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Front. Sports Act. Living, 15 November 2022
Sec. Elite Sports and Performance Enhancement
https://doi.org/10.3389/fspor.2022.1035190 (https://doi.org/10.3389/fspor.2022.1035190)

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Inter-set stretch: A potential time-e!cient strategy for enhancing

skeletal muscle adaptations

Brad J. Schoenfeld (https://www.frontiersin.org/people/u/455300)1*,

Henning Wackerhage (https://www.frontiersin.org/people/u/560924)2 and Eduardo

De Souza3

1
Department of Exercise Science and Recreation, Lehman College, Bronx, NY, United States
2
Department of Sport and Exercise Sciences, Technical University of Munich, Munich, Germany
3 Department of Health Sciences and Human Performance, The University of Tampa, Tampa, FL, United States

Time is considered a primary barrier to exercise adherence. Therefore, developing

time-e!cient resistance training (RT) strategies that optimize muscular adaptations
is of primary interest to practitioners. A novel approach to the problem involves
combining intensive stretch protocols with RT. Conceivably, integrating stretch into
the inter-set period may provide an added stimulus for muscle growth without
increasing session duration. Mechanistically, stretch can regulate anabolic signaling
via both active and passive force sensors. Emerging evidence indicates that both
:
 lengthening contractions against a high load as well as passive stretch can acutely
activate anabolic intracellular signaling pathways involved in muscle hypertrophy.
Although longitudinal research investigating the e"ects of stretching between RT
sets is limited, some evidence suggests it may in fact enhance hypertrophic
adaptations. Accordingly, the purpose of this paper is threefold: (1) to review how
the active force of a muscle contraction and the force of a passive stretched are
sensed; (2) to present evidence for the e"ectiveness of RT with inter-set stretch for
muscle hypertrophy (3) to provide practical recommendations for application of
inter-set stretch in program design as well as directions for future research.

Introduction

Skeletal muscle is intricately involved in human locomotion and mobility, as well as

playing an essential role in metabolic health (1). To accomplish its varied tasks,
skeletal muscle has a high degree of plasticity, allowing it to readily adapt to various
stimuli. Of note, skeletal muscle can hypertrophy, operationally defined as an
increase in the axial cross-sectional area of a myofiber or whole muscle (2), when
repeatedly subjected to external resistance. This capacity for growth has important
implications in wellness, functional capacity, athletic performance and aesthetic
pursuits (1, 2).

In humans, muscle hypertrophy is primarily achieved by the regimented performance

of resistance training (RT), where myofibers are dynamically shortened and
lengthened under load. Traditional RT programs involve performing multiple sets of a
variety of exercises that target the body's major muscle groups. In this scenario, a set
is performed for a given exercise and then the individual rests for a given amount of
time (usually 1 to 3 mins) before performing another set. Depending on the RT goal,
the amount of rest between sets can influence time-e!ciency, either increasing or
decreasing the length of the RT sessions duration.

Time is considered a primary barrier to exercise adherence (3, 4). Therefore,

developing time-e!cient RT strategies that optimize muscular adaptations is of
primary interest to practitioners. Several popular strategies have been devised to
achieve this objective, including the use of supersets and drop sets. However, while
these methods can reduce the duration of training sessions, their e!cacy for
enhancing muscular gains remains equivocal (2).

A novel approach to the problem involves combining intensive stretch protocols with
RT. Conceivably, integrating stretch into the inter-set period may provide an added
stimulus for muscle growth without increasing session duration. Accordingly, the
purpose of this paper is 3-fold: (1) to review how the active force of a muscle
contraction and the force of a passive stretch are sensed; (2) to present evidence for
:
 the e"ectiveness of RT with inter-set stretch for muscle hypertrophy (3) to provide
practical recommendations for application of inter-set stretch in program design as
well as directions for future research.

Theoretical basis for force and stretch-induced hypertrophy

The dry mass of a muscle is ≈70% protein (5); hence, the accretion of skeletal muscle
tissue is largely a function of net protein balance (i.e., protein synthesis > protein
breakdown). In this regard, muscles primarily hypertrophy when they synthesize
more protein than they break down over a given time frame. Mechanical force,
exercise-induced muscle damage, and metabolic stress are proposed candidate
hypertrophy stimuli (6). Of these putative mechanisms, mechanical force is generally
considered the most relevant to muscle growth. When a muscle is subjected to
mechanical stimuli, the body initiates a response via a process called
mechanotransduction, whereby these forces are transduced to chemical signals that
regulate anabolic and catabolic processes through a variety of intracellular enzymatic
pathways (7). Forces can be transduced longitudinally (from z-disk to z-disk within
each myofibril) and/or laterally via a series of radially oriented elastic elements
including the endomysium, epimysium and perimysium (8). For the purpose of this
review, we will focus on the sensing of contraction and stretch-induced force (i.e.,
mechanosensing) as hypertrophy stimuli. Two types of force sensors are
distinguished.

