Journal of Science in Sport and Exercise

https://doi.org/10.1007/s42978-022-00191-z

REVIEW ARTICLE

Long‑Lasting Stretching Induces Muscle Hypertrophy: A Meta‑Analysis

of Animal Studies
Konstantin Warneke1 · Philipp Alexander Freund2 · Stephan Schiemann1

Received: 17 February 2022 / Accepted: 7 August 2022

© The Author(s) 2022

Abstract
Muscular hypertrophy depends on metabolic exhaustion as well as mechanical load on the muscle. Mechanical tension
seems to be the crucial factor to stimulate protein synthesis. The present meta-analysis was conducted to determine whether
stretching can generate adequate mechanical tension to induce muscle hypertrophy. We used PubMed, Web of Science, and
Scopus to search for literature examining the effects of long-term stretching on muscle mass, muscle cross-sectional area, fiber
cross-sectional area, and fiber number. Since there was no sufficient number of studies investigating long-lasting stretching
in humans, we only included original animal studies in the current meta-analysis. Precisely, we identified 16 studies meeting
the inclusion criteria (e. g. stretching of at least 15 min per day). The 16 studies yielded 39 data points for muscle mass, 11
data points for muscle cross-sectional area, 20 data points for fiber cross-sectional area, and 10 data points for fiber number.
Across all designs and categories, statistically significant increases were found for muscle mass (d = 8.51; 95% CI 7.11–9.91),
muscle cross-sectional area (d = 7.91; 95% CI 5.75–10.08), fiber cross-sectional area (d = 5.81; 95% CI 4.32–7.31), and fiber
number (d = 4.62; 95% CI 2.54–6.71). The findings show an (almost) continuous positive effect of long-term stretching on
the listed parameters, so that it can be assumed that stretch training with adequate intensity and duration leads to hypertrophy
and hyperplasia, at least in animal studies. A general transferability to humans—certainly with limited effectiveness—can
be hypothesized but requires further research and training studies.

Keywords Muscle mass · Static stretching · Hyperplasia · Chronic stretching

Introduction corresponding mechanical stimulus can be initiated not only

by high loads in strength training, but also through stretching
To achieve muscular hypertrophy, strength training needs— with appropriate intensity. Smith et al. [55] demonstrated
in addition to metabolic exhaustion—a high mechanical load that mechanical stress generated by stretching can be suf-
on the muscle, which leads to micro-traumatization of the ficient to induce delayed onset muscle soreness (DOMS)
muscle fibers [63]. In this regard, the crucial factor is high [55]. Accordingly, it can be assumed that stretching stimuli
mechanical tension on the muscle. Resulting hypertrophy can cause adequate micro-traumatization. The resulting
effects depend on an increased (myofibrillar) protein syn- repair processes can trigger hypertrophy-stimulating sign-
thesis rate, which is stimulated via corresponding signal- aling pathways to increase protein synthesis rates [29]. The
ing pathways. In particular, activation of the Akt/mTOR/ resulting activation of stretch-activated channels alters the
p70S6K signaling pathway appears to be of high impor- cytoplasmic membrane and initiates signal transduction pro-
tance for the stimulation of muscular protein synthesis and cesses via mTOR [59, 61].
is primarily induced by mechanical loading [1, 17, 46]. A Against this background, the following hypothesis can
be derived: stretch training performed with sufficient inten-
* Konstantin Warneke sity leads to high mechanical load that can trigger muscular
[email protected] hypertrophy as a long-term training effect. This hypothesis
has already been discussed previously: “It is well known
1
Institute for Exercise, Sport and Health, Leuphana that application of chronic stretch is a very potent model
University, Universitätsallee 1, Lüneburg, Germany
for inducing muscle enlargement” [36]. However, to date,
2
Institute of Psychology, Leuphana University, Lüneburg,
Germany

