Physiology of Stretch0Mediated Hypertrophy and Strength Increases

Sports Medicine (2023) 53:2055–2075
https://doi.org/10.1007/s40279-023-01898-x
REVIEW ARTICLE
Physiology of Stretch‑Mediated Hypertrophy and Strength Increases:

A Narrative Review
Konstantin Warneke1,10,11 · Lars H. Lohmann2 · Camila D. Lima3 · Karsten Hollander4 · Andreas Konrad5,10 ·
Astrid Zech6 · Masatoshi Nakamura7 · Klaus Wirth8 · Michael Keiner9 · David G. Behm10
Accepted: 22 July 2023 / Published online: 9 August 2023

© The Author(s) 2023
Abstract
Increasing muscle strength and cross-sectional area is of crucial importance to improve or maintain physical function in musculo-
skeletal rehabilitation and sports performance. Decreases in muscular performance are experienced in phases of reduced physical
activity or immobilization. These decrements highlight the need for alternative, easily accessible training regimens for a sedentary
population to improve rehabilitation and injury prevention routines. Commonly, muscle hypertrophy and strength increases are
associated with resistance training, typically performed in a training facility. Mechanical tension, which is usually induced with
resistance machines and devices, is known to be an important factor that stimulates the underlying signaling pathways to enhance
protein synthesis. Findings from animal studies suggest an alternative means to induce mechanical tension to enhance protein
synthesis, and therefore muscle hypertrophy by inducing high-volume stretching. Thus, this narrative review discusses mechani-
cal tension-induced physiological adaptations and their impact on muscle hypertrophy and strength gains. Furthermore, research
addressing stretch-induced hypertrophy is critically analyzed. Derived from animal research, the stretching literature exploring
the impact of static stretching on morphological and functional adaptations was reviewed and critically discussed. No studies have
investigated the underlying physiological mechanisms in humans yet, and thus the underlying mechanisms remain speculative and
must be discussed in the light of animal research. However, studies that reported functional and morphological increases in humans
commonly used stretching durations of > 30 min per session of the plantar flexors, indicating the importance of high stretching
volume, if the aim is to increase muscle mass and maximum strength. Therefore, the practical applicability seems limited to set-
tings without access to resistance training (e.g., in an immobilized state at the start of rehabilitation), as resistance training seems
to be more time efficient. Nevertheless, further research is needed to generate evidence in different human populations (athletes,
sedentary individuals, and rehabilitation patients) and to quantify stretching intensity.
thickness [11, 15, 16]. However, force output also depends

1 Introduction on neuromuscular factors such as recruitment, rate coding,
and synchronization of related motor neurons [12, 14, 17].
In rehabilitation [1, 2] and sports practice [3–6], improv- Based on these mechanisms, increased maximal strength
ing maximal strength and muscle mass is well correlated induced by resistance training can be assumed to be the
with joint function and stability as well as sport-specific result of a wide range of neuromuscular and structural adap-
performance. Most commonly, resistance training is used tations in response to specific training stimuli.
to increase muscle strength capacity and induce significant Periods of immobilization or lower physical activity after
hypertrophy [7–11]. In the literature, Goldspink and Har- serious injury [2, 18, 19] or pandemic lockdowns [20–23] as
ridge [12] refer to muscle force as a reflection of myofila- well as aging in general [24, 25] result in significant loss of
ment cross-bridges working in parallel, suggesting a link muscular performance including atrophy, decrease in maxi-
between fiber cross-sectional area and muscle force [12]. mal strength, loss of flexibility as well as musculoskeletal
Accordingly, increases in maximal strength via resistance pain [26]. Morie et al. [27] and Hotta et al. [28] point out
training are often accompanied by hypertrophy [13, 14] the need for safe and efficient alternatives to commonly used
due to enhanced muscle cross-sectional area and/or muscle training and current therapies in rehabilitation settings to
prevent loss of muscle performance and to restore maxi-
mal strength and flexibility. While different types of stretch
Extended author information available on the last page of the article training are known to improve flexibility [29–33], muscle
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2056 K. Warneke et al.
hypertrophy of muscular tissue that counteracts performance