Active force sensors sense the force of a muscle contraction. Active force sensors
either localize to the Z-disc that anchors actin, or to protein complexes called
costameres that help to direct the force generated by contracting sarcomeres to the
extracellular matrix surrounding each myofiber (6).

Passive force sensors are activated by the passive stretch of a muscle. Passive force
sensors generally lie in parallel to the force generating actin and myosin and actually
“go slack” during a force-generating concentric action that pulls on active force
sensors (6). Passive force sensors generally experience force when muscles are
lengthened (e.g., during a stretch).

What follows is an overview of candidate force sensors in skeletal muscle. We first

discuss active force sensors and then consider passive force sensors.

There is some evidence of a filamin C-BAG3-mTORC1-protein synthesis pathway,

which to date is the only mechanism that links the active force produced during a
muscle contraction via an uninterrupted chain of events to muscle protein synthesis
(MPS) (6). Filamin C is a Z-disc bound protein and potential mechanosensor that is
coupled to BAG3. BAG3 not only regulates the degradation of filamin C by autophagy
:
 but is also linked to two mTORC1-activating mechanisms (6). The first of these
mTORC1-activating mechanisms involves the binding of BAG3 to the mTORC1
inhibitors TSC1 and TSC2 via its so-called WW domain (the “WW” signifies the two
tryptophans). Indeed, TSC1 appears to move near BAG3 after maximal eccentric
exercise of a human muscle and at the same time activation-related mTOR
phosphorylation increases (9). The second mTORC1-activating mechanism is that
mechano-activated BAG3 activates the Hippo e"ector YAP via an intermediate step.
Active YAP then increases the expression of the genes that encode the leucine
transporter LAT1 (10). Consistent with this process, resistance exercise increases the
expression of the leucine transporter LAT1 in human muscle (10). While this
mechanism is far from fully characterized, it suggests that mechanical stress causes
BAG3 to bind to mTOR and Hippo inhibitors, resulting in the activation of mTORC1-
dependent protein synthesis and a greater uptake of leucine, which would further
activate mTORC1 and the protein synthetic machinery.

Another candidate active force sensor is focal adhesion kinase (FAK). FAK is a
component of costameres that convey the forces generated by the cytoskeleton and
by sarcomeres out of the muscle fiber to the surrounding extracellular matrix (11).
FAK is involved in IGF-1-induced muscle hypertrophy via TSC2 and mTORC1 (12).
However, high load contractions of rat muscles do not appear to increase FAK Tyr397
phosphorylation (13). Thus, while FAK is a potential active force sensor capable of
mTORC1 activation, there is currently no evidence of a functional role during
hypertrophy-inducing contractions.

Regarding passive force sensors, titin is a candidate sensor as it has been proposed to
respond to mechanical stretch (14). Titin, the largest known protein in the human
body, possesses elastic properties that allow it to act as a molecular spring (14). Titin
is anchored to the Z-disc and extends to the M-line (14), and lies parallel to the
force-generating actin and myosin filaments. This suggests that titin should go slack
during a shortening contraction but generate tension during passive stretch of a
muscle (14). Titin has a force-activated kinase domain, so it possesses the ability to
phosphorylate other proteins (15). However, there is no known signaling link to the
master MPS regulator mTORC1 and consequently there is so far no evidence that the
activation of titin by passive stretch increases MPS.