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studies examining adaptations of stretch training have gener- especially to check if the applicability of the training method
ally focused either on increasing range of motion (ROM), or appears worthwhile for human studies [31]. There is one
on other parameters describing flexibility [38, 40]. Moreo- meta-analysis available from Kelley [33] that has addressed
ver, acute effects of stretching interventions on muscular this issue before. In Kelley’s meta-analysis, however, the
performance mostly show negative effects regarding maxi- muscular overload was not generated exclusively by stretch-
mum strength and explosive power [13, 71]. ing but also by other methods (weight training, ablation), so
Initial human studies show that long-term stretching that no conclusion could be drawn about the specific effects
interventions for several weeks can induce hypertrophic of long-term stretching. Moreover, comprehensive analysis
effects and/or increase maximum strength. For example, on distinct outcomes such as MM, MCSA, FCSA and FN
Simpson et al. [53] were able to achieve an average increase are not available in the study by Kelley [33]. Consequently,
of 5.6% in muscle cross-sectional area through a stretching a distinct base of empirical evidence needs to be researched
intervention with a duration of three minutes, three days to investigate the questions of the present meta-analysis. In
per week, for 6 weeks. Panidi et al. [44] found an increase particular, the present meta-analysis of animal studies aims
in muscle cross sectional area (MCSA) of 23% ± 14% after to provide a comprehensive and differentiated overview of
a 12-week stretching intervention with stretching durations the effects of (continuous) stretching interventions on MM,
up to 15 min per training session. Nelson et al. [43] demon- MCSA and FCSA, and on hyperplasia effects (FN). Sub-
strated a 29% increase in maximal strength after stretching sequently, the relevance of these results with regard to the
the calf muscles for 4 × 30 s, 3 days a week for 10 weeks. In potential use of stretching training with the goal of muscle
addition, Kokkonen et al. [34] achieved significant improve- and strength building in athletic and therapeutic training will
ments in various performance tests, such as 1 RM knee be discussed.
extension and knee flexion, standing long jump, and high
jump, with static stretching for 40 min per session, 3 days
per week for 10 weeks.
Longitudinal studies using animal experiments have been Methods
available for some time and have demonstrated significant
hypertrophy effects after continuous stretching from 30 min The following search terms were defined to search PubMed,
to 24 h per day over an intervention period of several weeks, Web of Science, and Scopus databases: [(“hypertrophy” OR
reflected by an increase in muscle mass (MM), MCSA, fiber “hyperplasia”) AND (“stretch-induced growth” OR “stretch-
cross sectional area (FCSA) and/or hyperplasia effects with induced hypertrophy” OR “fiber number” OR “fiber length”
an increased fiber number (FN) [8, 10, 15, 23, 25]. Data OR “sarcomere length” OR “sarcomere number”) AND
of muscle weight were collected by removing the connec- “skeletal muscle”) NOT (“exercise induced” OR “endo-
tive tissue and weighing the wet muscle weight. MCSA and crine” OR “nervous system” OR “electrical stimulation”
FN were investigated by placing the muscle in a solution OR “cardiomyocytes”]. The search strategy was limited to
in which the different muscle fibers were stained in differ- English language sources only.
ent colors (fast twitch fiber stained lightly, slow twitch fib- A total of 89 publications were found from this combi-
ers stained darkly). Subsequently, the muscle cross-section nation of terms. The references found in these publications
and fiber cross-sectional area were determined from a given were examined for further relevant studies. However, this
number of fibers (for example 500 slow twitch and 200 fast did not yield any additional studies. After reviewing the
twitch fibers in Antonio et al. [10] using light microgra- titles, 47 studies remained, which were then screened to
phy images and an image analysis computer program). In exclude studies that only indirectly investigated structural
addition, in vitro condition a significant increase in maxi- adaptations and those studies that focused more on hormonal
mum strength was demonstrated by continuous stretching, adaptations, muscle fiber distribution, or signal transduction
so that these hypertrophy effects are functional in animals pathways without collecting the target parameters of mus-
[3, 4]. The muscle fiber type was determined by ATPase- cle mass, muscle cross-sectional area, fiber cross-sectional
activity using an ATPase staining method and fiber number area, fiber length, or fiber number. After this step, 23 studies
was investigated by counting fibers running from origin to remained, which were then subjected to full-text analysis
insertion [10] using inclusion and exclusion criteria established in advance
Since animal studies play a vital role in research to inves- of the meta-analysis for the final selection.
tigate human health, and systematic reviews or meta-analy- The following parameters were defined as inclusion
ses provide a suitable basis for drawing evidence-based con- criteria:
clusions concerning a research topic, we decided to create a - Objective measurement of muscle mass and/or mus-
transparent overview of the available information on effects cle cross-sectional area and/or fiber count and/or fiber
of long-lasting stretching intervention on muscle tissue,

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Fig. 1  Searching method via