Key Points and muscle mass losses in rehabilitative settings. However,
as no studies were detected exploring the underlying physi-
Stretching is known to improve flexibility, while resist- ological mechanisms of stretch-induced muscle tension and
ance training is commonly used to increase strength its impact on muscle hypertrophy, this review discusses dis-
capacity and hypertrophy. crepancies found in the literature in the light of mechanical
Both training methods produce mechanical tension, tension. Furthermore, potential factors contributing to the
which is a factor known to stimulate anabolic signaling described heterogeneity in results will be highlighted.
and enhance protein synthesis.
In this review, results extracted from the current lit- 2 Impact of Mechanical Tension
erature considering physiological aspects indicate the on Physiological Muscle Adaptations:
possibility of using high-volume stretching durations From the Genotype to the Phenotype
to enhance muscle volume in humans. However, the
findings considered are biased by the different methods “Form follows function” is a well-known concept propagated
applied in the literature, particularly in regard to stretch- by the architect Louis H. Sullivan, which seems applicable to
ing intensity and duration. many biological adaptations: specific suprathreshold stimuli
cause specific adaptations in the human body. Following the
hypertrophy and strength increases are mainly associated response matrix model developed by Toigo and Boutellier
with resistance training [15, 34, 35]. However, recent litera- [51], anabolic responses can be traced back to a specific mix-
ture demonstrated that, on the one hand, resistance training ture of transcription and translation factors, determined by
performed over the full range of motion seems to provide the type of external stimuli [46, 52, 53]. Inducing mechani-
a sufficient stimulus to enhance flexibility [36, 37], while cal tension is one of the most ubiquitous external stimuli,
on the other hand, animal studies reported hypertrophy and specifically known from resistance training and seemingly
strength increases by attaching weight/resistance to one of crucial importance for adaptations of muscular tissue [13,
wing of a quail or using a stretching apparatus [34, 35] to 46]. When performing resistance training, mechanical ten-
apply (weighted) stretch to the muscles. Therefore, consider- sion is increased by enhancing the intensity, for example by
ing similar effects regarding the main outcomes of strength, adding weight to a barbell, leading to enhanced mechanical
muscle mass, and flexibility, shared underlying mechanisms stress-induced micro-traumatization and exercise-induced
between stretching and strength training could be hypoth- muscle damage [54–56]. To induce morphological changes,
esized. In humans, there is conflicting evidence regarding on a physiological level, gene expression is controlled via
stretch-induced muscle hypertrophy and strength gains. external stimuli such as suprathreshold mechanical ten-
Some recently published studies support a potential transfer- sion. By applying specific stress, specific signaling path-
ability to human models confirming stretch-mediated hyper- ways are enhanced, transducing external physical stimuli
trophy and maximal strength increases in humans following (such as mechanical overload) through biochemical signals
long-lasting stretching for up to 2 h per session using calf to control the corresponding net protein synthesis rate by
muscle orthoses or stretching boards [38–41]. These results causing either an anabolic or catabolic milieu [12, 57–59].
are opposed by findings showing neither maximal strength Accordingly, Coffey and Hawley [60] described that “[t]he
increases [42–44], nor hypertrophy [42, 45]. Therefore, the process of converting a mechanical signal generated dur-
question arises about the underlying mechanisms and the ing contraction to a molecular event that promotes adapta-
influence of training modalities such as stretching duration tion in a muscle cell involves the upregulation of primary
or intensity, which could be responsible for the similarities and secondary messengers that initiate a cascade of events
in adaptations between (full range of motion) strength train- that result in activation and/or repression of specific signal-
ing and (long-duration) static stretching training on muscle ing pathways regulating exercise-induced gene expression
hypertrophy. and protein synthesis/degradation”, which is considered of
In strength training, mechanical tension is known to play paramount importance when developing new tissue [61, 62].
a key role in morphological and functional adaptations, It has been shown that several signaling pathways, such as
such as hypertrophy and maximal strength [14, 46, 47], and the mammalian target of rapamycin/ribosomal protein/S6
has also been considered to be one of the most ubiquitous kinases/phosphoinositide 3-kinases (mTOR/p70s6K/PI3K)
explanatory processes contributing to stretch-mediated pathway stimulate anabolic responses for muscle protein
hypertrophy in animals [48–50]. Hypothesizing a potential synthesis [46, 47, 61, 63, 64] with the highly discussed influ-
transferability to humans opens up different opportunities ence of p70S6K in training-induced hypertrophy [65–68].
to passively apply sufficient mechanical tension to induce Based on this pathway, mechanical overload seems to be
Physiology of Stretch-Mediated Hypertrophy and Strength Increases 2057
responsible for the release of insulin-like growth factor-1 chicken and quail [48–50, 79, 80]. The results are provided
(IGF-1), a key factor in muscle growth [58, 69]. In papers and discussed in the following section and Table 1.
by Tidall [70] and Toigo and Boutellier [51], activation of
protein kinase B is described as a response to mechanical
overload, which activates the v-Akt murine thymoma viral 3 Stretch‑Mediated Hypertrophy: Results
oncogene/protein kinase-B/mammalian target of rapamycin from Animal Studies
pathway with a downstream phosphorylation of p70S6K [71,
72] contributing to anabolic responses mainly in two ways: If the aim is to answer an important research question, Hooi-
firstly, catabolic or anti-anabolic pathways such as glycogen jmans et al. [81] suggested investigating underlying mech-
synthase kinase 3 are reported to be inhibited. Secondly, anisms in animal studies first. From 1970 to 2000, a vast
further growth factors such as myogenin growth factor bind- number of studies using animal models aimed to explore
ing to IGF-related protein (IGFR) activate anabolic kinases underlying mechanisms of chronic stretch-induced muscle
by phosphorylation of insulin receptor substrate 1. Hence, mass increases [77, 82–85]. For that purpose, chickens or
a highly complex system of signaling pathways influencing quails typically had the muscle of one wing stretched by
the homeostasis between muscle protein synthesis and deg- applying a stretching device [79, 80, 86] or adding weight
radation is described for resistance training-induced muscle equivalent to 10–35% of the respective bodyweight to the
hypertrophy [12, 55–57]. wing [77, 87–89]. Most of the studies examined the influ-
The literature points out alternative ways to induce ence of different stretching times that ranged from 2 × 15 min
mechanical tension and therefore mechanical overload on per day (using an intermittent stretching protocol) [86, 90] to
muscular tissue. As early as 1887, Marey [73] demonstrated 24 h of stretching per day [50, 82, 90–95] on muscle mass,
the adaptability of muscle tissue in animals by surgically muscle cross-sectional area, fiber cross-sectional area, fiber
moving the triceps surae muscle insertion on the calcaneum length and/or fiber number. Some studies investigated phys-
farther from the origin leading to an elongation by increas- iological adaptations and changes in gene expression and
ing the distance between the muscle origin and insertion and muscle protein synthesis, myosin isoforms [83, 91, 96] as
consequently a stretch, which is reported to result in a signif- well as myosin heavy chains and myosin light chains [83, 92,
icant increase in the serial sarcomere number [74]. Hence, it 97]. To include a control condition, the contralateral non-
can be hypothesized that the increased muscle length caused stretched muscle was used as an intra-individual control
