|

Received: 29 December 2018    Accepted: 22 January 2020

DOI: 10.1111/cpf.12622

REVIEW ARTICLE

Does stretch training induce muscle hypertrophy in humans?

A review of the literature

João Pedro Nunes1  | Brad J. Schoenfeld2 | Masatoshi Nakamura3  |

Alex S. Ribeiro1,4 | Paolo M. Cunha1 | Edilson S. Cyrino1

1
Metabolism, Nutrition, and Exercise
Laboratory, Physical Education and Sport Abstract
Center, Londrina State University, Londrina, Stretch training is widely used in a variety of fitness-related capacities such as in-
Brazil
2 creasing joint range of motion, preventing contractures and alleviating injuries.
Department of Health Sciences, Lehman
College, Bronx, NY, USA Moreover, some researches indicate that stretch training may induce muscle hyper-
3
Institute for Human Movement and Medical trophy; however, studies on the topic have been primarily relegated to animal and in
Sciences, Niigata University of Health and
Welfare, Niigata, Japan
vitro models. The purpose of this brief review was to evaluate whether stretch train-
4
Center for Research in Health Sciences, ing is a viable strategy to induce muscle hypertrophy in humans. An extensive litera-
University of Northern Paraná, Londrina, ture search was performed using PubMed/MEDLINE, SciELO and Scopus databases,
Brazil
using terms related to stretching and muscle hypertrophy. Only human trials that
Correspondence evaluated changes in measures of muscle size or architecture following training pro-
João Pedro Nunes, Metabolism, Nutrition,
and Exercise Laboratory, Physical Education tocols that it was performed stretching exercises were selected for inclusion. Of the
and Sport Center, Londrina State University, 10 studies identified, 3 observed some significantly positive effects of stretch train-
Londrina, Brazil.
Email: [email protected] ing on muscle structure. Intriguingly, in these studies, the stretching was carried out
with an apparatus that aided in its performance, or with an external overload. In all
studies, the subjects performed stretching at their own self-determined range of mo-
tion, and no effect was observed. Of the 5 available studies that integrated stretching
into a resistance training programme, 2 applied the stretching in the interset rest pe-
riod and were the ones that showed enhanced muscle growth. In conclusion, passive,
low-intensity stretch does not appear to confer beneficial changes in muscle size and
architecture; alternatively, albeit limited evidence suggests that when stretching is
done with a certain degree of tensile strain (particularly when loaded, or added be-
tween active muscle contractions) may elicit muscle hypertrophy.

KEYWORDS

fascicle length, flexibility, muscle thickness, pennation angle, protein synthesis, resistance
training, stretching

1 | I NTRO D U C TI O N tolerance (Magnusson, Simonsen, Aagaard, Sørensen, & Kjaer,

1996; Medeiros & Lima, 2017). Stretching may also reduce the
Stretch training is widely used to prevent contractures, facil- stiffness of the muscle–tendon unit (Riley & Van Dyke, 2012), al-
itate recovery from injuries and muscle–tendon shortening though this effect appears to last about half an hour (Magnusson
and is effective for improving joint range of motion and stretch & Renström, 2006). Also, despite stretching exercises may acutely

© 2020 Scandinavian Society of Clinical Physiology and Nuclear Medicine. Published by John Wiley & Sons Ltd

|
148     
wileyonlinelibrary.com/journal/cpf Clin Physiol Funct Imaging. 2020;40:148–156.
NUNES et al. |
      149

impair the subsequent muscle performance (Magnusson & with ultrasound, magnetic resonance imaging or computed tomog-
Renström, 2006), some studies indicate that carrying out stretch- raphy. Those involving proprioceptive neuromuscular facilitation
ing in a chronic fashion may improve the efficiency of several were excluded from consideration. The citations on Google Scholar
exercise tasks (Medeiros & Lima, 2017). Evidence suggests that and the reference lists of the selected articles were subsequently
a loaded stretching provides mechanical and metabolic stimuli to screened by the lead author (JPN) to uncover any additional articles
muscle and produces cellular biomarkers that are important for that met inclusion criteria.
muscle growth (Goldberg, Etlinger, Goldspink, & Jablecki, 1975;
Tatsumi, 2010; Wisdom, Delp, & Kuhl, 2015) so that maintaining
a muscle in a lengthened position may help to preserve muscle 3 | R E S U LT S
mass in clinical atrophic conditions (Lowe & Alway, 2002). In this
way, it is speculated that muscle hypertrophy could be enhanced A total of 10 studies (Akagi & Takahashi, 2014; Blazevich et al.,
when performing stretching between exercise sets in a resistance 2014; Freitas & Mil-Homens, 2015; Konrad & Tilp, 2014a, 2014b;
training programme (Mohamad, Nosaka, & Cronin, 2011). Lima, Carneiro, Alves, Peixinho, & Oliveira, 2015; Mizuno, 2019;
The conceptual basis for stretching-mediated hypertro- Moltubakk, 2019; Nakamura, Ikezoe, Takeno, & Ichihashi, 2012;
phic effects dates back to studies in animal and in vitro models Simpson, Kim, Bourcet, Jones, & Jakobi, 2017) met inclusion crite-
(Goldberg et al., 1975; Tatsumi, 2010). Research shows that acute ria. A summary of the results of the included studies is presented
stretching may trigger mechanisms that are important for muscle in Table 1, and Table 2 summarizes the studies' designs. As all
hypertrophy, such as insulin-like and myogenic growth factors, studies used B-mode ultrasound to assess the changes in skeletal
stretch-activated channels, the AKT/mTOR pathway and protein muscle tissue, Table 2 presents which measurement techniques
synthesis (Mohamad et al., 2011; Riley & Van Dyke, 2012; Tatsumi, were adopted in each individual study. Results for the effects of
2010; Wisdom et al., 2015). Indeed, seminal longitudinal studies stretch training are presented as subsections, MT and muscle ar-
in animals showed robust hypertrophy and perhaps hyperplasia chitecture (FL and PA). Studies that included both measures were
after several weeks of intervention (Antonio & Gonyea, 1993b; discussed in each subsection; however, the methodology was
Goldberg et al., 1975; Goldspink, Tabary, Tabary, Tardieu, & included only in the first subsection and not repeated when dis-
Tardieu, 1974). For instance, Goldspink et al. (1974) observed pro- cussed subsequently.
nounced sarcomerogenesis in soleus muscle of cats after 4 weeks
of hind limb denervation and plaster-cast immobilization with
muscles in the stretched position. Similarly, Antonio and Gonyea 3.1 | Effects of stretch training on muscle thickness
(1993a) observed huge muscle growth in quails following a period
of progressive stretch overload of wing muscles, in which it was Lima et al. (2015) investigated the effects of stretch training on bi-
added a cuff weight filled with lead pellets secured around the ceps femoris architecture and MT of the vastus lateralis in 24 healthy,
wing of each bird. physically active men (stretching group, n = 12; control group, n = 12).
Nonetheless, given the nature of these protocols in animal mod- Subjects in the stretching group performed 3 × 30 s of static stretch-
els, the results should not necessarily be extrapolated to humans ing for knee extensors and flexors muscles three times a week for
performing traditional passive stretching. A number of subsequent 8 weeks. The stretching exercise for knee extensors consisted of
studies have endeavoured to investigate the effect of stretch train- trunk extension with the knee flexed while seated on the floor, while
ing on human muscles. Therefore, the purpose of this paper is to stretching for the knee flexors consisted of maximal trunk flexion,
review the evidence as to whether stretch training is capable of elic- knee extension and ankle dorsiflexion in the sitting position. Results
iting muscle hypertrophy in humans. indicated no significant change in vastus lateralis MT (stretching:
pre = 28 mm, post = 26 mm; control: pre = 25 mm, post = 24 mm)
or biceps femoris MT (stretching: pre = 25 mm, post = 26; control:
2 |  M E TH O DS pre = 23 mm, post = 23 mm).
To explore potential time-dependent effects over longer dura-
An extensive literature search was performed using PubMed/ tions, Moltubakk (2019) investigated the effects of 24 weeks of tri-
MEDLINE, SciELO and Scopus databases for all dates up to and in- ceps surae stretching. Stretching was self-performed daily (4 × 60 s
cluding September 2019. Searches were performed using the follow- per day) with subjects instructed to stretch the leg as far posteriorly
ing terms, both in English and in Portuguese, alone or in combination: as possible while pushing the heel down to the ground and point-
“stretch,” “stretching,” “flexibility,” “muscle hypertrophy.” “muscle ing the forefoot forward. At the study's end, no significant effect
growth,” “muscle volume,” “muscle thickness,” “fascicle length” and was observed for the trained limb compared with the contralateral
“muscle architecture.” As inclusion criteria, studies would be se- control limb on medial gastrocnemius MT (stretching: pre = 20 mm,
lected if were performed in healthy humans and directly measured post = 21 mm; control: pre = 20 mm, post = 22 mm), nor on the soleus
muscle thickness (MT), muscle architecture (fascicle length—FL or MT (stretching: pre = 18 mm, post = 19 mm; control: pre = 18 mm,
pennation angle—PA), muscle cross-sectional area or muscle volume, post = 18 mm). On the other hand, in an attempt to ascertain the
150      | NUNES et al.