The nuclei of muscle fibers “flatten” during passive stretch of a muscle fiber (16).
Research in non-muscle cells indicates that such nuclear flattening can activate the
Hippo e"ector YAP, which drives the expression of the genes that encode the leucine
LAT1 transporter (17). Although there are no experimental data on this as yet, we can
speculate that passive stretch may cause nuclei to flatten, followed by YAP influx and
activity, increased LAT1 expression (18), greater leucine influx, mTORC1 activation and
ultimately greater MPS. This hypothesis remains to be tested.
:
 Finally, when a membrane is stretched, mechanosensitive channels or stretch-
activated channels (SAC) open and become permeable to ions such as Ca2+ and Na+
(19). The main function of stretch-activated channels is to “inform” the nervous
system about mechanical stimuli such as the touch of the skin. It has been proposed
that stretching of muscle fibers opens SACs, initiating an intracellular signaling
cascade that induces calcium-dependent anabolic signaling via mTOR and its
downstream target enzymes (20). The role of SACs in mechanostransduction has
been demonstrated in multiple studies. For example, Spangenburg and McBride (21)
showed that inhibition of SACs blunted activation of p70S6K, a key regulator of
protein translation, after performance of eccentric actions. Moreover, Mirozev et al.
(19) established a link between SAC activity and downstream anabolic signaling
pathways in rat skeletal muscle during acute recovery following a period of
mechanical unloading. These findings are consistent with evidence that blockage of
SACs correlates with a blunted hypertrophic response to training in rodents (22).
Although a mechanistic rationale for SAC-induced anabolic e"ects remains
undetermined, possible mediators include enhanced cytoskeletal remodeling,
calmodulin/calcineurin interactions, and elevated levels of heat shock protein-70
(23).

In summary, myofibers contain active force sensors and passive force sensors. Thus,
the force generated by a muscle contraction and the force that results from a passive
stretch are likely sensed by di"erent mechanosensors. While evidence is lacking for
the role of passive sensors mediating anabolic responses, it is plausible to assume
that their stimulation might activate mTORC1 and MPS or other events that
contribute to muscle hypertrophy via several mechanisms (see Figure 1).

Figure 1
:
 (https://www.frontiersin.org/files/Articles/1035190/fspor-04-1035190-HTML-r1/im-
age_m/fspor-04-1035190-g001.jpg)

Figure 1. Putative mechanisms of passive sensors mediating anabolic responses.

The e"ects of passive stretch on acute anabolic factors

Given that there are several candidate force sensors that respond to passive stretch,
the question arises: Can passive stretching between sets be employed as a strategy
to further enhance the protein synthetic response achieved with traditional RT
protocols? Emerging evidence indicates that both lengthening contractions against a
high load as well as passive stretch can activate anabolic intracellular signaling
pathways involved in muscle hypertrophy. In particular, an extensive body of research
in rodents demonstrates increased mTOR signaling in response to passive tension.
These studies include models employing stretched cultured myoblasts and myotubes
(24–26) as well as ex vivo stretch of isolated muscles (27–29).

Evidence strongly implicates mitogen-activated protein kinase (MAPK) as an

upstream mediator of stretch-induced intracellular signaling. Increased
phosphorylation of MAPK has consistently been observed following passive stretch,
although the degrees of involvement of specific kinases in the MAPK family (i.e., ERK
1/2, p38 and JNK) have been inconsistent (30–33). Martineau and Gardiner (32)
demonstrated that p54-JNK signaling was responsive to both peak tension and the
time/tension interval following in situ passive stretch of the rat medial
:
 gastrocnemius; the application of a contraction stimulus to the stretched muscle
amplifies JNK signaling, with the greatest e"ects observed following eccentric
actions (33).

The PI3K/Akt pathway also appears to be involved in muscle stretch. In vitro research
by Sasai et al. (26) found that primary cultures of chick skeletal myotubes subjected
to passive cyclic stretching for 72 h activates the PI3K/Akt pathway, resulting in
muscle hypertrophy. Similarly, Sakamoto et al. (34) demonstrated that passively
stretching the incubated fast-twitch dominant extensor digitorum longus muscle for
10 min elicited a 2-fold increase in Akt activity. Intriguingly, stretch did not
significantly increase Akt activation in the predominantly slow-twitch soleus muscle
despite a marked phosphorylation of MAPK p38. Adding further context to the
evidence, Russ (35) electrically stimulated the tibialis anterior muscles of rats in situ at
di"erent frequencies across di"erent lengths. Results showed that active contractile
force had a more pronounced e"ect on Akt phosphorylation than passive tension;
however, increasing passive tension during active muscle contraction had a
synergistic e"ect on the degree of Akt activation. These findings suggest a potential
additive role for passive tension, whereby the combination of stretch and contractile
forces heighten Akt signaling and thus potentially produce a greater hypertrophic
response.

Other intracellular signaling molecules have been implicated with stretch protocols,
as well. Wang et al. (36) found that passive stretch (15 min, 5 days a week for 2
weeks) of the gastrocnemii in mice upregulated mRNA expression of the anabolic
factors p70S6K and myogenin while downregulating the catabolic factors MuRF1,
MAFbx, myostatin, and 4E-BP1. Moreover, Czerwinski et al. (37) showed that IGF-I
mRNA abundance of the avian patagialis muscle increased 3-fold after 11 days of
load stretch of the birds' wings. Upregulation of IGF-1 mRNA expression following
passive stretch by plaster cast immobilization of the lower limb muscles in a
lengthened position have been reported elsewhere in rabbits (38).

