerontol. A Biol. Sci. Med. Sci. 55: M641-8, 2000.
24. Kosek, DJ, Kim, JS, Petrella, JK, Cross, JM, and Bamman, MM. Efficacy of 3 days/wk
resistance training on myofiber hypertrophy and myogenic mechanisms in young vs. older
adults. J. Appl. Physiol. 101: 531-544, 2006.
C
C
25. Kraemer, WJ, and Ratamess, NA. Fundamentals of resistance training: progression and
exercise prescription. Med. Sci. Sports Exerc. 36: 674-688, 2004.
26. Kraemer, WJ, Adams, K, Cafarelli, E, Dudley, GA, Dooly, C, Feigenbaum, MS, Fleck, SJ,
Franklin, B, Fry, AC, Hoffman, JR, Newton, RU, Potteiger, J, Stone, MH, Ratamess, NA, and
Triplett-McBride, T. American College of Sports Medicine position stand. Progression models in
resistance training for healthy adults. Med. Sci. Sports Exerc. 34: 364-380, 2002.
27. Krieger, JW. Single vs. multiple sets of resistance exercise for muscle hypertrophy: a metaanalysis. J. Strength Cond Res. 24: 1150-1159, 2010.
28. Leger, B, Cartoni, R, Praz, M, Lamon, S, Deriaz, O, Crettenand, A, Gobelet, C, Rohmer, P,
Konzelmann, M, Luthi, F, and Russell, AP. Akt signalling through GSK-3beta, mTOR and
Foxo1 is involved in human skeletal muscle hypertrophy and atrophy. J. Physiol. 576: 923-933,
2006.
29. Maughan, RJ, Watson, JS, and Weir, J. Strength and cross-sectional area of human skeletal
muscle. J. Physiol. 338: 37-49, 1983.
30. Maughan, RJ, Watson, JS, and Weir, J. Muscle strength and cross-sectional area in man: a
comparison of strength-trained and untrained subjects. Br. J. Sports Med. 18: 149-157, 1984.
20
TE
34. Miyatani, M, Kanehisa, H, Ito, M, Kawakami, Y, and Fukunaga, T. The accuracy of volume
estimates using ultrasound muscle thickness measurements in different muscle groups. Eur. J.
Appl. Physiol. 91: 264-272, 2004.
35. Netreba, AI, Popov, DV, Liubaeva, EV, Bravyi, I, Prostova, AB, Lemesheva, I, and
Vinogradova, OL. Physiological effects of using the low intensity strength training without
relaxation in single-joint and multi-joint movements. Ross. Fiziol. Zh. Im. I. M. Sechenova. 93:
27-38, 2007.
EP
36. Nogueira, W, Gentil, P, Mello, SN, Oliveira, RJ, Bezerra, AJ, and Bottaro, M. Effects of
power training on muscle thickness of older men. Int. J. Sports Med. 30: 200-204, 2009.
37. Ogasawara, R, Loenneke, JP, Thiebaud, RS, and Abe, T. Low-Load Bench Press Training to
Fatigue Results in Muscle Hypertrophy Similar to High-Load Bench Press Training. Int J Clin
Med. 4: 114-121, 2013.
C
C
38. Ogasawara, R, Thiebaud, RS, Loenneke, JP, Loftin, M, and Abe, T. Time course for arm and
chest muscle thickness changes following bench press training. Interventional Medicine and
Applied Science 4: 217-220, 2012.
39. Ogasawara, R, Kobayashi, K, Tsutaki, A, Lee, K, Abe, T, Fujita, S, Nakazato, K, and Ishii,
N. mTOR signaling response to resistance exercise is altered by chronic resistance training and
detraining in skeletal muscle. J. Appl. Physiol. , 2013.
40. Ogborn, D, and Schoenfeld, BJ. The role of fber types in muscle hypertrophy: Implications
for loading strategies. Strength Cond J 36: 20-25, 2014.
41. Peterson, MD, Rhea, MR, and Alvar, BA. Maximizing strength development in athletes: a
meta-analysis to determine the dose-response relationship. J. Strength Cond Res. 18: 377-382,
2004.
42. Peterson, MD, Rhea, MR, and Alvar, BA. Applications of the dose-response for muscular
strength development: a review of meta-analytic efficacy and reliability for designing training
prescription. J. Strength Cond Res. 19: 950-958, 2005.
21
45. Pretorius, A, and Keating, JL. Validity of real time ultrasound for measuring skeletal muscle
size. Phys Ther Rev 13: 415-426, 2008.
TE
46. Rana, SR, Chleboun, GS, Gilders, RM, Hagerman, FC, Herman, JR, Hikida, RS, Kushnick,
MR, Staron, RS, and Toma, K. Comparison of early phase adaptations for traditional strength
and endurance, and low velocity resistance training programs in college-aged women. J. Strength
Cond Res. 22: 119-127, 2008.
47. Rantanen, T, Avlund, K, Suominen, H, Schroll, M, Frandin, K, and Pertti, E. Muscle strength
as a predictor of onset of ADL dependence in people aged 75 years. Aging Clin. Exp. Res. 14:
10-15, 2002.
EP
48. Reeves, ND, Maganaris, CN, and Narici, MV. Ultrasonographic assessment of human
skeletal muscle size. Eur. J. Appl. Physiol. 91: 116-118, 2004.