TA B L E 1   Effects of stretch training on

Muscle structure adaptations
muscle hypertrophy
Pennation Fascicle Muscle
Studies Stretch training type angle length thickness

Konrad and Tilp (2014a) Dynamic, self-performed ↔ ↔ n/a

Konrad and Tilp (2014b) Static, self-performed ↔ ↔ n/a
Nakamura et al. (2012) Static, self-performed n/a ↔ n/a
Blazevich et al. (2014) Static, self-performed n/a ↔ n/a
Akagi and Takahashi (2014) Static, stretching board n/a n/a ↔
Lima et al. (2015) Static, self-performed ↔ ↔ ↔
Freitas and Mil-Homens (2015) Static, machine-assisted ↔ ↑ ↔
Simpson et al. (2017)a  Static, machine-loaded ↓↑ ↑ ↔
Moltubakk (2019) Static, self-performed ↔ ↔ ↔
Mizuno (2019) Static, stretching board ↔ n/a ↑

Note: Summary of the findings of the studies met the inclusion criteria. ↑ = the outcome variable
was improved with the stretch training. ↓ = the outcome variable was decreased with the
stretching training. ↔ = the outcome variable presented similar responses between the stretch
training and the control conditions. n/a = not applicable, unavailable data.
a
In the study of Simpson et al. (2017), values of pennation angle of the triceps surae altered
depending on the location of measurement; please see the results section for further details.

TA B L E 2   Summary of the methodology of the studies met the inclusion criteria

Training duration Measurement protocols of the outcomesd 

Duration TTUS Probe type and FL

Study Sample (n; mean age) (weeks) (min) Muscles analysed width Field of view extrapolation?

Konrad and Tilp (2014a) 48 (30 men); 23 years 6 60 Gastrocnemius medialis Linear array, 10 cm 74 mm depth No
Konrad and Tilp (2014b) 49 (35 men); 23 years 6 60 Gastrocnemius medialis Linear array, 10 cm 74 mm depth No
Nakamura et al. (2012) 18 men; 21 years 4 56 Gastrocnemius medialis Linear array, 5 cm 30 mm depth Yesa
Blazevich et al. (2014) 24 men; 19 years 3 84 Gastrocnemius medialis Linear array, 4.5 cm 40 mm depth No
Akagi and Takahashi (2014) 19 men; 24 years 5 180 Plantar flexors Convex array n/a n/a
Lima et al. (2015) 24 men; 19 years 8 36 Biceps femoris and vastus Linear array, 8 cm 50–80 mm Yesb
lateralis depth
Freitas and Mil-Homens 10 men; 21 years 8 210 Biceps femoris Linear array, 6 cm n/a Yesc
(2015)
Simpson et al. (2017) 21 men; 22 years 6 90 Gastrocnemii medialis and Linear array, 5.8 cm 50 mm depth No
lateralis
Moltubakk (2019) 26 (9 men); 22 years 24 672 Gastrocnemii medialis and Linear array, 5 cm 30–70 mm No
soleus depth
Mizuno (2019) 20 (12 men); 18 years 8 48 Gastrocnemius medialis Linear array, 4.5 cm 39 mm depth n/a

Abbreviations: FL, fascicle length; n/a, not applicable; TTUS, total time under stretching.
a
Calculation of the FL was based on the formula of Kumagai et al. (2000).
b
Calculation of the FL was based on the formula of Potier et al. (2009).
c
Calculation of the FL was based on the formula of Noorkoiv et al. (2010).
d
All studies used B-mode ultrasound to assess the changes in skeletal muscle architecture.