PRISMA method

cross-sectional area and/or fiber length and/or number of information provided on blinding of the “care provider” and
muscle fibers. “outcome provider.” Only Czerwinski et al.[23] provided
- Stretching interventions of at least one week. information on randomization.
- Stretching times of at least 15 min per day.
- Specification of mean values and standard deviations. Meta‑analytic Procedure
- Studies on animals.
Accordingly, the following were considered exclusion Using the meta-analysis software RevMan, version 5.4.1
criteria: [22], 5 separate analyses were performed for the follow-
- No measurement of muscle mass and/or muscle cross- ing parameters: muscle mass, muscle cross-sectional area,
section and/or fiber number and/or fiber cross-section and/ muscle fiber cross-sectional area, muscle fiber length, and
or fiber length and/or number of muscle fibers. number of muscle fibers. The following parameters from
- Missing or insufficient information on the duration of each of the studies were included in the analysis: number of
the intervention and on the stretching times. experimental animals, and the respective mean values and
- Missing data concerning mean values and standard standard deviations of the experimental and control condi-
deviations, absence of absolute values. tions. Since several studies involved different durations, the
- Missing data of number of test animals. studies were listed in alphabetical order with a lowercase
- Missing control group/control condition. letter to allow assignment of the elongation period to the
The final sample in the meta-analysis included 16 stud- respective representation in the forest plot. We used a ran-
ies, whereby some studies with multiple effect sizes were dom effects model to take into account any heterogeneity
included in the analysis because they either included differ- resulting from the use of different species in the studies and
ent variables (e.g. muscle mass, fiber cross-section and/or all other potential between-study differences (study charac-
hyperplasia effects) or because they described the effects of teristics are summarized in Table 1).1
different intervention periods (a few days to several months). Tables 3–7 report the empirical M, SD and N for the
Figure 1 illustrates the procedure for study selection and parameters muscle mass, muscle cross sectional area, fiber
Table 1 details the included studies. cross sectional area, muscle fiber number, and fiber length.
For all analyses, the standardized mean difference (with
Quality Assessment inverse variance weighting) and its 95% confidence interval
were computed as the effect size of interest in RevMan.2
The quality assessment was based on the Delphi list [62]. Since for the evaluation of MM, MCSA, FCSA, FN and FL
The Delphi method was chosen as a reliable and valid tool in laboratory studies, animals had to be dissected and flight
for the assessment of the quality of the included studies [54]. muscles (ALD, PAT) had to be removed, no pre-post com-
The assessment items for the current meta-analyses can be parison of the same subjects could be performed. Therefore,
found in Table 2. The evaluation was performed by two inde-
pendent raters. If question 2 received an affirmative answer,
it was assumed that the age of the test animals, the species 1
In addition, we provide funnel plots for each outcome parameter as
or breed of the animal as well as the initial weight were
supplemental material to illustrate potential publication bias.
given. In all studies listed, mean values and standard devia- ( )
2
Using the formulae.SMDi = 1i s 2i 1 − 4N3−9 and
m −m
tions were given (see inclusion criteria) and the objective √ i i

of the study was clearly stated. In none of the studies was Ni SMD2i
+ 2 N −3.94 .
{ }
SE SMDi = n1i n2i ( i )

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 Table 1  Description of included studies
Source Subjects Muscle Group Intervention Measured Parameters

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Alway et al. [7] N = 63 ALD 7-day stretching intervention, nine animals MM: + 64% ± 8.4%
examined every day MCSA: + 29.9% ± 12.3%
FL: + 40.2% ± 2.2%
FN: + 27.3% ± 3%
Alway [2] N = 36, 22 in intervention group ALD 30-day stretching with 12% of bodyweight MM: + 161.5% ± 7.9%
FL: + 25.4% ± 4.6%
Alway [3] N = 24, 12 young (Y) and 12 old (O) quails ALD 30 days of unilateral stretching with 12% MM: YCG:26.7 ± 1.2 mg; YIG:71.6 ± 3.0 mg
bodyweight OCG:28.5 ± 1.5 mg; OIG: 67.4 ± 4.4 mg
maximal strength: YCG:58.3 ± 2.8 mN; YIG
115.4 ± 5.9 mN
OCG:57.4 ± 3.1 mN; OIG:112.1 ± 6.1 mN
Alway [4] N = 18, 12 in intervention group, 6 in ALD Unilateral stretching with 12% of body MM: + 162.5% ± 3.4%
control group weight FN: + 48.4% ± 3.2%
relative maximal strength 23.6 ± 0.9 mN vs.
18.9 ± 0.6 mN
absolute maximal strength 95% increase vs.
control muscle
Antonio et al. [10] N = 26 ALD Intermittent stretching protocol with Maximal values
progressive weight increase, followed by MM: + 318% ± 31.5%
continuous stretching at 35% of own body FN: + 82.2% ± 17.1%
weight MCSA: + 141.6% ± 32.5%
Intervention period 37 days
Antonio and Gonyea [8] N=7 ALD Stretching with 10% of bodyweight; inter- MM: + 53.1% ± 9%
mittent stretching protocol FCSA: + 27.8% ± 6%
FL: + 26.1% ± 7.3%
Antonio and Gonyea [9] N = 18 ALD 28-day stretching intervention with 29% MM: day16: + 188.1% ± 15.6%;
of bodyweight, animals examined after day28: + 294.3% ± 39.1%
16 days and 28 days FL: day16: + 80.4% ± 11.8%.
day28: + 74.6% ± 9.7%
FN: day16: − 6.7% ± 4.6%;
day28: + 29.7% ± 6.8%
Barnett et al. [14] N = 63 PAT, Unilateral stretching for up to 10 days, ani- MM:
biceps brachii mals examined after 1, 2, 3, 7 and 10 days PAT: CG: 0.1474 ± 0.0142 g IG:
0.2461 ± 0.0239 g
Biceps brachii: CG: 0.5914 ± 0.0607
IG:0.7644 ± 0.0646
Brown et al. [18] N = 40 PAT 16-day stretching intervention, animals Muscle mass increased for 61% in 6-week-old
were examined after 6 days and 16 days chicken and 34% in l0-month-old chicken.
28-month-old animals’had an 18% loss of
muscle mass during passive stretch
Journal of Science in Sport and Exercise
 Table 1  (continued)
Source Subjects Muscle Group Intervention Measured Parameters