49. Rhea, MR, Alvar, BA, Burkett, LN, and Ball, SD. A meta-analysis to determine the dose
response for strength development. Med. Sci. Sports Exerc. 35: 456-464, 2003.
C
C
52. Sale, DG, MacDougall, JD, Alway, SE, and Sutton, JR. Voluntary strength and muscle
characteristics in untrained men and women and male bodybuilders. J. Appl. Physiol. (1985) 62:
1786-1793, 1987.
53. Sale, DG. Neural adaptation to resistance training. Med. Sci. Sports Exerc. 20: S135-45,
1988.
54. Schoenfeld, BJ. The mechanisms of muscle hypertrophy and their application to resistance
training. J. Strength Cond Res. 24: 2857-2872, 2010.
55. Schuenke, MD, Herman, JR, Gliders, RM, Hagerman, FC, Hikida, RS, Rana, SR, Ragg, KE,
and Staron, RS. Early-phase muscular adaptations in response to slow-speed versus traditional
resistance-training regimens. Eur. J. Appl. Physiol. 112: 3585-3595, 2012.
22
TE
59. Vollestad, NK, Vaage, O, and Hermansen, L. Muscle glycogen depletion patterns in type I
and subgroups of type II fibres during prolonged severe exercise in man. Acta Physiol. Scand.
122: 433-441, 1984.
EP
C
C
62. Weiss, LW, Coney, HD, and Clark, FC. Differential functional adaptations to short-term low, moderate- and high-repetition weight training. J Strength Cond Res 13: 236-241, 1999.
63. Weiss, LW, Coney, HD, and Clark, FC. Gross measures of exercise-induced muscular
hypertrophy. J. Orthop. Sports Phys. Ther. 30: 143-148, 2000.
64. Wilkinson, SB, Phillips, SM, Atherton, PJ, Patel, R, Yarasheski, KE, Tarnopolsky, MA, and
Rennie, MJ. Differential effects of resistance and endurance exercise in the fed state on
signalling molecule phosphorylation and protein synthesis in human muscle. J. Physiol. 586:
3701-3717, 2008.
65. Young, A, Stokes, M, Round, JM, and Edwards, RH. The effect of high-resistance training
on the strength and cross-sectional area of the human quadriceps. Eur. J. Clin. Invest. 13: 411417, 1983.
23
LL-PRE
LL-POST
ES
HL-PRE
42.4 6.6
44.5 6.8
54.6 10.9
101.0 25.6
122.1 39.7
1282.8 220.8
46.0 7.1*
46.9 7.4*
59.8 9.2*
103.0 23.3
132.8 36.5*
1496.0 104.9*
0.54
0.33
0.51
0.08
0.28
1.23
46.6 6.3
45.6 5.4
57.1 4.2
101.5 20.5
121.0 36.6
1438.4 311.7
HL-POST
49.1 6.2*
48.3 3.9*
62.3 5.2*
108.1 21.0*
144.7 27.4*
1421.0 257.0
C
C
EP
TE
ES
0.40
0.57
1.10
0.32
0.73
-0.06
TE
D
80
70
Pre
Post
EP
50
40
30
10
C
C
20
Low Load
High Load
Condition
60
80
Pre
Post
TE
D
60
50
40
30
EP
20
10
Low Load
High Load
C
C
Condition
70
TE
D
100
90
Pre
Post
70
EP
60
50
40
20
10
0
C
C
30
Low Load
High Load
Condition
80
160
TE
D
140
Pre
Post
100
80
EP
60
40
20
C
C
Low Load
High Load
Condition
120
TE
D
220
200
Pre
Post
180
EP
140
120
100
80
40
20
0
C
C
60
Low Load
High Load
Condition
160
2200
2000
Pre
Post
TE
D
1800
1400
1200
1000
800
600
EP
400
200
0
Low Load
High Load
Condition
A
C
1600
Figure Captions
Figure 1 Graphical representation of muscle thickness values of the biceps brachii pre- and postintervention for LL and HL, mean (SD). Values expressed in mms. *Significantly greater than
corresponding pre-training value.
Figure 2 Graphical representation of muscle thickness values of the triceps brachii pre- and postintervention for LL and HL, mean (SD). Values expressed in mms. *Significantly greater than
corresponding pre-training value.
Figure 3 Graphical representation of muscle thickness values of the quadriceps femoris pre- and
post-intervention for LL and HL, mean (SD). Values expressed in mms. *Significantly greater
than corresponding pre-training value.
TE
Figure 4 Graphical representation of 1RM bench press values pre- and post-intervention for LL
and HL, mean (SD). Values expressed in kgs. *Significantly greater than corresponding pretraining value.
Figure 5 Graphical representation of 1RM back squat values pre- and post-intervention for LL
and HL, mean (SD). Values expressed in kgs. *Significantly greater than corresponding pretraining value. #Significantly greater than corresponding group.
C
C
corresponding group.
EP
Figure 6 Graphical representation of 50% bench press to falure values pre- and post-intervention
for LL and HL, mean (SD). Values expressed as total volume-load (total number of repetitions
x load) in kgs. *Significantly greater than corresponding pre-training value. #Significantly greater than