a chronic stretch of the muscle. Accordingly, Alder et al. [75] condition [82, 87, 91, 92]. Thus, increases in the intervened
reported that stretching a muscle increased muscle length, muscle were reported under the assumption of no change in
whereas, in contrast, immobilizing a muscle in a shortened the control muscle of the contralateral side.
position decreased the serial sarcomere number [75, 76]. As Most studies investigated the influence of prolonged
the influence of modified innervation patterns (contracting stretching interventions in the anterior latissimus dorsi,
against an attached immobilization device) was discussed in because of its high percentage of slow-twitch fibers, assum-
studies investigating the influence of stretching on muscle ing higher responses to chronic stimulation [98]. In 1973,
morphology, Sola et al. [77] applied chronic stretch to den- Sola et al. [77] used 100-g and 200-g weights attached to
ervated bird muscle to exclude the impact of central nerv- the wing of chickens to induce a stretch stimulus to the ante-
ous innervation and was consequently able to attribute the rior latissimus dorsi, posterior latissimus dorsi, and teres
measured hypertrophy effects to passive mechanical tension. minor of one wing leading to an increase in muscle weight
In addition to known sarcomere accumulation in series, the of up to 169%. Other authors investigated the influence of
authors reported significant hypertrophy effects through par- mechanical overload via stretching in a flight muscle (pata-
allel accumulation of myofibrils. Even though the healthy gialis muscle). Muscle mass results for applicable studies
comparison group showed (slightly) higher hypertrophic are provided in Table 1.
effects, the authors reported significant muscle growth in The results in Table 1 show progressive muscle mass
the denervated muscle. Consequently, the authors hypoth- growth over time with increases of 53.1% after 7 days to
esized that skeletal muscle stretch would induce sufficient 318.6% after 5 weeks, using daily stretching [87]. In their
mechanical tension as the underlying assumed mechanism meta-analysis, Warneke et al. [49] reported increases in ani-
to induce hypertrophy in parallel [54, 58, 69, 78]. In the mal fiber cross-sectional area of up to 141.6% [89, 93, 94,
following years, stretch-mediated hypertrophy was exten- 99] and an enhanced fiber number of up to 82.2% [82, 84,
sively researched by applying chronic stretching (24 h per 87, 89, 93, 94]. These adaptations were accompanied by
day 7 days per week for up to 35 weeks) to one wing of fiber length increases of up to 50% [87, 89, 99] in chicken
Table 1 Characteristics of longitudinal animal studies investigating stretch-induced increase in muscle mass in chickens and quails
2058
Study Animals Intervention period (Daily) stretching time and Weight added to stretch or Results (muscle mass increase)
stretch application frequency stretching apparatus
Alway [88] Japanese quail (n = 20) 30 days 24 h per day, every day 12% of body weight 161.5 ± 7.9%
Alway [99] Japanese quail (n = 70) 30 days 24 h per day, every day 12% of body weight Young: 45.1 ± 2.1%, aged:
24.1 ± 2.6%
Alway [92] Japanese quail (n = 24) 30 days 24 h per day, every day 12% of body weight Young: 168.2%, aged: 136.5%
Alway [91] Japanese quail (n = 12) 30 days 24 h per day, every day 12% of body weight 162.5 ± 4.3%
Alway et al. [100] Japanese quail (n = 73) 1–7 days 24 h per day, every day 10% of body weight 6.9 ± 2.7% after 1 day,
21.3 ± 4.7% after 2 days,
41.7 ± 4.7 after 3 days,
44.1 ± 4.4% after 4 days,
59.8 ± 6.5% after 5 days,
58.9 ± 9.6 after 6 days and
64 ± 8.4% after 7 days
Muscle mass difference at
baseline: 0.5 ± 0.02%
Alway et al. [89] Japanese quail (n = 34) 30 days 24 h per day, every day 10% of body weight 171.8 ± 13.5%
Antonio and Gonyea [101] Japanese quail (n = 18) 16 and 28 days (intermittent 24 h of stretching on day 1 Progressing from 10 to 35% 188.1 ± 15.6% after 16 days and
stretching protocol) with 10%, day 4 with 15%, of body weight 294.3 ± 39.1% after 28 days
day 8 with 20%, days 11–14
with 25% and days 17–37
with 35% of bodyweight.
The days in-between
included no intervention
(rest)
Antonio and Gonyea [50] Japanese quail (n = 26) 28 days (intermittent stretch- 24h of stretching on day 1 Progressing from 10 to 35% 318.6 ± 31.5%
ing protocol) with 10%, day 4 with 15%, of body weight
day 8 with 20%, days 11–14
with 25% and days 17–37
with 35% of bodyweight.
The days in-between
(rest)
Antonio and Gonyea [82] Japanese quail (n = 7) 15 days (intermittent stretch- 24 h of stretching on day 1, 10% of body weight 53.1 ± 9%
ing protocol) day 4, day 8, day 11, and
day 15 The days in-between
(rest)
Ashmore [102] Chicken (n = N/A) [dys- 42 days 24 h per day, every day Cardboard sleeve 200%
trophic]
Barnett et al. [79] Chicken (n = N/A) 1, 3, 5, 7, and 10 days 24 h per day, every day Stretch apparatus 16.9% after 1 day, 35.86% after
3 days, 59.48% after 5 days,
59.74% after 7 days, and
67.35% after 10 days
K. Warneke et al.
Table 1 (continued)
Bates [86] Quail (n = 16) 35 days 0.5 h, 1 h, 2 h, 4 h, or 8 h per Cardboard sleeve Daily stretching time of 0.5 h, 1
day, respectively, every day h, 2 h, 4 h, and 8 h
0.5 h = 57%, 1 h = 60%,
2 h = 67%, 4 h = 72%, 8
h = 150%
Brown et al. [103] Chicken (n = 46) 4, 6, 11, and 16 days 24 h per day, every day Wing band Male, 6 weeks old: 66.67%
after 6 days Female, 10
months old: 27.12% after
4 days, 32.95% after 11,
and 43.23% after 16 days
Muscle mass differences at
baseline 1.09% Female, 28
months old: 6.06% after 4
days, 12.64% after 11 days,
and − 17.65% after 16 days.
Muscle mass differences at
baseline: 3.51%
Physiology of Stretch-Mediated Hypertrophy and Strength Increases
Carson and Alway [84] Japanese quail (n = 30) 7 and 14 days 24 h per day, every day 10% of body weight Young: 98.7 ± 12% after 7 days
and 141.6 ± 9.5% after 14
days, aged: 83.9 ± 6.6% after
7 days and 106.9 ± 11% after
14 days
Carson et al. [93] Japanese quail (n = 94) 7 and 14 days 24 h per day, every day 10% of body weight Adult: 94.1 ± 7.4% after 7 days
and 134.7 ± 5.8% after 14
days, aged: 82.1.1 ± 4.9%
after 7 days and 102.4 ± 6.2%
after 14 days. Muscle mass
difference at baseline:
1.8 ± 0.6%
Carson et al. [94] Japanese quail (n = 32) 30 days 24 h per day, every day 10% of body weight Young: 178.7 ± 7.1%, aged:
142.8 ± 7.9%
Czerwinski et al. [104] Chicken (n = 22) 2 and 11 days 24 h per day, every day Cardboard sleeve 13% after 2 days and 44% after
11 days
2059
Table 1 (continued)
2060
Devol et al. [105] Chicken (n = 101) 5, 10, 15, 20, and 25 days 4 h or 24 h per day, respec- Cardboard sleeve Stretching 4 h per day: 8.62%
tively, every day after 5 days, 25.32% after 10
Stretching 24 h per day: 35.51%
after 5 days, 42.76% after 10
Frankeny et al. [90] Chicken (n = 54) 42 days 2 × 0.25 h, 0.5 h, 1 h, 2 h, 2 × 4 Cardboard sleeve Daily stretching time var-
h, 6 h, 8 h or 24 h, respec- ies between 2 × 15 min
tively, every day and 24 h per day. 2 × 0.25
h = 52.9%, 1 × 0.5 h = 70%,
1 × 1 h = 68.8%, 1 × 2
h = 64.41%, 2 × 2 h = 97.83%,
1 × 4 h = 88.46%, 1 × 6
h = 105.77%, 1 × 8 h = 90.45%
and 1 × 24 h = 122.43%
Holly et al. [80] Chicken (n = 14) 35 days 24 h per day, every day Spring-loaded device 63% in PAT muscle, 81% in
ALD muscle, and 28% in
biceps muscle
Kennedy et al. [83] Chicken (n = 32) 35 days 24 h per day, every day 10% of body weight 100% after 35 days
Laurent and Sparrow [106] Fowl gallus domesticus 6 days 24 h per day, every day Attaching a weight (N/A) 74% after 6 days
(n = N/A)
Lee and Alway [95] Quail (n = N/A) 30 days 24 h per day, every day 12% of body weight Young: 44.1 ± 3.1%, adult:
32.6 ± 3.9%, old: 25.7 ± 4.3%
Matthews et al. [96] Japanese quail (n = 10) 33 days 24 h per day, every day 10% of body weight 247 ± 91%
Roman and Alway [97] Japanese quail (n = 28) 7, 14, and 21 days 24 h per day, every day 10% of body weight 53.6 ± 2.9% after 7 days,
67 ± 4.4% after 14 days and
70.2 ± 4% after 21 days
Sola et al. [77] Chicken (n = N/A) 14, 28, and 56 days 24 h per day, every day Attaching a weight of 100 g Peak increase in the ALD with
and 200 g 200-g overload: 158%, 100-g
overload: 93%
Peak increase in the PLD with
200g overload: 29%, 100g
overload: − 4%
Peak increase in the TM with
200-g overload: 169%, 100-g
overload: 64%
Summers et al. [98] Chicken (n = N/A) 5 days 24 h per day, every day Cardboard sleeve Up to 50%
K. Warneke et al.
and quails. While the literature provides extensive evidence