Carson et al. [20] N = 94, YA and OA ALD 30-day stretching intervention with 10% of MM: YA: 7d: 94.1% ± 7.4%; 14d:
bodyweight, animals were examined after 134.7% ± 5.8%
7, 14 and 30 days in both ages OA: 7d: 82.1% ± 4.9%; 14d:102.4% ± 6.5%
FL: YA: 7d: 37.7% ± 2.0%; 14d:
28.9% ± 4.0%
OA: 7d: 39.8% ± 4.1%; 14d: 21.3% ± 5.3%
FN: YA: 14d: 31.6% ± 2.1%; OA:14d:
19.2% ± 2.2%
FCSA: YA:14d: 51.6% ± 7%; OA:14d:
Journal of Science in Sport and Exercise

39.6% ± 8.5%
Carson et al. [20] N = 32, YA n = 16 vs. OA n = 16 ALD Unilateral stretching with 10% of body- MM: YA: + 178.7% ± 7.1%
weight, contralateral muscle was control OA: + 142.8% ± 7.9%
muscle FN:
YA: IG: 22.5 ± 0.4 vs. CG: 18.5 ± 0.4
OA: IG: 22.8 ± 1.2 vs. CG:18.4 ± 0.9
MCSA: YA: + 63.8% ± 7.8%;
OA: + 49.1% ± 5.4%
FN: YA: + 59.6% ± 8%; OA: + 47.2% ± 8.1%
Carson and Alway [19] N = 30, YA n = 15 vs. OA n = 15 ALD Unilateral stretching for 7 and 14 days MM: YA;7d: + 98.7% ± 12%
YA;14d: + 141.4% ± 9.5%
OA;7d: + 83.9% ± 6.6%
OA;14d: + 106.9% ± 11%
Czerwinski et al. [23] N = 57, chicken PAT 11-day intervention, stretched muscle vs. MM: CG: 1.3 ± 0.07 g vs. IG: 1.88 ± 0.09 g
control muscle, banded stretch for one
wing
Frankeny et al. [25] N = 54 PAT 6 week stretching intervention with several MM: 24 h: + 121%
stretching protocols, 8, 4, 2 + 2, 2, 1, 0.5 MCSA: up to + 111%
und 0.25 + 0.25 h of intermittent stretch- FCSA: up to + 110%
ing and 24 h of permanent stretching
Matthews et al. [37] N = 10 PAT 33-day stretching intervention with 10% of MM: + 247% ± 91%
bodyweight FCSA: IG: 985 ± 291 µm2 CG: 520 ± 96 µm2
Roman and Alway 1995 [47] N = 28 ALD 21 days stretching intervention, animals MM: 7 days: CG:37.2 ± 1.8 mg IG:
examined after 7, 14 and 21 days 54.6 ± 2.9 mg
14 days: CG:43.5 ± 2.7 mg IG:67.8 ± 4.3 mg
21 days: CG:42.6 ± 3.2 mg IG:71.2 ± 3.7 mg
Sparrow [56] N = 60 ALD 30-day stretching intervention, 30 animals MM: CG:0.928 ± 0.026 g; IG:1.850 ± 0.07 g
examined after 3, 7, 13 and 29 days,
remaining animals examined after 5, 13,
25 and 35 days after stretching without
intervention to investigate regression