effects of a higher intensity stretching protocol on muscular adap- passive ROM and intensity for 450 s. After 8 weeks, ultrasound mea-
tations, Freitas and Mil-Homens (2015) allocated 10 young, phys- sures of the biceps femoris showed no significant effect of stretch
ically active adults into stretching or control groups, whereby the training on MT.
stretch training targeted the knee flexors with an average frequency Akagi and Takahashi (2014) investigated the effect of stretch
of 3.1 times per week. The stretching protocol required that subjects training on gastrocnemii muscle architecture. Employing a with-
lie on the floor with the hips flexed at 90° and engage a stretch of the in-subject design, 19 young men performed stretch training on one
hamstrings by employing a knee extension at the highest tolerable leg, and the other leg was assigned to serve as a non-training control
NUNES et al. |
      151

condition. Six of the subjects were sedentary, and the others re- post-stretch-training changes were observed for both stretching
ported engaging in a moderate amount of weekly recreational sport- (n = 20) and control (n = 18) groups in FL (stretching: pre = 62 mm,
ing activities. The protocol consisted of static stretching (3 × 120 s) post = 62 mm; control: pre = 61 mm, post = 62 mm) and PA
performed 6 times a week using a calf-stretching board for 5 weeks. (stretching: pre = 19°, post = 19°; control: pre = 18°, post = 18°).
Subjects were instructed to stand erect with one foot on the stretch- The same laboratory investigated a protocol involving against-the-
ing board. Pre-to-post analysis indicated no effect (p = .66) of stretch wall ballistic calf stretch training (4 × 330 s moving up and down
training on MT (stretching = pre: 76 mm, post: 76 mm; control = pre: with the front knee once a second, 5 sessions/week) in a cohort of
76 mm, post: 76 mm). In another study that employed a stretching 48 police cadets (Konrad & Tilp, 2014a); no statistical changes in
board, Mizuno (2019) investigated the effects of an 8-week protocol FL (stretching: pre = 64 mm, post = 63 mm; control: pre = 61 mm,
on the medial gastrocnemius in 20 university students (stretching post = 62 mm) and PA (stretching: pre = 17°, post = 18°; control:
group, n = 11; control group, n = 9). The supervised stretch training pre = 18°, post = 18°) were observed with stretch training com-
sessions were carried out 3 sessions/week and consisted of 4 × 30 s pared with a non-training control after the 6 weeks of interven-
of stretching with 30 s rest between sets. The intensity of stretching tion. Moltubakk (2019) also did not observe a significant effect on
was set as the highest inclination of the board whereby subjects per- architectural changes of the medial gastrocnemius (FL: stretching:
ceived their calf muscles to be fully stretched. Post-training results pre = 54 mm, post = 54 mm; control: pre = 54 mm, post = 54 mm.
showed a significant (p = .04) improvement in MT for the experi- PA: stretching: pre = 22°, post = 24°; control: pre = 22°, post = 24°)
mental group (+5.8%), whereas the control group showed no change or soleus (FL: stretching: pre = 33 mm, post = 35 mm; control:
(p = .41). pre = 38 mm, post = 36 mm. PA: stretching: pre = 24°, post = 22°;
Alternatively, Simpson et al. (2017) explored the effects of a control: pre = 27°, post = 27°) after the 24-week intervention (self-
loaded stretch training. Therefore, 21 young men were randomized performed daily stretching, 4 × 360 s per day). Similarly, Mizuno
to either stretching (n = 11) or control (n = 10) groups. The stretch (2019) did not observe significant changes in PA of the medial gas-
training was carried out on the non-dominant leg and consisted trocnemius, despite an increase in MT, after 8 weeks of stretch
of 180 s of static stretching in a leg press loaded with 20% of the training (4 × 330 s, 3 sessions/week).
maximum voluntary isometric contraction. Training was carried out With respect to the thigh musculature, Lima et al. (2015) found
5 times a week over the 6-week study period. The authors reported no significant benefit to performing 8 weeks of stretch training on
a statistically significant increase of 5.6% in MT for the stretch train- both vastus lateralis FL (stretching: pre = 90 mm, post = 83 mm; con-
ing group; however, raw data were not shown (Simpson et al., 2017). trol: pre = 78 mm, post = 69 mm) and biceps femoris FL (stretching:
Subsequent to a letter to the editor that questioned the study's pre = 81, post = 78; control: pre = 88, post = 83 mm) in 24 healthy,
findings (Nunes, Nakamura, Schoenfeld, & Cyrino, 2018), raw data physically active men (stretching group, n = 12; control group,
supplied by the authors indicated that the average increases be- n = 12). In a pilot study of 10 young adults (stretching group, n = 5;
tween baseline and week 6 were actually very similar, equating to control group, n = 5), Freitas and Mil-Homens (2015) reported that
5.9% (1.06 mm) for the stretch training group and 7.6% (1.19 mm) 8 weeks of intense, long-duration (450 s per set), passive stretch
for the control group (Jakobi, Simpson, Smart, & O'Connor, 2018; training had no effect on PA (p = .13), but increased biceps femoris
Table 1). Thus, the conclusions of this study should be interpreted FL of 13.7% when compared with baseline's value (p = .04), whereas
with caution. no significant differences were seen in a non-training control group.
Although the ability to draw strong inferences from these data was
limited by the low sample size (5 subjects each group), it should be
3.2 | Effect of stretch training on muscle noted that stretch training-induced increases in FL were higher than
architecture the minimal detectable change. Similarly, Simpson et al. (2017) re-
ported significant increases of 25% and 5% on FL near the gastroc-
Blazevich et al. (2014) assigned 24 men to stretch training (n = 15) nemii muscle–tendon junction and in the muscle belly, respectively,
or control (n = 9) groups. The stretch training group stretched their after 6 weeks of loaded stretch training in the leg press machine.
calf muscles against a wall for 4 × 30 s with 15 s of rest. This proto- Interestingly, PA significantly decreased in the lateral gastrocnemius
col was performed twice daily (morning and evening) for 3 weeks. while increasing in the medial gastrocnemius near the muscle–ten-
Results showed stretch training had no significant effect on FL don junction but remaining the same in the muscle belly (Simpson
(stretching: pre = 48 mm, post = 46 mm; control: pre = 49 mm, et al., 2017).
post = 49 mm). The relatively short duration of the protocol raises Nakamura et al. (2012) randomly assigned 18 men to either
questions as to whether the time frame was sufficient to realize 4 weeks of gastrocnemii static stretch training (n = 9) or control
significant results. In a longer duration study, Konrad and Tilp (n = 9) groups. The stretch training programme consisted of having
(2014b) investigated the effects of static stretch training (against- subjects stand with arms supported against a wall, keeping the fore-
the-wall static calf stretches, 4 × 30 s, 5 sessions/week for foot resting on a platform and the ankle joint progressively dorsi-
6 weeks) in a cohort of 49 police cadets. Consistent with the find- flexed by leaning towards the wall until their self-perceived largest
ings of Blazevich et al. (2014), no statistical differences in pre- to tolerable stretch. Results showed no significant effect of stretch
 |
152       NUNES et al.