Results (muscle mass increase) for stretch-mediated hypertrophy in birds, there are only a
33.6 ± 7.1% after 5 days and

115.3 ± 8% after 30 days
small number of studies in mammals, showing partially con-
trasting results.
De Jaeger et al. [100] performed a 4-week stretch train-
ing program on rabbit plantar flexors 3 days per week and
found significant muscle mass increases of 13.6%. Stretch-
ing immobilized muscle, Coutinho et al. [101] found
decreases in rat soleus muscle mass following 3 weeks
ALD anterior latissimus dorsi muscle, n number of subjects, N/A not available, PAT patagialis muscle, PLD posterior latissimus dorsi muscle, TM teres minor muscle
of immobilization of one limb and stretching for 40 min

every 3 days during the immobilization phase, while
Weight added to stretch or
stretching alone without immobilization resulted in no sig-

nificant changes. Similarly, Gomes et al. [102] determined
10% of body weight
stretching apparatus
a significant decrease in muscle mass in rat soleus muscle

after 3 weeks of immobilization and stretching for 40 min
once weekly. In a subsequent study by Coutinho et al.
[103], the authors found that daily stretching for 3 weeks
after a prior 4-week immobilization period resulted in a
significant increase in rat soleus muscle mass compared
stretch application frequency
with the contralateral control. The methodology used in

(Daily) stretching time and
these studies is therefore not comparable to the procedure

24 h per day, every day
used in studies including birds, which are listed in Table 1.