ALD anterior latissimus dorsi muscle, PAT patagialis muscle, MM muscle mass, MCSA muscle cross-sectional area, FCSA fiber cross sectional area, FL fiber length, FN fiber number, YA young
animals, OA old animals, YCG​young control group, YIG young intervention group, OCG old control group, OIG old intervention group

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Table 2  Quality Assessment using the Delphi List

Study Rand- Treatment Groups were Eligibility Blinding of Blinding Blinding of Point Esti- Intention to
omiza- Allocation Similar at Criteria Outcome of Care Patient? mates and Treat Analysis
tion? Concealed? Baseline? Specified? Assessor? Provider? Measures of Included?
Variability
Presented?

Czerwinski Y Y Y DN DN Y Y
et al. [23]
Alway et al. DN Y Y DN DN Y Y
[7]
Alway et al. DN Y Y DN DN Y Y
[2]
Alway [3] DN DN Y DN DN Y Y
Alway [4] DN Y Y DN DN Y Y
Antonio DN Y Y DN DN Y Y
et al. [10]
Antonio and DN Y Y DN DN Y Y
Gonyea [8]
Antonio and DN DN Y DN DN Y Y
Gonyea [9]
Barnett et al. DN DN Y DN DN Y Y
[14]
Brown et al. DN Y Y Dn DN Y Y
[18]
Carson et al. DN Y Y DN DN Y Y
[20]
Carson et al. DN Y Y DN DN Y Y
[20]
Carson and DN Y Y DN DN Y Y
Alway [19]
Frankeny DN Y Y DN DN Y Y
et al. [25]
Matthews DN Y Y DN DN Y Y
et al. [37]
Roman and DN DN Y DN DN Y Y
Alway [47]
Sparrow [56] DN Y Y DN DN Y Y

For “treatment allocation concealed?” and “blinding of patient” an assessment was not possible

the SMD was calculated for the comparison of the post- over up to 6 weeks [25]. There were positive effects found
treatment experimental and a respective control group. on muscle mass in most studies, expect for one interven-
tion performed by Brown et al. [18], see Table 3. The
highest increases in muscle mass in the listed studies
Results were obtained by Antonio and Gonyea [8] with a 37-day
stretching intervention and an increase of 318% ± 39.1%
Muscle Mass and d = 7.01, 95% CI 3.77–10.24. Other high percentage
increases were obtained by Antonio and Gonyea [9] with
The included studies show that in animal experiments a an increase of 294.3% ± 39.1% with d = 11.96, 95% CI
significant increase in muscle mass can be achieved by 7.27–16.66 in muscle mass, Alway [2] with an increase of
stretching intervention over several weeks. The effect 161.5% ± 7.9% with d = 6.64, 95% CI 5.43–7.85, and Car-
size across all studies was d = 8.51, P < 0.001, 95% CI son et al. [20, 21] with 178.7% ± 7.1% d = 20.82, 95% CI
7.11–9.91. Stretching was performed with varying dura- 15.44–26.32.
tions per day (minimum 2 × 15 min) up to 24 h stretching

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Table 3  Forest plot for muscle mass

Table 4  Forest plot for muscle cross-sectional area

Muscle Cross‑Sectional Area the control muscle. Alway [3] also recorded muscle cross-
sectional increases of 100% (see Table 4).
Changes in muscle cross-sectional were all positive.
Here, an effect strength of d = 7.91, P < 0.001, 95% CI Fiber Cross‑Sectional Area
5.75–10.08 was recorded. Frankeny et al. [25] measured
an increase in muscle cross-section of 111% compared to For the effects on fiber hypertrophy, an increase due to
the stretching intervention was also determined (almost)

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Table 5  Forest plot for fiber cross-sectional area