training on FL of the gastrocnemii, either when measured with the intensity of the protocol seems to be an important factor in pro-
ankle joint positioned at 0° or 30°. There were also noted no ef- moting stretch-induced muscle hypertrophy in humans (Freitas &
fects on resolved-FL, whereas there was observed an increase in Mil-Homens, 2015; Mizuno, 2019; Simpson et al., 2017). Moreover,
factors associated with muscle–tendon unit flexibility other than FL results from Moltubakk (2019) further indicate that longer inter-
(i.e., muscle–tendon junction displacement - ∆ resolved FL) for the ventions do not compensate for the low intensity of the self-per-
stretch training compared with the control group. formed stretching, as training lasted 24 weeks without any observed
changes in muscle architecture.
Of the eight studies that investigated changes in FL, only two
4 |  D I S CU S S I O N A N D CO N C LU S I O N S observed a pre- to post-stretch-training effect, and both were those
that the stretching was aided by some device (Freitas & Mil-Homens,
The purpose of this review was to evaluate the effect of stretch 2015; Simpson et al., 2017). In the other six studies (Blazevich et al.,
training on inducing muscle hypertrophy based on data from the 2014; Konrad & Tilp, 2014a, 2014b; Lima et al., 2015; Moltubakk,
literature. Of the 10 studies that met inclusion criteria, 3 observed 2019; Nakamura et al., 2012), the stretching was self-performed and
positive effects in some measure of muscle growth (Freitas & Mil- at an intensity within the tolerance level of the subjects. Indeed, a po-
Homens, 2015; Mizuno, 2019; Simpson et al., 2017). Therefore, it is tential explanation for the null findings in these studies may be that
suggested that the stretch training can induce muscle hypertrophy; self-performed passive stretching is an insufficient stimulus (Fowles
however, the way that the stretching is performed seems to influ- et al., 2000) for triggering important mechanisms for muscle hy-
ence the adaptations. pertrophy (Dankel et al., 2017; Wackerhage, Schoenfeld, Hamilton,
Various factors in the experimental designs differed between Lehti, & Hulmi, 2019). Moreover, although research in animal and in
the studies (Table 2). Intervention protocol of the works included vitro models has demonstrated that stretching can increase anabolic
in this review had a duration ranging from 3 to 24 weeks, and the signalling (Atherton et al., 2009; Sakamoto, Aschenbach, Hirshman,
total time under stretching (TTUS) of the programmes ranged from & Goodyear, 2003), human studies have failed to show significant
36 min to about 11 hr (i.e., time of each stretching set × number of elevations in the fractional muscle protein synthetic rate after maxi-
sets of each session × number of sessions). Results from Simpson mum tolerable stretching exercise (Fowles et al., 2000).
et al. (2017), in which TTUS was 90 min (6 weeks), indicated an in- In both the works of Akagi and Takahashi (2014) and Mizuno
crease in the FL, while other studies with similar TTUS (Blazevich et (2019), stretching was performed for the calf muscles using stretch-
al., 2014) or duration in weeks (Konrad & Tilp, 2014a, 2014b) did not. ing boards; however, only the latter study observed a hypertrophic
In addition, Moltubakk (2019) also did not see any effect on muscle effect. Some differences in experimental protocols may explain
structure after 24 weeks of self-performed stretch training. Thus, discrepancies between findings. First, subjects in Mizuno (2019)
within fairly wide limits, the training volume does not seem to be trained for 8 weeks, while the duration of the training for Akagi and
a sole determining factor, and other elements seem to play larger Takahashi (2014) lasted just 5 weeks. Moreover, Akagi and Takahashi
roles in promoting hypertrophic responses, such as the type or the (2014) used a convex probe to measure the MT of the ankle flexors
intensity of the stretching exercise. (from the adipose tissue–muscle interface of the posterior lower leg
The stretch training protocols of the studies analysed herein can to the muscle–bone interface), whereas Mizuno (2019) used a lin-
be classified as either (a) dynamic or static, and (b) self-performed ear probe and measured only the lateral gastrocnemius. Considering
or aided by a device. Regarding the first category, it is difficult to that muscle portions may respond non-uniformly to the muscle
draw inferences, given that only one study investigated dynamic lengthening (Franchi, Raiteri, et al., 2018; Franchi, Ruoss, et al., 2018;
stretch training (Konrad & Tilp, 2014a); nonetheless, results showed Simpson et al., 2017), this may at least in part help to explain such
no effect on muscle architecture. With regard to the second fac- differences. Additionally, in the study of Mizuno (2019), the stretch
tor, the 3 studies that demonstrated a hypertrophic effect, all used was carried out at an ankle angle whereby the calf muscles were
some external apparatus to confer an overload to stretching (Freitas fully stretched, while the stretch in Akagi and Takahashi (2014) was
& Mil-Homens, 2015; Mizuno, 2019; Simpson et al., 2017). Freitas performed with a ~10% reduction in this angle and intensity. This
and Mil-Homens (2015) employed an intense, long-duration passive reinforces the potential importance of ensuring high tensile strain
stretch training, and the design of Simpson et al. (2017) incorporated and/or muscle stress during the stretching if the goal is to induce
a loaded stretch, and in the study of Mizuno (2019), the stretching changes in muscle structure.
was performed using a calf-stretching board (“loaded” by the body- Another noteworthy point is that the work of Mizuno (2019) was
weight). Two studies observed a significant effect of stretch training the only one that actually observed increases in MT, although the
on MT (Mizuno, 2019; Simpson et al., 2017), albeit with the caveat PA remained unchanged. It is important to consider that increases in
that raw data of the study of Simpson et al. that demonstrated MT may occur without changes in PA (in the case of FL elongation),
similar adaptive responses on MT between stretching and control whereas increases in MT may occur without changes in FL (in the
groups (Jakobi et al., 2018; Nunes et al., 2018), and two studies case of PA elevation), and no change in MT may occur concurrently
observed positive adaptations in FL (Freitas & Mil-Homens, 2015; with increases in FL (in the case of PA diminution). Thus, although the
Simpson et al., 2017). The results of these studies indicate that the FL was not measured, it remains conceivable that increases would
NUNES et al. |
      153

also be observed in this outcome (Mizuno, 2019). It is customary to TA B L E 3   Effects of stretch training associated with resistance
use equations for estimating the FL when analysing muscles that are training programmes on muscle hypertrophy