Accordingly, lower frequencies and/or stretching durations
per session were used, resulting in less hypertrophy, if any.
Consequently, the impact of modifying stretching param-
eters such as duration, frequency, and intensity is more
extensively debated in Sect. 5. Apart from differences in
load control, a species-dependent difference in adaptations
could also be assumed, raising questions about the trans-
ferability of stretch-mediated hypertrophy from birds to
humans, particularly given presumed differences in the
Intervention period
overall protein synthesis rate [49, 104, 105].

5 and 30 days
4 Stretch‑Mediated Hypertrophy: Results

from Human Studies
Winchester and Gonyea [107] Corturnix quail (n = 27)
As early as 1983, after finding stretch-mediated hypertrophy

in animals, Frankeny et al. [90] and Bates [86] requested
the investigation of transferability of stretch-induced muscle
growth to humans. However, in the only systematic review
Animals
found on the topic [45], no studies with comparable train-

ing parameters could be included. Accordingly, in 2020, the
available literature illustrated a lack of muscle hypertrophy
with stretching durations of up to 4.5 min per session [106]
for up to 24 weeks [107] with a weekly volume of up to
36 min [108]. Most studies with longer stretching dura-
Table 1 (continued)
tions that found static stretch-mediated hypertrophy were

performed after 2020 [38–41, 109–113] and were therefore
not included in the aforementioned review [45]. The cur-
rent state of the literature investigating stretch-mediated
Study
hypertrophy is listed in Table 2, with changes in maximal tension on the myofibrils is present. The tension may be
strength listed in Table 3. actively or passively conveyed to the contractile proteins.
The animal research on birds in Table 1 shows consistent It seems likely that rate of muscle growth is proportional
effects regarding stretch-induced hypertrophy using different to the time that tension is applied to the muscle fiber.” Fur-
stretching techniques such as attaching weight to the wing thermore, similar stretching-induced signaling pathways,
and using cardboard sleeves or stretching bands to apply biochemical, and physical changes [80, 118–122] includ-
stretching times that ranged from 0.5 h to 24 h per day every ing growth hormone and IGF-1 [117, 123] were frequently
day (except for the three studies by Antonio and Gonyea reported in animal experiments, supporting the importance
[50, 82, 114]) for up to 56 days. Conversely, Tables 2 and of stretching-induced mechanical overload to activate ana-
3 illustrate conflicting results in human studies regarding bolic pathways possibly via stretch-activated channels [72,
stretch-induced hypertrophy and strength using stretching 124, 125]. Increases in muscle mass were accompanied by
devices such as stretching boards or orthoses, while only an increase in deoxyribonucleic acid and ribonucleic acid
Simpson et al. [44] used external weights via a leg press to levels [79, 126], growth promoting factors [61, 98, 127]
perform 3 min of stretch per session. While hypertrophy and and IGF-1 messenger ribonucleic acid [128] as well as
strength increases can be found with stretching times of as increases in the rates of muscle protein synthesis [61, 119,
little as 4 × 30 s per day, 3 days per week for 8 weeks [115], 126, 129]. Interestingly, comparable with strength train-
other interventions with similar or longer stretching dura- ing, Sasai et al. [69] demonstrated that mTOR/p70S6K
tions per session and overall intervention periods reported is activated acutely after stretching. Furthermore, Agata
no significant changes [38, 42, 116]. Studies using a com- et al. [130] showed significantly increased phosphorylation
parable load control to animal studies were performed in of Akt and p70S6K due to a stretching intervention period
2022 and 2023 applying high stretching volumes of 7–14 h of 2 weeks in which the rat soleus was stretched with an
per week with daily stretching of 1–2 h [40, 109, 112, 113] intermittent protocol for 15 min per day.
showing more consistent effects regarding hypertrophy and In the literature on birds, further specific adaptations
strength increases. following stretching interventions were reported. Roman
and Alway [97], Alway [88] and Kennedy et al. [83]
showed a shift in myosin heavy chain expression toward
5 Discussion slower myosin isoform of slow myosin 2, decreasing con-
traction velocity [118], which was explained as a more
The aim of this narrative review was to discuss the impact favorable energetic state of the muscle to handle chronic
of stretch-induced mechanical tension as an underly- mechanical overload [96]. Furthermore, an increase in
ing mechanism on muscle hypertrophy and strength fiber number was reported frequently [48, 77, 82, 114]
enhancements. and may be attributable to hyperplasia effects. The authors
discussed different theories including splitting of existing
muscle fibers and activation of satellite cells [50, 77, 85,
5.1 Stretching‑Induced Muscle Hypertrophy 96]. Just one study by Alway [91] determined an approxi-
and Maximal Strength in Animals mately 95% maximal strength increase following 30 days
of chronic stretch. Finally, Sola et al. [77] concluded that
There are different approaches to explain stretch-mediated “[i]f stretch can be maintained, there appears to be little
hypertrophy. Most prominently, the majority of the authors limit to extent and duration of the hypertrophy” (p. 95).
from animal studies attributed increases in muscle mass Based on these studies, the transferability to mammals
to stretching-induced mechanical overload [77, 90]. Devol remains speculative, as there were substantial differences
et al. [117] referred to mechanical tension per sarcomere from the studies on birds [100–103].
as an important factor inducing hypertrophy, which would There are different limitations, complicating a con-
explain the rapid increase in fiber cross-sectional area in clusive final statement attributing adaptations exclu-
the first days [97, 99]. When the stretch stimulus was not sively to mechanical tension. Chicken and quail stretched
re-adjusted, hypertrophy plateaued after these first few one side only, partially using weights attached to the
days [79, 97], which was suggested to be the result of wing. Thus, it is difficult to distinguish between passive
longitudinal hypertrophy (increased number sarcomeres mechanical overload and contraction of the bird against
in series) causing a reduction of mechanical tension per the weight causing muscle mass increases [89]. However,
sarcomere. Accordingly, supporting the initially stated consideration of studies from Holly et al. [80] and Barnett
hypothesis of shared mechanisms between strength and [79] describing no increased electromyographic activity
stretch training, Ashmore [118] indicated that “[t]he com- in the stretched muscles and Sola et al. [77] reporting
mon factor present in all cases of muscle growth is that reduced but still significant stretch-induced hypertrophy
Table 2 Characteristics of longitudinal human studies on muscle hypertrophy