Table 6  Forest plot for fiber number

consistently. The effect size here was d = 5.81, P < 0.001, the number of fibers was also recorded in this study (see
95% CI 4.32–7.31. The changes in fiber cross-section Table 6).
ranged from − 0.75% to 141.6% (± 32.6%), with these two
values being more of an outlier, as all other results ranged Fiber Length
from + 27.8% to + 63.8% (see Table 5).
The fiber length was only taken into account in three stud-
Fiber Number (Hyperplasia) ies. The effect size determined was d = 7.86, P < 0.001,
95% CI 4.00–11.72. Here, percentage increases were
With regard to the number of fibers, the studies also show 26.1% ± 7.3% (d = 3.31, 95% CI 1.52–5.09 [8]. Studies by
significant increases as an adaptation to permanent stretch- Alway [3] determined muscle length changes of approx.
ing. Here, the calculated effect size across the studies is 25% compared to the control muscle due to the stretching
d = 4.62, P < 0.001, 2.54–6.71. In two studies, a decrease intervention (see Table 7).
in the number of fibers − 0.7% ± 3.6% with d = − 0.29,
95% CI − 1.34–0.77 in Antonio and Gonyea [8] and
− 6.7% ± 4.6% with d = − 1.3, 95% CI − 2.6–0.0 in Anto- Discussion
nio and Gonyea [9] was initially determined after a certain
intervention period, which, however, was no longer pre- Based on the studies and the effect sizes determined in this
sent at a later test in the same study, so that an increase in meta-analysis, it can be assumed that (continuous) stretch-
ing (from 30 min to 24 h per day in a longitudinal section

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Table 7  Forest plot for fiber length

over several days to weeks) induces muscular tension in effects achieved by stretching, such that the mechanical
animal muscles, which leads to the following morphologi- stimulus on the muscle is the adaptation-inducing stressor
cal adaptations of the stretched muscles: an increase in and thus the crucial stimulus for muscle mass gains [49,
muscle mass, muscle cross-section, fiber cross-section, 67]. The muscle responds to this stimulus by increasing
fiber length, and/or number of muscle fibers. This is con- its serial sarcomere number [66] and the accumulation of
firmed by the results of other studies whose experimental myofibrils triggers an increase in cross-sectional area [4,
investigations were similar to the analyzed studies, but 8, 20, 25]. The increase in muscle mass due to long dura-
which could not be included in the statistical analyses due tion stretching interventions has been clearly demonstrated
to exclusion criteria or missing information in the method in animal studies. Various studies with animals have also
description [6, 12, 15, 35, 69]. demonstrated an increased rate of protein synthesis by
Several studies show that there seems to be a correla- stretching [16, 28, 29]. Whether and to what extent the
tion between stretching time and stretching intensity with results of this study are transferable to humans have not
achieved muscle mass increase [15, 25, 35], assuming an yet been adequately investigated. Several of the studies
upper limit or optimum of stretching duration. In studies integrated in this meta-analysis specifically request this
by Frankeny et al. [25] and Bates [15], although further step [15, 29]. Critically, protein synthesis differs between
increases due to an increase in stretching duration can be humans and animals. Garibotto et al. [27] and Tessari et al.
detected, the stretching optimum (effort relative to return) [58] list protein synthesis rates of 2% and 1.5%, respec-
seems to be 30 min: “We conclude that daily stretching for tively, for leg muscles. Early experiments made by Wil-
as little as 30 min per day is a powerful inducer of growth liams and Goldspink indicate 2–3 days for length adapta-
in normal and dystrophic muscle” [25]. Antonio et al. tion of muscle in mice, but 2–3 weeks in cats and humans
[10] achieved maximal muscle mass gains of 318% with [67]. For the species primarily studied in this meta-anal-
a progressively increased stretching load and an intermit- ysis (chickens/quail), Sayegh and Lajtha [50] indicate a
tent stretching protocol. The increases in muscle mass is lower protein synthesis rate compared to mice. However,
consistent in almost all studies listed in this meta-analysis the protein synthesis rate is dependent on the species, but
except for one measured parameter by Brown et al. [18] also on other factors such as gender or hormones (e.g.,
due to stretching the PAT for 16 days in old female chicken testosterone) [60], age, and muscle fiber distribution or
(28 month old). the expression of myosin heavy chains [42, 52]. The high-
The muscle mass gains are attributed by most authors to est increases in muscle length reported in the literature
muscle fiber hypertrophy and muscle fiber hyperplasia. For were found to be up to 60% depending on the duration
muscle hyperplasia, uninterrupted continuous stretching of stretching by Antonio and Gonyea [8] or up to 77%
seems to be the initiating stimulus, since the muscle fiber by Antonio et al. [9].
is not given sufficient time to regenerate. This stimulates With regard to fiber hypertrophy in animal experiments,
increased satellite cell activation, which leads to the for- no uniform statement can be made. Antonio et al. [10] found
mation of new muscle fibers [8]. Another explanation is an increase in the cross-section of FT as well as ST fib-
that reaching a critical muscle fiber size by hypertrophy ers, whereas Alway, [3] and Roman and Alway [47], for
effects leads to the splicing of the muscle fiber into several example, do not highlight any increase in the muscle cross-
muscle fibers. This could be responsible for hyperplasia section of FT fibers. The hypertrophy of ST fibers seems to
[8, 10]. be regulated by the calcineurin/NFAT signal transduction
pathway [48]. This is significant as the studies listed in this
meta-analysis are primarily concerned with prolonged exer-
Hypertrophy cise leading to ST fiber adaptations [10, 20, 21, 25, 29, 30].