large in length or when using a probe with a small width or field of Hypertrophic
view, as employed in some studies included herein (Freitas & Mil- Studies effects Observations
Homens, 2015; Lima et al., 2015; Nakamura et al., 2012). However, Kubo et al. (2002b) ↔ Away from RT session
some equations display better validity than others (Kumagai et Ferreira-Júnior et al. ↔ Before RT session
al., 2000; Noorkoiv, Stavnsbo, Aagaard, & Blazevich, 2010; Potier, (2019) (↔vol)
Alexander, & Seynnes, 2009); thus, extrapolation of FL results Moriggi Junior et al. ↓ Before RT session
should be made with caution (Ando et al., 2014; Franchi, Raiteri, et (2017) (↓vol)
al., 2018), as well as the comparison of findings between studies that Silva et al. (2014) ↑ Interset-rest of RT
employ this procedure but use different equations. Future studies exercises
should consider the use of appropriate measurement techniques de- Evangelista et al. (2019) ↑ Interset-rest of RT
pending on the size of the muscle to be analysed (Franchi, Raiteri, et exercises

al., 2018). Note: Summary of the findings of the studies available in the literature.
Despite the observed beneficial effects of stretch training on FL, ↑ = the outcome variable was improved with the addition of stretching
to an RT programme. ↓ = the outcome variable was blunted with the
the findings are not necessarily due to an increase in muscle fibre
addition of stretching to an RT programme. ↔ = the outcome variable
length. After eccentric training, Franchi et al. (2014); Franchi, Ruoss, presented similar responses between the RT + stretching training and
et al. (2018) observed higher growth in vastus lateralis cross-sectional the stretch training only. ↔vol = no effect on training volume when
area at the distal site compared with the muscle belly. Likewise, they stretching was added before the RT. ↓vol = negative effect on training
volume when stretching was added before the RT.
reported that the content of activated costamere-associated proteins
Abbreviation: RT, resistance training.
increased more prominently at the distal site. This phenomenon, as
suggested by the authors, may indicate a potential to preferentially
increase the amount of myofibril Z-bands in series (thus reflecting an same resistance training programme in the contralateral leg without
FL increase) at the distal portion of the muscles (Franchi, Ruoss, et al., preworkout stretching. The authors speculated that the diminished
2018). Although stretch training seems to elicit different responses at hypertrophic effect seems to be mediated by the reduction in the
different sites along the muscle (Simpson et al., 2017), fascicle elon- total resistance training volume, which is postulated to be a primary
gation also may be a consequence of decreases in stiffness of the driver of training-induced muscle growth (Figueiredo, Salles, & Trajano,
connective tissue, particularly the perimysium, which is the largest 2018; Schoenfeld & Grgic, 2018). Alternatively, Ferreira-Júnior et al.
extracellular contributor of tissue stiffness (Akagi & Takahashi, 2014; (2019) explored the influence of adding static or dynamic stretching
Purslow, 1989). Alterations may be attributed to increases in non-con- exercises for biceps femoris (~80 s, 2 sessions/week) before the seated
tractile properties, viscoelastic components of the muscle–tendon leg curl resistance exercise in untrained young men. Compared with
unit, collagen fibres and/or in other factors/components of the mus- a third group, which performed only bouts of resistance exercise (i.e.,
cle–tendon unit beside the muscle fibre length. Moreover, increases in non-stretch control group), stretch training groups demonstrated no
these viscoelastic components are more abundant in close proximity impairments in biceps femoris growth after the 8-week training period.
to the muscle–tendon junction (DeDeyne, 2001; Franchi, Atherton, Also, in this case, performing stretching did not diminish the total resis-
Maganaris, & Narici, 2016; Kubo, Kanehisa, & Fukunaga, 2002a; tance training volume (Ferreira-Júnior et al., 2019). Conversely, Kubo
Morse, Degens, Seynnes, Maganaris, & Jones, 2007; Nakamura et al., et al. (2002b) found that performing stretching twice daily 5 × 45 s did
2012; Purslow, 1989). It remains to be determined whether the ad- not impair the hypertrophic response when combined with resistance
aptations obtained after bouts of passive stretching without muscle training (unilateral plantar flexion at 70% of 1RM with 5 × 10 repeti-
contraction are the same as with lengthening contraction training. It tions, 4 sessions/week) compared with a resistance training-only pro-
is necessary to explore the influence of applying an external overload tocol (2.9% vs. 3.1% for resistance training and resistance + stretch
during stretching, especially at the molecular level (Haun et al., 2019), training, respectively). The discrepant findings between studies make
in order to establish the adaptative response of skeletal muscle fibres it difficult to draw strong conclusions as to how the combination of
in response to this kind of training. stretch training and resistance training impacts the relationship be-
In an effort to provide additional insights on the topic, several stud- tween volume and skeletal muscle hypertrophy.
ies have endeavoured to determine how stretch training impacts hyper- In a non-peer-reviewed work presented as a conference abstract,
trophy in conjunction with regimented resistance training (Evangelista Silva et al. (2014) randomly assigned 24 trained men to interset-stretch-
et al., 2019; Ferreira-Júnior et al., 2019; Kubo, Kanehisa, & Fukunaga, ing or non-stretching control conditions in combination with a resis-
2002b; Moriggi Junior, Berton, Souza, Chacon-Mikahil, & Cavaglieri, tance training programme. In both conditions, subjects performed
2017; Silva et al., 2014; Table 3). For example, Moriggi Junior et al. 4 sets of plantar flexion exercise for 8-12RM on a leg press machine
(2017) found that performing 2 × 25 s of static stretching immediately twice a week for 5 weeks. For the interset-stretching condition, sub-
before resistance training for 10 weeks blunted quadriceps muscle jects maintained the weight of the leg press in a dorsiflexed position for
hypertrophy in young, healthy males compared with performing the 30 s between sets, while subjects in the control group passively rested
 |
154       NUNES et al.