Study Subjects Interven- Muscle and stretching volume Methods Results (hypertrophy increase)
tion period mean % increasea
(weeks)
Longo et al. [122] n = 30 12 Plantar flexors Image acquisition: B-mode No significant changes
2 exercises: each 5 × 45 s/day 5 ultrasound
days/week Control condition: separate pas-
sive control group
Mizuno [123] n = 35 8 Plantar flexors Image acquisition: B-mode Muscle thickness in SS
SS (with additional ES) 4 × 30 ultrasound group + 5.8%
s/day 3 days/week Control condition: separate pas- Muscle thickness in SS + ES
sive control group group + 13.4%
Additional information: 2 IGs,
one with SS only and one com-
bining SS with ES
Panidi et al. [41] n = 21 12 Plantar flexors Image acquisition: EFOV ultra- MCSA in intervened leg + 23%
5 days/week starting with 540 sound MCSA in control leg + 13%
s/day and increasing to 900 Control condition: contralateral
s/day control leg
Sato et al. [42] n = 24 6 Plantar flexors Image acquisition: B-mode No significant changes
Group 1: 3 × 120 s/day 3 days/ ultrasound
week Control condition: uncontrolled
Group 2: 1 × 360 s/day 1 day/ Additional information: 2 IGs, 1
week with 120 s/day and 1 with 360
s/day
Simpson et al. [44] n = 21 6 Plantar flexors Image acquisition: B-mode Muscle thickness + 5.6%
3 min/day 5 days/week ultrasound
Control condition: separate pas-
sive control group
Yahata et al. [38] n = 16 5 Plantar flexors Image acquisition: B-mode No significant changes
6 × 5 min/day 2 days/week ultrasound
Control condition: contralateral
control leg
Warneke et al. [117] n = 52 6 Plantar flexors Image acquisition: B-mode Muscle thickness in lateral head
60 min/day 7 days/week ultrasound of gastrocnemius + 15.3%
Control condition: separate pas-
sive control group and contralat-
eral control leg
Warneke et al. [120] n = 69 6 Plantar flexors Image acquisition: B-mode Muscle thickness in lateral head
60 min/day 7 days/week ultrasound of gastrocnemius + 4.68%
Control condition: separate pas- Muscle thickness in medial head
sive control group and contralat- of gastrocnemius + 7.72%
eral control leg
Warneke et al. [121] n = 45 6 Plantar flexors Image acquisition: B-mode ultra- MCSA via MRI in lateral head of
60 min/day 7 days/week sound and MRI gastrocnemius + 8.8%
Control condition: separate pas- MCSA via MRI in medial head of
sive control group and contralat- gastrocnemius + 5.68%
eral control leg Muscle thickness via ultrasound
in lateral head of gastrocne-
mius + 7.9%
Muscle thickness via ultrasound
in medial head of gastrocne-
mius + 7.29%
Wohlann et al. [124] n = 44 6 Lower extremity muscles, i.e., Image acquisition: B-mode ultra- No significant changes
quadriceps, hamstrings, glu- sound in vastus lateralis
teal muscles, plantar flexors Control condition: separate pas-
4 exercises: each 5 min/day 7 sive control group and contralat-
days/week eral control leg
B-mode brightness mode, EFOV extended field-of-view, ES electrical stimulation, IGs intervention groups, min minutes, MCSA muscle cross-
sectional area, MRI magnetic resonance imaging, n number of participants, sec seconds, SS static stretching
a
Because of inconsistency in reporting, the % change is only stated as the mean value
in denervated muscle, indicates that voluntary contractions in addition to regular volleyball training. Of note, no pas-
against the stretching device may not be a pre-condition sive control group was included in either study. In 2022
for hypertrophy. While Hotta et al. [28] pointed out a sig- and 2023, Warneke et al. [40, 109, 112, 113] examined the
nificantly reduced blood flow to the stretched muscle, fur- effects of daily 1-h stretch training on muscle growth and
ther explanatory approaches, such as hypoxia, however, strength parameters, reporting significant hypertrophy. With
were not discussed in most animal studies. Notably, in enhancements in muscle thickness of approximately 5–15%,
humans, muscle blood flow restriction training has shown the results seem to be comparable to those reported from
hypertrophic responses [131–134]. Additionally, because resistance training, with hypertrophy of up to 15% following
animals had to be dissected for examination, there was 12 weeks of resistance training [139–141], but limited to the
no real control group included in any of the listed stud- calf muscles. Consequently, performing long-lasting stretch-
ies, lowering the study quality. Finally, the recently per- ing could be hypothesized as an alternative when aiming to
formed meta-analysis [49] shows significant limitations, as induce muscle hypertrophy. Interestingly, the effects of 1 h
high study heterogeneity as well as dependency of effects of daily stretching were directly compared to a 3-days per
were ignored. Although high consistency in the direction week 5 × 12 repetition calf raise training, with no signifi-
of effects can be assumed (almost all showed increases), cant difference in adaptations regarding maximum strength,
multiple effect sizes originating from the same study were hypertrophy, and flexibility observed. In a randomized trial
not pooled, leading to an overestimation of the concluded in humans, there was an 18% gain in plantar flexor strength
effect size. In summary, while in mammals results are in full knee extension following stretching and a 13% gain
contradictory, high consistency in chronic stretch-induced in strength following a 5 × 12 repetition calf raise training
muscle hypertrophy in birds was reported. The explanatory 3 times a week with an extended knee joint. When the knee
approach including mechanical tension as the only stimu- was bent to 90 degrees for isometric testing, the results sug-
lus seems to be limited and one-dimensional. gested a ~ 10% increase for both conditions [112]. Although
the participants were described as “recreationally trained”, it
5.2 Stretching‑Induced Muscle Hypertrophy can be assumed that the participants were not familiar with
and Maximal Strength in Humans long-lasting stretching, but were likely familiar with strength
training. Therefore, the unexpectedly high stretch-induced
While animal studies exhibit high effect consistency, there increases could possibly be attributed to a new training stim-
are contrasting results in humans raising questions about ulus. Because exclusively long-lasting static stretching stud-
the transferability of animal results [49, 86, 90]. Previous ies were able to show hypertrophy, the practical applicability
reviews by Shrier [135], Medeiros and Lima [136] and seems to be limited to special circumstances. For example,
most recently Arntz et al. [137] reported small magnitude it would be difficult to practically implement and compare
effects of stretch training on maximal strength; however, 7 h of calf stretch training to 3 × 15 min of calf raise training
no systematic review found stretch-induced hypertrophy in per week [142]. Furthermore, almost all studies reporting
humans. Accordingly, in 2020, Nunes et al. [45] were not increases in maximal strength or hypertrophy used external
able to demonstrate stretch-induced hypertrophy. Of note, in devices, either a stretching board [38, 41, 110] or stretching
contrast to animal research, most human studies used short orthoses [109, 112, 113] and applied long-lasting stretch-
stretching durations showing no significant hypertrophy. ing to isolated, lower extremity muscle groups, typically the
Only Simpson et al. [44] and Mizuno et al. [115] reported plantar flexors [38, 41, 44, 109, 112, 113] or the hamstrings
muscle thickness increases of 5.6% using 3 min of plantar [143]. Aiming to transfer measured adaptations to larger
flexor stretching 5 times per week for 6 weeks and 5.8% fol- muscle groups and more complex movements, Wohlann
lowing 8 weeks of plantar flexor stretching 2 min per session et al. [116] applied 4 static stretches to the lower extrem-
with 3 sessions per week, respectively. However, the statisti- ity, including the plantar flexors, hamstrings, quadriceps,
cal procedure in the Simpson et al. [44] study was criticized, and gluteal muscles (each for 5 min daily) without using
with a request for a recalculation of the results [138]. Since stretching devices. Even though maximal strength increased
2020, new evidence emerged examining longer stretching slightly (by about 4%), no muscle hypertrophy was found.
durations and/or higher training frequencies. Yahata et al. There are different explanatory approaches for the dis-
[38] used session durations of 30 min (divided into 6 × 5 min crepancies in the results. Firstly and most likely, following
performed 2 days per week) and reported significant maxi- the theory of mechanically-induced protein synthesis stimu-
mal strength increases without accompanied hypertrophy. lation, stretching intensity can be assumed to play a key role
Panidi et al. [41] observed plantar flexors muscle hypertro- in adaptations [144]. In line with this theory, van der Pjil
phy of 23% in the intervened leg and approximately 13% in et al. [145, 146] described the importance of titin unfolding
the control leg using stretching durations of up to 15 min, for stimulating anabolic signaling leading to hypertrophic
5 days per week in adolescent female volleyball players responses. As titin can be assumed to unfold with high
Table 3 Characteristics of longitudinal human studies on maximal strength
Study Subjects Intervention period Muscle and stretching volume Methods Results (maximal strength increase)
mean % c hangea
Abdel-Aziem and Mohammad [125] n = 75 6 weeks Plantar flexors Contraction type: dynamic (concentric Trained participants:
5 × 30 s twice/day 5 days/week and eccentric) concentric peak torque 30°/sec + 9.11%,
Control condition: separate passive concentric peak torque 120°/
control group sec + 11.64%, eccentric peak torque
Additional information: 1 IG with 30°/sec + 11.92%, eccentric peak
trained participants, 1 IG with torque 120°/sec + 12.54%
untrained participants Untrained participants:
concentric peak torque 30°/sec –
0.25%, concentric peak torque 120°/
sec + 14.51, eccentric peak torque 30°/
sec + 11.96%, eccentric peak torque
120°/sec + 9.38%
Akagi and Takahashi [116] n = 19 5 weeks Plantar flexors Contraction type: isometric No significant changes
3 × 2 min/day on 6 days/week Control condition: contralateral control
leg
Caldwell et al. [126] n = 30 2 weeks Quadriceps and hamstrings Contraction type: isometric IG with 2 sessions/day + 7.1% in quadri-
3 × 30 s, either once or twice per day Control condition: contralateral control cep MVC
for 14 days leg No significant increase for both groups