Induced tension or mechanical stress on the individual sar-

comeres are thought to be responsible for the hypertrophy

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Hyperplasia explain all of the 60% decline in shortening velocity, unless

ATPase activity also declined in SM1 or slow-p fibers. Our
Referring to the finding of Antonio et al. [10] that the preliminary data suggest that ­Ca2+ activated ATPase activity
amount of increase in fiber count is related to the duration was − 20% lower in the SM2 isoform than the SM1 isoform,
as well as the amount of the stretching stimulus, it can be and ATPase activity decreased in both isoforms after stretch
hypothesized that traditional strength training methods do overload” [4]. If these results are transferable to humans, it
not achieve adequate stretching of the muscles. A stretching can be assumed that an increase in the ST-fiber content and
intervention lasting several hours to several days, as per- thus a reduction in high-speed power output (e.g. jumps,
formed in animal experiments, has of course not been car- sprints) is due to muscle plasticity and a reduced ATPase
ried out. The proliferation and activation of satellite cells is activity.
held responsible for the hyperplasia effects [8, 10, 29]. This For the investigated parameters MM, FCSA and FN, het-
seems to occur—at least in animal experiments—when a erogeneity was relatively large (I2 > 90%), suggesting that
muscle is seriously damaged by mechanical stress [36, 57]. moderator variables could explain some of the differences
between the true effect sizes of the included individual stud-
Maximum Strength and Speed Strength ies. The forest plots for MCSA, FCSA, FN and FL provide
graphical information of which effect sizes differ the most
Studies by Alway [3, 4] found a significant increase in mus- from the weighted averages, but systematic subgroup analy-
cle cross-sectional area (approximately 100%), in muscle ses where studies are grouped with respect to moderators,
mass (260%) and in maximum strength (95%) in animal such as muscle group or fiber distribution within the muscle,
muscles. Stretching the muscle can be assumed to lengthen gender of the test animal, age of the animal or stretching
the muscle fiber through serial accumulation of sarcomeres duration, does not seem feasible due to the (still) relatively
[4, 10]. In animal experiments, muscle lengthening of up to small number of effect sizes. Using only birds as experimen-
60% depending on the stretch duration was found by Antonio tal animals and including ALD and PAT in the analysis of
and Gonyea [8] or up to 77% by Antonio and Gonyea [9]. this meta-analysis, we already tried to account for potential
According to Goldspink and Harridge [29], this can lead to a heterogeneity by controlling these variables in the selection
faster contractile capacity of the muscle and thus an increase of studies (in contrast to Kelley [33].
in fast or explosive power capacity. This hypothesis is con-
firmed by Medeiros and Lima [39] who identified 14 studies Practical Implications
with a positive influence on “muscle performance” through
chronic stretching. Muscle performance was recorded in Although the results from animal experiments presented
the studies by functional tests such as jumps or sprints or here are conclusive, they may not be directly transferable to
by isometric or isotonic contractions. This is contradicted humans. First evidence that stretching training can induce
by data on the change in myosin heavy chain expression in micro-traumatization in humans if appropriate intensity of
stretched muscle as demonstrated in the animal experiment the stimulus is given was provided by Smith et al. [55]. Sch-
by Roman and Alway [47]. Myosin isoform SM2 increased oenfeld [51, p. 2862] also refers to the possibility to induce
from a level of 43.1% (± 1.7%) in the control muscle to 55% sufficient mechanical tension to induce morphological adap-
(± 1.2%) in the stretched muscle. It was shown that sustained tations using stretching training: "Mechanically induced ten-
stretching resulted in increased expression of SH2 myosin sion produced both by force generation and stretch is con-
heavy chains and decreased expression of SH1 myosin heavy sidered essential to muscle growth, and the combination of
chains. Thus, due to the decreased ATPase activity in hyper- these stimuli appears to have a pronounced additive effect”.
trophied type I fibers after stretching, a negative effect on Consequently, there are some studies pointing out improve-
muscle contraction speed can be assumed, which was con- ments in sport-specific parameters as jumping and sprinting
firmed by Alway [2]. [34, 44], maximal strength [41, 43, 70] and muscle thick-
Contraction time increased significantly from 149 ms ness [44, 53] using stretching durations of up to 6 × 5 min
(± 9 ms) to 162 ms (± 7 ms) in young animals and from [70] for up to 12 weeks [44]. However, there is still a lack
174 ms (± 16 ms) to 215 ms (± 14 ms) in old animals by of human studies on the effects of long-lasting stretching
continuous stretching with 12% of their own body weight. interventions for many weeks on muscular hypertrophy,
“Overload increased twitch contraction time by 36% in mus- hyperplasia, and force development. Because frequency,
cles from … birds” [5]. There was a measurable shift from magnitude, and especially intensity of stretching appear to
SM1 myosin isoform to SM2 myosin isoform. “Neverthe- play an important role in adaptive responses, further studies
less, the slowing of V, and Vmax in the ALD was related to need to focus on load controls via these load normatives.
the decrease in SMl and slow muscle fibers. The explana- Apostolopoulos et al. [11] hypothesized that below the pain
tion for a shift in fiber type or myosin isoforms is unable to threshold stretches in the muscle are compensated via the