during this period. Results showed markedly greater hypertrophic in- assessing changes at different sites of the muscles (i.e., proximal,
creases favouring the group that performed loaded interset-stretch belly and distal) would be helpful for elucidating subtle alterations in
compared with control (23% vs. 9%, respectively). Evangelista et al. skeletal muscle adaptations, particularly with respect to FL. Further,
(2019) recently compared the effects of an 8-week traditional whole investigations at the microstructural level (i.e., via biopsy sampling)
body resistance training programme (6 exercises, 4 × 8–12 repetitions, may shed more light on the potential effect of stretch training on
2 sessions/week) to the same protocol employing interset-stretch human skeletal muscle hypertrophy (Haun et al., 2019).
training (30 s of passive stretching during each 90 s interset-rest pe- In conclusion, the available literature indicates that passive,
riod) on muscular adaptations in men without experience on resis- low-intensity stretch does not appear to confer beneficial changes
tance training. Results showed that MT increased similarly between in muscle size and architecture. Alternatively, the current evidence
conditions for the biceps brachii, triceps brachii and rectus femoris. suggests that intense stretch training, particularly when loaded, or
Alternatively, significantly greater increases in vastus lateralis MT were added between active muscle contractions, may elicit muscle hy-
observed favouring the interset-stretch training group. pertrophy; however, the relative paucity of research implementing
Notably, both studies that showed an hypertrophic effect em- such protocols precludes the ability to draw strong inferences on
ployed stretching in the interset-rest period (Evangelista et al., 2019; the topic. Future studies using high-intensity passive stretch training
Silva et al., 2014), while those that did not observe a positive effect protocols should be carried out to fill existing gaps in the literature
involved performing stretching immediately before resistance exer- and thus provide greater insight into the potential adaptations and
cise (Ferreira-Júnior et al., 2019; Moriggi Junior et al., 2017) or away their practical applicability.
from the resistance training session time period (Kubo et al., 2002b).
In this regard, it seems that stretch training when performed in com- C O N FL I C T O F I N T E R E S T
bination with resistance training may induce some additional mus- The authors declare no conflict of interest.
cle growth. Together, these results indicate that stretch-induced
structural adaptations only occur after a minimum threshold of ORCID
stimulus is reached either by stretching itself or by adding stretch- João Pedro Nunes  https://orcid.org/0000-0001-8144-5906
ing exercises between the executions of resistance training sets. Masatoshi Nakamura  https://orcid.org/0000-0002-8184-1121
The underpinning mechanical and metabolic factors that may play
a role in stretch training-induced muscle growth are described in REFERENCES
several works (Antonio & Gonyea, 1993a, 1993b; DeDeyne, 2001; Akagi, R., & Takahashi, H. (2014). Effect of a 5-week static stretching pro-
Goldberg et al., 1975; Kelley, 1996; Lowe & Alway, 2002; Riley & gram on hardness of the gastrocnemius muscle. Scandinavian Journal
of Medicine & Science in Sports, 24, 950–957. https​://doi.org/10.1111/
Van Dyke, 2012; Tatsumi, 2010; Wisdom et al., 2015). On the other
sms.12111​
hand, the potentiated muscle growth effect of stretching added be- Ando, R., Taniguchi, K., Saito, A., Fujimiya, M., Katayose, M., & Akima,
tween resistance training sets seems to be related to the restriction H. (2014). Validity of fascicle length estimation in the vastus lat-
of blood flow on interset-rest period, conceivably by augmenting eralis and vastus intermedius using ultrasonography. Journal
of Electromyography and Kinesiology, 24, 214–220. https​ ://doi.
metabolite accumulation (Dankel et al., 2017) and/or regulating ana-
org/10.1016/j.jelek​in.2014.01.003
bolic-related signalling pathways (Mohamad et al., 2011). Antonio, J., & Gonyea, W. J. (1993a). Progressive stretch overload of
When assessing gaps in the current literature, future works should skeletal muscle results in hypertrophy before hyperplasia. Journal
consider the use of a device for application of external overload, such of Applied Physiology, 75, 1263–1271. https​ ://doi.org/10.1152/
jappl.1993.75.3.1263
as those used by Freitas and Mil-Homens (2015) and Simpson et al.
Antonio, J., & Gonyea, W. J. (1993b). Skeletal muscle fiber hyperplasia.
(2017), or the use of an apparatus that aids in the performance of the Medicine & Science in Sports & Exercise, 25, 1333–1345. https​://doi.
stretching protocol, as used by Mizuno (2019). Measuring the elec- org/10.1249/00005​768-19931​2000-00004​
tromyographic signal is also a potentially viable strategy to help en- Atherton, P. J., Szewczyk, N. J., Selby, A., Rankin, D., Hillier, K., Smith, K.,
sure that exercise involves a truly passive stretch rather than active … Loughna, P. T. (2009). Cyclic stretch reduces myofibrillar protein
synthesis despite increases in FAK and anabolic signalling in L6 cells.
muscle contraction, as employed by Simpson et al. (2017). Additional
The Journal of Physiology, 587, 3719–3727. https​://doi.org/10.1113/
studies are needed that combine stretch training with resistance jphys​iol.2009.169854
training to assess whether the combination potentiates muscular ad- Blazevich, A. J., Cannavan, D., Waugh, C. M., Miller, S. C., Thorlund, J.
aptations compared with resistance training alone. Moreover, there B., Aagaard, P., & Kay, A. D. (2014). Range of motion, neuromechan-
ical, and architectural adaptations to plantar flexor stretch training
is evidence that stretch training possibly activates resident myogenic
in humans. Journal of Applied Physiology, 117, 452–462. https​://doi.
stem cell (DeDeyne, 2001; Tatsumi, 2010) and their differentiation org/10.1152/jappl​physi​ol.00204.2014
into myonuclei, which has been implicated in increasing muscle hy- Conceição, M. S., Vechin, F. C., Lixandrão, M., Damas, F., Libardi, C.
pertrophy capacity (Conceição et al., 2018). A cross-limb study, using A., Tricoli, V., … Ugrinowitsch, C. (2018). Muscle fiber hypertrophy
and myonuclei addition: A systematic review and meta-analysis.
one limb for a high-intensity stretch training period and another for
Medicine & Science in Sports & Exercise, 50, 1385–1393. https​://doi.
control, in which both would be thereafter submitted to a resistance org/10.1249/MSS.00000​0 0000​0 01593
training programme, could help to determine whether stretch train- Dankel, S. J., Mattocks, K. T., Jessee, M. B., Buckner, S. L., Mouser, J. G.,
ing enhances resistance training-induced muscle growth. In addition, & Loenneke, J. P. (2017). Do metabolites that are produced during
NUNES et al. |
      155