Additional information: 2 IGs, in hamstring MVC and quadricep
1 IG stretched once/day, MVC for IG with 1 session/day
1 IG stretched twice/day
Chen et al. [127] n = 30 8 weeks Hamstrings Contraction type: isokinetic concentric MVC in flexion (hamstrings) + 8.67%
30 × 30 s/day 3 days/week at 60°/sec MVC in extension (quadriceps) + 3.04%
Control condition: separate passive
control group
Additional information: MVC tested in
quadriceps as well
Guissard and Duchateau [128] n = 12 6 weeks Plantar flexors Contraction type: ballistic and isometric No significant changes
10 min/day 5 times/week Control condition: contralateral control
leg
Ikeda and Ryushi [129] n = 25 6 weeks Knee extensors Contraction type: isometric MVC + 10.16%
6 × 30 s/day 3 days/week Control condition: separate passive
control group
Kokkonen et al. [130] n = 38 10 weeks Lower extremity muscles, i.e., Contraction type: dynamic 1RM Knee flexion 1RM + 15.3%
quadriceps, hamstrings, gluteus, Control condition: separate passive Knee extension 1RM + 32.4%
abductors, adductors, plantar flex- control group
ors, dorsiflexors
15 exercises: each 3 × 15 s/day 3
days/week
Konrad and Tilp [131] n = 41 6 weeks Plantar flexors Contraction type: isometric No significant changes
4 × 30 s/day 5 days/week Control condition: separate passive con-
trol group and contralateral control leg
2065
Table 3 (continued)
2066
mean % c hangea
LaRoche et al. [132] n = 29 4 weeks Hip extensors Contraction type: isokinetic hip exten- Peak torque + 5.3%
10 × 30 s/day 3 days/week sion at 60°/sec
control group
Longo et al. [122] n = 30 12 weeks Plantar flexors Contraction type: isometric No significant changes
2 exercises: each 5 × 45 s/day 5 days/ Control condition: separate passive
week control group
Marshall et al. [133] n = 22 4 weeks Hamstrings Contraction type: isokinetic at 30°/sec No significant changes
4 exercises: each 3 × 30 s/day 5 days/ and 120°/sec
week Control condition: separate passive
control group
Mizuno [123] n = 35 8 weeks Plantar flexors Contraction type: dynamic 1RM SS + 20.8%
4 × 30 s/day 3 days/week Control condition: separate passive SS&ES + 22.4%
control group
Additional information: 2 IGs, 1 with SS
only and
1 combining SS with ES
Nakamura et al. [43] n = 40 4 weeks Plantar flexors Contraction type: isometric No significant changes
3 × 60 s/day 3 days/week Control condition: separate passive
control group
Additional information: high-intensity
and low-intensity static stretching
groups
Nakao et al. [134] n = 30 4 weeks Hamstrings Contraction type: isometric and isoki- Peak torque 60°/sec + 10.55%
1 × 5 min/day 3 days/week netic at 60°/sec and 180°/sec Peak torque 180°/sec + 13.61%
Control condition: separate passive No significant changes in isometric
control group testing
Nelson et al. [130] n = 25 10 weeks Plantar flexors Contraction type: dynamic 1RM 1RM in intervened leg + 29%
4 × 30 s/day 3 days/week Control condition: separate passive con- 1RM in control leg + 11%
Sato et al. [42] n = 24 6 weeks Plantar flexors Contraction type: isometric at 30° plan- No significant changes
Group 1: 120 s/day 3 days/week tar flexion, neutral and 15° dorsiflexion
Group 2: 360 s/day 1 day/week Control condition: uncontrolled
Additional information: 2 IGs, 1 with
120 s/day 3 days/week and 1 with 360
s/day 1 day/week
Simpson et al. [44] n = 21 6 weeks Plantar flexor Contraction type: isometric No significant changes
3 min/day 5 days/week Control condition: separate passive
control group
K. Warneke et al.
Table 3 (continued)
mean % c hangea
Warneke et al. [117] n = 52 6 weeks Plantar flexors Contraction type: dynamic 1RM and MVC + 16.2%
60 min/day 7 days/week isometric 1RM + 25.3%
Control condition: separate passive con-
Warneke et al. [120] n = 69 6 weeks Plantar flexors Contraction type: isometric with MVC with extended knee joint + 18%
60 min/day 7 days/week extended and bent knee joint MVC with bent knee joint + 9.58%
control group
Warneke et al. [121] n = 45 6 weeks Plantar flexors Contraction type: isometric with MVC with extended knee joint + 9.44%
60 min/day 7 days/week extended sand bent knee joint MVC with bent knee joint + 4.84%
Control condition: separate passive con-
Warneke et al. [40] n = 70 6 weeks Plantar flexors Contraction type: isometric MVC in 60-min group + 15.3%
60 min/day or 120 min each 7 days/ Control condition: separate passive con- MVC in 120-min group + 22.9%
week trol group and contralateral control leg
Additional information: 2 IGs,
1 IG stretched 60 min/day,
1 IG stretched 120 min/day
Warneke et al. [118] n = 35 6 weeks Plantar flexors Contraction type: isometric MVC + 10.5%
10 min/day 7 days/week Control condition: separate passive
control group
Wohlann et al. [124] n = 44 6 weeks Lower extremity muscles, i.e., Contraction type: isometric in leg press MVC + 4.4%
quadriceps, hamstrings, gluteus, Control condition: separate passive con-
plantar flexors trol group and contralateral control leg
4 exercises: each 5 min/day 7 days/
week
Worrell et al. [135] n = 19 3 weeks Hamstrings Contraction type: isokinetic at 60°/sec Eccentric peak torque 60°/sec + 8.5%,
4 × 15–20 s/day 5 days/week and 120°/sec eccentric peak torque 120°/
Control condition: uncontrolled sec + 13.5%, concentric peak torque
120°/sec + 11.2%, no significant
increase for concentric peak torque
60°/sec
Yahata et al. [38] n = 16 5 weeks Plantar flexors Contraction type: isokinetic at 30°/ MVC isometric with neutral ankle
6 × 5 min/day 2 days/week sec and 120°/sec and isometric at 30° angle + 6.4
plantar flexion and neutral MVC isokinetic at 120°/sec + 7.8%
Control condition: contralateral control Other tests showed no significant
leg changes
1RM one repetition maximum, ES electrical stimulation, IG Intervention group, min minutes, MVC maximal voluntary contraction, MVIC maximal voluntary isometric contraction, n number of
participants, sec seconds
a
Because of inconsistency in reporting, the % change is only stated as the mean value
2067
sarcomere lengths only, reaching high muscle lengths seems since those increases were higher than or similar to reported
essential, assuming the activation of titin kinase involving adaptations from strength training [7], which should still be
downstream pathways [46]. Furthermore, Wackerhage et al. considered the gold standard when maximizing strength and
[46] described titin unfolding related interactions with pro- muscle mass.
tein synthesis influencing factors such as muscle RING-fin- Unexpectedly high hypertrophy effects could possibly be
ger protein 1 and 2—proteasome and autophagy signaling, attributed to limitations of the measurement devices. Stretch-
which are hypothesized to be involved in muscle protein mediated hypertrophy was measured via ultrasound in most
synthesis regulation. Apart from titin unfolding, mechani- of the listed studies [41, 49, 112, 116, 153]. Even though
cally overloading filamin is described to be associated with ultrasound is stated as highly reliable and valid [158], Eng-
further biochemical responses known to be linked to hyper- lish et al. [159] as well as Hebert et al. [160] raised concerns
trophy, such as the Chaperon regulator-3, mammalian target about using sonography to assess muscle thickness, refer-
of rapamycin complex-1 (Bag3/mTORC1/YAP) [transcrip- ring to low quality and emphasizing the dependency of the
tional co-activator] pathway [46]. If quantified, most studies measured muscle thickness on the investigator-dependent
used a subjective measure of pain threshold to determine pressure applied on the transducer. Accordingly, magnetic
the stretching intensity. Unfortunately, however, in a cross- resonance imaging is described as the gold standard meas-
sectional study, Lim and Park [147] found no correlation urement investigating hypertrophy [161], which was exclu-
between passive torque (passive tension of the muscle devel- sively used in Warneke et al. [113]. Further heterogeneity
oped versus the stretching) and subjective pain perception. in study design arises from potential sex-related differences.
When assuming mechanical tension (measured via passive Panidi et al. [41] assessed female participants only, while
torque) to be of high importance for muscle hypertrophy most authors performed stretching with male participants
induced via stretching, a more objective intensity documen- [38, 42–44, 108, 143, 157]. Assuming differences in hormo-
tation by quantifying passive torque could be an appropriate nal status influence muscle hypertrophy [162], only Warneke
option. For example, stretching intensity could be stated as et al. [111] explored sex-related differences in stretch-medi-
a percentage value of the measured maximal passive torque ated hypertrophy and maximal strength increases showing
to improve objectivity—comparable with strength training significantly higher effects in male participants. In accord-
stating the percentage of the one repetition maximum. How- ance with the present explanatory approach, the authors
ever, as no studies could be found using this procedure, the hypothesized that lower flexibility baseline values in male
practical applicability remains unclear. individuals would enhance the applied absolute mechanical
Further heterogeneity is observed with intervention peri- tension with stretching compared to female individuals and
ods. In 2018, Freitas et al. [148] reviewed the available lit- therefore enhance the magnitude of adaptations.
erature and concluded that it seems unreasonable to assume Apart from mechanical tension-induced signaling cas-
structural adaptations following about 20 min of stretching cades, other explanations such as blood flow restriction [28,
per week for less than 8 weeks. Consequently, no hyper- 131, 133] or neuromuscular adaptations [109, 151] should
trophic adaptations could be assumed in most of the listed be considered in further studies. Literature exploring the
studies using intervention periods of less than 6 weeks and/ underlying physiological mechanisms in humans is scarce.
or weekly stretching volumes of below 20 min [44, 45, 115, In accordance with hypotheses from animal studies, Kremer
149, 150]. In contrast, considering underlying physiologi- [72] described the activation of growth factors such as fibro-
cal mechanisms as well as animal research, using high vol- blast growth factor, IGF-1, and mTOR leading to stimulation
umes of stretching as evaluated by Warneke et al. [112, 113], of anabolic pathways such as Akt/PKB, tuberous sclerosis
Wohlann et al. [116], Panidi et al. [41] and Yahata et al. [38] 1 and 2 due to stretch training via stretch-activated chan-
seems logical. nels [69, 125] linked to further physiological responses [68].
It is well known that there are further factors influenc- It is hypothesized that there are contractile and metabolic
ing training adaptations. Almost all of the studies included adaptations due to changes in protein synthesis via changes
untrained or recreationally active participants [38, 42–44, in protein kinases and transcription factors after mechani-
108, 143, 151–156]. In another study, participants were cally overloading the muscle, independent of resistance or
described as “trained” when performing aerobic activity stretch training [124]. Only Fowles et al. [56] investigated
3 times per week for more than 20 min per session [157]; the acute effects of 33 min of stretching on protein synthesis
however, this categorization of “highly trained” may be and failed to show any changes. Furthermore, Smith et al.
debatable. Consequently, studies showing stretch-mediated [163] reported stretching to be sufficient to induce micro-
hypertrophy in well-trained participants or even elite ath- traumatization of the muscular tissue in the long term. How-
letes are lacking. Therefore, maximal strength increases of ever, Wohlann et al. [116] were not able to reproduce these
up to 29% using 4 × 30 s of stretch 3 times per week [151] results using 4 exercises, performed for 5 min per day. More-
or 23% of hypertrophy [41] must be considered critically, over, the authors described the changes in creatine kinase
values reported by Smith et al. [163] as clinically irrelevant. conclude muscle hypertrophy may occur using long-lasting
Therefore, no comprehensive conclusions because of high stretching interventions (> 30 min per muscle per session)
heterogeneity in the study design as well as the outcomes in humans as well as in animals. Unfortunately, because no
can be drawn. study explored signaling pathways or muscle protein synthe-
sis rate, the underlying mechanisms can only be discussed
5.3 Effects of Long‑Term Stretching on Longitudinal based on animal research, and therefore remain speculative
Hypertrophy in humans.
Increases in muscle mass could also be attributed to 5.5 Practical Applications