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Journal of Science in Sport and Exercise

elastic components and only stretches above the pain thresh- not possible. This is different in studies using human par-
old lead to inflammation, which is normal after a fatiguing ticipants. With regard to the conducted quality assessment,
load [32] and/or delayed onset muscle soreness. In addition an important limitation appears to be the fact that in most
to intensity, a minimum amount and duration of stretching is studies, the assessors (of the outcome parameters) were not
essential, as Fowles et al. [24] showed that a single bout of blinded with regard to which animals were assigned to the
stretching does not seem to be sufficient to increase protein experimental or control group. Also, visual inspection of
synthesis. In accordance, Freitas et al. [26] pointed out that the funnel plots performed for each outcome parameter sug-
interventions of less than 8 weeks with a stretching duration gested slight deviations from a symmetric distribution in
of less than 20 min per week would not be expected to pro- some cases. However, this could be due to the rather small
duce statistically significant structural changes in humans. effect sizes and should be interpreted with caution. Further-
Therefore, stretching duration may play an important role, more, also due to the rather small number of studies, it was
too. Only one study using daily long-lasting stretching train- not possible to reliably investigate the potential influence
ing for the plantar flexors could be determined, showing sig- of moderator variables, such as duration of stretching, for
nificant increases in maximal strength, muscle thickness and instance. Finally, it needs to be highlighted that most studies
flexibility [64]. Since in animal studies, apparatuses were were performed about 30–40 years ago.
used to achieve long-lasting stretching durations, stretch-
Funding Open Access funding enabled and organized by Projekt
ing devices (as used by Warneke et al. [64]) could also be DEAL.
recommended to achieve long-lasting stretching durations
in humans. Otherwise, it can be assumed that stretching Data Availability The datasets generated and analyzed during the cur-
durations lasting several hours are not feasible. If a certain rent study are available from the corresponding author upon request.
degree of transferability to humans is assumed, the studies
analyzed here can be seen to have particular relevance in
Declarations
rehabilitation [29], as immobilization due to injury is known Conflict of Interest On behalf of all authors, the corresponding author
to lead to significant muscle atrophy [45]. If the hypertro- states that there is no conflict of interest.
phy effects from animal studies are assumed to be transfer-
able to humans, aid-based continuous stretching for several Consent for publication All authors have read and agreed to the pub-
lished version of the manuscript.
hours could counteract atrophy and, if necessary, support
muscle mass gain. “The therapeutic applications of stretch Open Access This article is licensed under a Creative Commons Attri-
should therefore be borne in mind when designing regimens bution 4.0 International License, which permits use, sharing, adapta-
for rehabilitation or improved athletic performance” [29]. tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
Furthermore, if voluntary muscle activation is not possible, provide a link to the Creative Commons licence, and indicate if changes
stretching intervention would already be applicable. This were made. The images or other third party material in this article are
could minimize muscle atrophy and loss of strength through included in the article's Creative Commons licence, unless indicated
immobilization due to injuries or illnesses [65, 68]. otherwise in a credit line to the material. If material is not included in
the article's Creative Commons licence and your intended use is not
For an examination of the results in humans, moderator permitted by statutory regulation or exceeds the permitted use, you will
variables should be taken into account to be able to examine need to obtain permission directly from the copyright holder. To view a
their influence. copy of this licence, visit http://​creat​iveco​mmons.​org/​licen​ses/​by/4.​0/.
If transferability of our results to humans is given, we see
a high potential in using long-lasting stretching to achieve
muscle hypertrophy. But it remains controversial whether
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