resistance exercise enhance muscle hypertrophy? European Journal Medicine & Science in Sports, 28, 2769–2771. https​://doi.org/10.1111/
of Applied Physiology, 117, 2125–2135. https​ ://doi.org/10.1007/ sms.13300​
s00421-017-3690-1 Kelley, G. (1996). Mechanical overload and skeletal muscle fiber hyper-
DeDeyne, P. G. (2001). Application of passive stretch and its implication plasia: A meta-analysis. Journal of Applied Physiology, 81, 1584–1588.
for muscle fibers. Physical Therapy, 81, 819–827. https​://doi.org/10.1152/jappl.1996.81.4.1584
Evangelista, A. L., De Souza, E. O., Moreira, D. C. B., Alonso, A. C., Konrad, A., & Tilp, M. (2014a). Effects of ballistic stretching training on
Teixeira, C. V. L. S., Wadhi, T., … Greve, J. M. D. A. (2019). Inter-set the properties of human muscle and tendon structures. Journal of
stretching versus traditional strength training: Effects on muscle Applied Physiology, 117, 29–35. https​://doi.org/10.1152/jappl​physi​
strength and hypertrophy in untrained individuals. Journal of Strength ol.00195.2014
and Conditioning Research, 33, S159–S166. https​://doi.org/10.1519/ Konrad, A., & Tilp, M. (2014b). Increased range of motion after static
JSC.00000​0 0000​0 03036 stretching is not due to changes in muscle and tendon structures.
Ferreira-Júnior, J. B., Benine, R. P. C., Chaves, S. F. N., Borba, D. A., Martins- Clinical Biomechanics, 29, 636–642. https​://doi.org/10.1016/j.clinb​
Costa, H. C., Freitas, E. D. S., … Bottaro, M. (2019). Effects of static iomech.2014.04.013
and dynamic stretching performed before resistance training on mus- Kubo, K., Kanehisa, H., & Fukunaga, T. (2002a). Effect of stretching train-
cle adaptations in untrained men. Journal of Strength and Conditioning ing on the viscoelastic properties of human tendon structures in vivo.
Research.https​://doi.org/10.1519/JSC.00000​0 0000​0 03283 Journal of Applied Physiology, 92, 595–601. https​://doi.org/10.1152/
Figueiredo, V. C., de Salles, B. F., & Trajano, G. S. (2018). Volume for jappl​physi​ol.00658.2001
muscle hypertrophy and health outcomes: The most effective vari- Kubo, K., Kanehisa, H., & Fukunaga, T. (2002b). Effects of resistance
able in resistance training. Sports Medicine, 48, 499–505. https​://doi. and stretching training programmes on the viscoelastic proper-
org/10.1007/s40279-017-0793-0 ties of human tendon structures in vivo. Journal of Physiology, 538,
Fowles, J. R., MacDougall, J. D., Tarnopolsky, M. A., Sale, D. G., Roy, B. 219–226.
D., & Yarasheski, K. E. (2000). The effects of acute passive stretch Kumagai, K., Abe, T., Brechue, W. F., Ryushi, T., Takano, S., & Mizuno,
on muscle protein synthesis in humans. Canadian Journal of Applied M. (2000). Sprint performance is related to muscle fascicle length
Physiology, 25, 1. https​://doi.org/10.1139/h00-012 in male 100-m sprinters. Journal of Applied Physiology, 88, 811–816.
Franchi, M. V., Atherton, P. J., Maganaris, C. N., & Narici, M. V. (2016). https​://doi.org/10.1152/jappl.2000.88.3.811
Fascicle length does increase in response to longitudinal resistance Lima, K. M. M., Carneiro, S. P., Alves, D. D. S., Peixinho, C. C., & de Oliveira,
training and in a contraction-mode specific manner. Springerplus, 5, L. F. (2015). Assessment of muscle architecture of the biceps femoris
1–3. https​://doi.org/10.1186/s40064-015-1548-8 and vastus lateralis by ultrasound after a chronic stretching program.
Franchi, M. V., Atherton, P. J., Reeves, N. D., Flück, M., Williams, J., Clinical Journal of Sport Medicine, 25, 55–60. https​://doi.org/10.1097/
Mitchell, W. K., … Narici, M. V. (2014). Architectural, functional JSM.00000​0 0000​0 00069
and molecular responses to concentric and eccentric loading in Lowe, D. A., & Alway, S. E. (2002). Animal models for inducing muscle
human skeletal muscle. Acta Physiologica, 210, 642–654. https​://doi. hypertrophy: Are they relevant for clinical applications in humans?
org/10.1111/apha.12225​ Journal of Orthopaedic & Sports Physical Therapy, 32, 36–43. https​://
Franchi, M. V., Raiteri, B. J., Longo, S., Sinha, S., Narici, M. V., & Csapo, doi.org/10.2519/jospt.2002.32.2.36
R. (2018). Muscle architecture assessment: Strengths, shortcom- Magnusson, P., & Renström, P. (2006). The European college of sports
ings and new frontiers of in-vivo imaging techniques. Ultrasound in sciences position statement: The role of stretching exercises in
Medicine & Biology, 44, 2492–2504. https​://doi.org/10.1016/j.ultra​ sports. European Journal of Sport Science, 6, 87–91. https​ ://doi.
smedb​io.2018.07.010 org/10.1080/17461​39060​0617865
Franchi, M. V., Ruoss, S., Valdivieso, P., Mitchell, K. W., Smith, K., Magnusson, S. P., Simonsen, E. B., Aagaard, P., Sørensen, H., & Kjaer, M.
Atherton, P. J., … Flück, M. (2018). Regional regulation of focal ad- (1996). A mechanism for altered flexibility in human skeletal muscle.
hesion kinase after concentric and eccentric loading is related to re- The Journal of Physiology, 497, 291–298. https​ ://doi.org/10.1113/
modelling of human skeletal muscle. Acta Physiologica, 223, e13056. jphys​iol.1996.sp021768
https​://doi.org/10.1111/apha.13056​ Medeiros, D. M., & Lima, C. S. (2017). Influence of chronic stretching on
Freitas, S. R., & Mil-Homens, P. (2015). Effect of 8-week high-intensity muscle performance: Systematic review. Human Movement Science,
stretching training on biceps femoris architecture. Journal of Strength 54, 220–229. https​://doi.org/10.1016/j.humov.2017.05.006
and Conditioning Research, 29, 1737–1740. https​://doi.org/10.1519/ Mizuno, T. (2019). Combined effects of static stretching and electrical
JSC.00000​0 0000​0 00800 stimulation on joint range of motion and muscle strength. Journal
Goldberg, A. L., Etlinger, J. D., Goldspink, D. F., & Jablecki, C. (1975). of Strength and Conditioning Research, 33, 2694–2703. https​://doi.
Mechanism of work-induced hypertrophy of skeletal muscle. org/10.1519/JSC.00000​0 0000​0 02260
Medicine and Science in Sports and Exercise, 7, 185–198. https​://doi. Mohamad, N. I., Nosaka, K., & Cronin, J. (2011). Maximizing hypertro-
org/10.1249/00005​768-19750​0730-00016​ phy: Possible contribution of stretching in the interset rest period.
Goldspink, G., Tabary, C., Tabary, J. C., Tardieu, C., & Tardieu, G. (1974). Strength and Conditioning Journal, 33, 81–87. https​://doi.org/10.1519/
Effect of denervation on the adaptation of sarcomere number and SSC.0b013​e3181​fe7164
muscle extensibility to the functional length of the muscle. The Moltubakk, M. M. H. (2019). Effects of long-term stretching training on
Journal of Physiology, 236, 733–742. https​://doi.org/10.1113/jphys​ muscle-tendon morphology, mechanics and function (Papers 3 and 4;
iol.1974.sp010463 unpublished). PhD Dissertation from the Norwegian School of Sport
Haun, C. T., Vann, C. G., Roberts, B. M., Vigotsky, A. D., Schoenfeld, Sciences, p. 214. ISBN: 978-82-502-0564-2.
B. J., & Roberts, M. D. (2019). A critical evaluation of the biologi- Moriggi Junior, R., Berton, R., de Souza, T. M. F., Chacon-Mikahil, M. P. T.,
cal construct skeletal muscle hypertrophy: Size matters but so & Cavaglieri, C. R. (2017). Effect of the flexibility training performed
does the measurement. Frontiers in Physiology, 10, 247. https​://doi. immediately before resistance training on muscle hypertrophy, max-
org/10.3389/fphys.2019.00247​ imum strength and flexibility. European Journal of Applied Physiology,
Jakobi, J. M., Simpson, C. L., Smart, R. R., & O'Connor, B. (2018). Response 117, 767–774. https​://doi.org/10.1007/s00421-016-3527-3
to Nunes and colleagues letter: The data do not seem to support the Morse, C. I., Degens, H., Seynnes, O. R., Maganaris, C. N., & Jones, D. A.
effect of stretch training in increasing MT. Scandinavian Journal of (2007). The acute effect of stretching on the passive stiffness of the
 |
156       NUNES et al.