increases in muscle fascicle length by increases in serial
sarcomere number. Currently, there is only indirect evidence Practical applicability is limited assuming that commonly
for longitudinal hypertrophy in humans, derived from theo- used resistance training would result in similar adaptations
retical or simulated muscle models [164, 165]. However, with significantly less time effort compared with long-dura-
there are some indications for enhanced muscle lengths. tion stretch training [112]. However, in situations without
Chen and colleagues and LaRoche et al. [143, 156] reported access or the possibility to perform strength training, the
beneficial effects on force development, while Yahata et al. possibility of stretch-induced muscle hypertrophy should
[38] showed improved maximal strength production in long be kept in mind especially in situations of reduced physi-
muscle lengths, while no adaptations were reported in short cal activity or prolonged phases of immobilization after
muscle lengths after stretching interventions. As stated by injury or surgery. Because no active movement is required
Kruse et al., “[…] it is assumed that stretching treatments in stretching, it could provide a sufficient alternative or early
may also induce such adaptations in humans […]. However, step in rehabilitation processes to avoid muscle atrophy and
the scientific evidence has not yet confirmed this assumption performance losses. However, hypertrophic effects were
and the overall effectiveness of stretching in humans is still exclusively reported using long-lasting static stretching in
in question […].”[165]. the calf muscle. Additionally, when muscles are immobi-
lized in a lengthened position for high-volume stretching,
5.4 Limitations the antagonist muscles are immobilized in a shortened posi-
tion. Since Williams et al. [76] reported a rapid decrease in
As this is a narrative review, the authors attempted to reflect serial sarcomeres and considering mechanical stretch as an
the essential state of the literature by performing an extended underlying mechanism of hypertrophy, atrophic effects of
study search. However, because there is a vast number of the antagonist might be assumed. However, further studies
studies, especially regarding the effects of stretching on are needed in this area, as well as on the effects in different
flexibility in humans and on hypertrophy in animals, it was populations, such as patients.
necessary to focus the literature search, which possibly led
to some studies missing in the review article. To analyze
studies addressing our research question, we started by
screening recent systematic review articles addressing the 6 Conclusions
topic [45, 48, 49, 135, 136]. Subsequently, related articles
and reference lists were screened to find articles excluded in To date, the underlying mechanisms of stretch-mediated
the aforementioned systematic reviews. Furthermore, only muscle strength increases and muscle hypertrophy in
studies investigating the effects of stretching on strength or humans are not fully clarified. Even though some studies
strength-related parameters, such as peak torque, maximal using long-lasting stretching of ≥ 30 min per session showed
voluntary contractions (eccentric, isometric, or concentric), strength increases and hypertrophy, these studies are lim-
or muscle mass-related parameters, were considered in this ited to the calf muscle and opposed by a high number of
review. For a comprehensive review of the literature, system- studies using shorter stretching durations without any effect.
atic reviews are needed, focusing on the effects of stretch- Because of high study heterogeneity and limited intensity
ing on different outcomes—flexibility, maximal strength and quantification in stretching studies, further literature seems
muscle hypertrophy—separately. Obviously, there are fun- necessary to state a final conclusion.
damental differences regarding strength training and stretch- Because in humans only a few studies have explored first
ing. Although there seem to be similarities in the outcomes physiological parameters, the discussion is based on ani-
(hypertrophy, maximal strength) induced by (high-volume) mal results. The scarce evidence from human investigations
stretching or strength training, a narrative review does not addressing protein synthesis and muscle damage provides
provide mean effect sizes or confidence intervals. However, only an inconclusive picture that seems to be in conflict
based on the studies discussed above, it seems reasonable to with indications from animal studies. Therefore, the role of
mechanical tension in humans needs further clarification by 2. Maestroni L, Read P, Bishop C, Turner A. Strength and power
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Funding Open access funding provided by University of Klagenfurt. vertical jump height in elite soccer players. Br J Sports Med.
2004;38:285–8.
Conflicts of interest/competing interests Konstantin Warneke, Lars 7. Schoenfeld BJ, Peterson MD, Ogborn D, Contreras B, Sonmez
H. Lohmann, Camila D. Lima, Karsten Hollander, Andreas Konrad, GT. Effects of low- vs. high-load resistance training on muscle
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David G. Behm have no conflicts of interest that are directly related Res. 2015;29:2954–63.
to the content of this article. There were no financial interests of any 8. Schoenfeld BJ, Pope ZK, Benik FM, Hester GM, Sellers J,
authors contributing to the paper. Nooner JL, et al. Longer interset rest periods enhance muscle
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Consent to participate Not applicable. relationship between resistance training volume and muscle
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Open Access This article is licensed under a Creative Commons Attri- 2007;37:225–64.
bution 4.0 International License, which permits use, sharing, adapta- 14. Behm DG. Neuromuscular implications and applications of
tion, distribution and reproduction in any medium or format, as long resistance training. J Strength Cond Res. 1995;9:264–74.
as you give appropriate credit to the original author(s) and the source, 15. Schoenfeld BJ, Grgic J, Ogborn D, Krieger JW. Strength and
provide a link to the Creative Commons licence, and indicate if changes hypertrophy adaptations between low- vs. high-Load resist-
were made. The images or other third party material in this article are ance training: a systematic review and meta analysis. J Strength
included in the article's Creative Commons licence, unless indicated Cond Res. 2017;31:3508–23.
otherwise in a credit line to the material. If material is not included in 16. Lixandrão ME, Ugrinowitsch C, Berton R, Vechin FC, Con-
the article's Creative Commons licence and your intended use is not ceição MS, Damas F, et al. Magnitude of muscle strength and
permitted by statutory regulation or exceeds the permitted use, you will mass adaptations between high-load resistance training ver-
need to obtain permission directly from the copyright holder. To view a sus low-load resistance training associated with blood-flow
copy of this licence, visit http://creativecommons.org/licenses/by/4.0/. restriction: a systematic review and meta-analysis. Sports Med.
2018;48:361–78.
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Authors and Affiliations
Konstantin Warneke1,10,11 · Lars H. Lohmann2 · Camila D. Lima3 · Karsten Hollander4 · Andreas Konrad5,10 ·
Astrid Zech6 · Masatoshi Nakamura7 · Klaus Wirth8 · Michael Keiner9 · David G. Behm10
6
* Konstantin Warneke Department of Human Motion Science and Exercise
[email protected]; Physiology, Friedrich Schiller University, Jena, Germany
[email protected] 7
Faculty of Rehabilitation Sciences, Nishi Kyushu University,
1 Ozaki, Kanzaki, Saga, Japan
Institute for Exercise, Sport and Health, Leuphana
8
University, Universitätsallee 1, 21335 Lüneburg, Institute of Sport Science, University of Applied Sciences
Deutschland, Germany Wiener Neustadt, Wiener Neustadt, Austria
2 9
University Sports Center, Carl von Ossietzky University Department of Sport Science, German University of Health
Oldenburg, Oldenburg, Germany and Sport, Ismaning, Germany
3 10
School of Medical and Health Sciences, Edith Cowan School of Human Kinetics and Recreation, Memorial
University, Joondalup, WA, Australia University of Newfoundland, St. John’s, NL, Canada
4 11
Institute of Interdisciplinary Exercise Science and Sports Institute of Sport Science, Alpen-Adria University
Medicine, MSH Medical School Hamburg, Hamburg, Klagenfurt, Klagenfurt, Germany
Germany
5
Institute of Human Movement Science, Sport and Health,
University of Graz, Graz, Austria

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Sports Medicine (2023) 53:2055–2075

Physiology of Stretch‑Mediated Hypertrophy and Strength Increases:

Accepted: 22 July 2023 / Published online: 9 August 2023

thickness [11, 15, 16]. However, force output also depends

hypertrophy of muscular tissue that counteracts performance

and quails. While the literature provides extensive evidence

33.6 ± 7.1% after 5 days and

of immobilization of one limb and stretching for 40 min

stretching alone without immobilization resulted in no sig-

a significant decrease in muscle mass in rat soleus muscle

with the contralateral control. The methodology used in

these studies is therefore not comparable to the procedure

used in studies including birds, which are listed in Table 1.

overall protein synthesis rate [49, 104, 105].

4 Stretch‑Mediated Hypertrophy: Results

As early as 1983, after finding stretch-mediated hypertrophy

found on the topic [45], no studies with comparable train-

tions that found static stretch-mediated hypertrophy were

Table 2 Characteristics of longitudinal human studies on muscle hypertrophy

for 14 days leg No significant increase for both groups

Increases in muscle mass could also be attributed to 5.5 Practical Applications

Authors and Affiliations

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4 Stretch‑Mediated Hypertrophy: Results

Increases in muscle mass could also be attributed to 5.5 Practical Applications