human gastrocnemius muscle tendon unit. The Journal of Physiology, and Conditioning Journal, 40, 107–112. https​ ://doi.org/10.1519/
586, 97–106. https​://doi.org/10.1113/jphys​iol.2007.140434 SSC.00000​0 0000​0 00363
Nakamura, M., Ikezoe, T., Takeno, Y., & Ichihashi, N. (2012). Effects of a 4-week Silva, J., Lowery, R., Antonio, J., McCleary, S., Rauch, J., Ormes, J., …
static stretch training program on passive stiffness of human gastrocne- Wilson, J. (2014). Weighted post-set stretching increases skeletal
mius muscle-tendon unit in vivo. European Journal of Applied Physiology, muscle hypertrophy (NSCA 2014 annual meeting). The Journal of
112, 2749–2755. https​://doi.org/10.1007/s00421-011-2250-3 Strength and Conditioning Research, 28, 65.
Noorkoiv, M., Stavnsbo, A., Aagaard, P., & Blazevich, A. J. (2010). In vivo Simpson, C. L., Kim, B. D. H., Bourcet, M. R., Jones, G. R., & Jakobi, J. M.
assessment of muscle fascicle length by extended field-of-view ul- (2017). Stretch training induces unequal adaptation in muscle fasci-
trasonography. Journal of Applied Physiology, 109, 1974–1979. https​ cles and thickness in medial and lateral gastrocnemii. Scandinavian
://doi.org/10.1152/jappl​physi​ol.00657.2010 Journal of Medicine & Science in Sports, 27, 1597–1604. https​://doi.
Nunes, J. P., Nakamura, M., Schoenfeld, B. J., & Cyrino, E. S. (2018). The org/10.1111/sms.12822​
data do not seem to support the effect of stretch training on increas- Tatsumi, R. (2010). Mechano-biology of skeletal muscle hypertrophy and
ing muscle thickness. Scandinavian Journal of Medicine & Science in regeneration: Possible mechanism of stretch-induced activation of
Sports, 28, 2767–2768. https​://doi.org/10.1111/sms.13285​ resident myogenic stem cells. Animal Science Journal, 81, 11–20. https​://
Potier, T. G., Alexander, C. M., & Seynnes, O. R. (2009). Effects of eccen- doi.org/10.1111/j.1740-0929.2009.00712.x
tric strength training on biceps femoris muscle architecture and knee Wackerhage, H., Schoenfeld, B. J., Hamilton, D. L., Lehti, M., & Hulmi, J.
joint range of movement. European Journal of Applied Physiology, 105, J. (2019). Stimuli and sensors that initiate skeletal muscle hypertro-
939–944. https​://doi.org/10.1007/s00421-008-0980-7 phy following resistance exercise. Journal of Applied Physiology, 126,
Purslow, P. P. (1989). Strain-induced reorientation of an intramus- 30–43. https​://doi.org/10.1152/jappl​physi​ol.00685.2018
cular connective tissue network: Implications for passive mus- Wisdom, K. M., Delp, S. L., & Kuhl, E. (2015). Use it or lose it: Multiscale
cle elasticity. Journal of Biomechanics, 22(1), 21–31. https​ ://doi. skeletal muscle adaptation to mechanical stimuli. Biomechanics and
org/10.1016/0021-9290(89)90181-4 Modeling in Mechanobiology, 14, 195–215. https​://doi.org/10.1007/
Riley, D. A., & Van Dyke, J. M. (2012). The effects of active and pas- s10237-014-0607-3
sive stretching on muscle length. Physical Medicine and Rehabilitation
Clinics of North America, 23, 51–57. https​ ://doi.org/10.1016/j.
pmr.2011.11.006
How to cite this article: Nunes JP, Schoenfeld BJ, Nakamura
Sakamoto, K., Aschenbach, W. G., Hirshman, M. F., & Goodyear, L. J.
(2003). Akt signaling in skeletal muscle: Regulation by exercise
M, Ribeiro AS, Cunha PM, Cyrino ES. Does stretch training
and passive stretch. American Journal of Physiology-Endocrinology induce muscle hypertrophy in humans? A review of the
and Metabolism, 285, 1081–1088. https​ ://doi.org/10.1152/ajpen​ literature. Clin Physiol Funct Imaging. 2020;40:148–156. https​://
do.00228.2003 doi.org/10.1111/cpf.12622​
Schoenfeld, B. J., & Grgic, J. (2018). Evidence-based guidelines for re-
sistance training volume to maximize muscle hypertrophy. Strength