While intracellular signaling activity provides insights into anabolic and catabolic
processes, skeletal muscle hypertrophy is ultimately a function of protein balance,
whereby the rate of MPS exceeds that of proteolysis over time. Animal models
consistently show marked increases in MPS following passive stretch protocols
(39–41). These trials demonstrate that tension alone, in the absence of contractile
activity, can stimulate a protein synthetic response. A notable exception to these
findings by Atherton et al. (42) reported a reduction in MPS in cultured L6 skeletal
muscle cells for up to 120 min after a bout of cyclic stretch. Paradoxically, results
occurred despite an increased phosphorylation of intracellular signaling kinases
associated with anabolism. In contrast to animal models, Fowles et al. (43) failed to
demonstrate an increase in soleus MPS following ~27 min intermittent passive plantar
:
 flexor stretch in a cohort of 8 healthy men. Given the discrepancies between human
and animal models, further research is required to better understand the e"ects of
varied stretching protocols on anabolic signaling and MPS in human skeletal muscle.

Hypertrophic adaptations to longitudinal isolated stretch protocols

Compelling evidence shows that when animals have a limb immobilized in a

lengthened position it results in muscle hypertrophy while immobilizing the limb in a
shortened position induces atrophy (44). A unique finding of these studies is that at
least some of the hypertrophy occurs by adding sarcomeres in series (i.e., along the
longitudinal axis) (44), as opposed to traditional RT protocols where a majority of
hypertrophy occurs from the addition of sarcomeres in parallel (45).

Avian models consistently show marked hypertrophy when the wings of birds are
stretched and then subjected to an external load (46–49). These protocols involve
attaching a tube filled with lead pellets to the birds' stretched wings either on a
continuous or intermittent basis over the course of several weeks. The magnitude of
hypertrophy associated with such interventions appears to be greater in type I
compared to type II myofibers (46). Moreover, murine research showed that manual,
passive dorsiflexion stretch performed for 15 min, 5 days per week significantly
increased cross-sectional area CSA of the gastrocnemii across a 2-week study period
(36).

Although animal research provides interesting insights into the role of passive tension
in hypertrophic adaptations, the extreme protocols employed in these models
cannot necessarily be generalized to ecologically valid human stretch training.
Indeed, longitudinal passive stretch interventions in humans generally do not show
appreciable increases in muscle mass when performed using traditional protocols.
For example, Sato et al. (50) subjected healthy males to 360 s static stretch of the
plantar flexors per week for 6 weeks. Participants were randomized to perform the
stretch protocol either as one weekly session or to distribute the stretch training
across three sessions; total time under stretch was equated between groups. Results
showed no post-study change in muscle thickness measures of the medial
gastrocnemius; stretch frequency had no e"ect on the response. A follow-up study
from the same lab reported no appreciable changes in plantar flexor hypertrophy in
healthy young men after 6 weeks of two weekly 30-min stretch training sessions (51).
Moreover, Junior et al. (52) reported that performing two, 25-s sets of passive stretch
immediately prior to leg extension exercise attenuated muscle attenuated increases
in vastus lateralis CSA compared to a RT-only condition (12.7 vs. 7.4%, respectively)
over a 10-week study period. This finding was associated with a decrease in the
number of repetitions performed, suggesting that negative e"ects of stretching prior
to RT may be related to reductions in volume load (i.e., repetitions x load).
:
 Alternatively, some evidence does show a hypertrophic benefit to passive stretch
protocols. Simpson et al. (53) found that loaded stretch of the plantar flexors
(performed on a leg press machine) increased gastrocnemius muscle thickness vs. a
non-stretched control after 3 weeks (~11% versus ~5%, respectively); however,
adaptations were similar at the end of the 6-week study period (~9% for both
conditions). It is curious that a non-training control would experience appreciable
hypertrophy over a relatively short time frame, raising questions as to the validity of
these findings. Recently, Warneke et al. (54) reported robust increases in muscle
thickness (~15%) of the lateral gastrocnemius following a 6-week stretch protocol for
the calf muscles using a specially designed orthosis. It should be noted that
participants stretched for an hour every day over the study period with an individual
rating of discomfort of 8 on a scale of 1 to 10. Although the study provides evidence
that passive stretch training can in fact promote hypertrophy in free-living humans,
the high volume, frequency and intensity required to achieve results would be
impractical for the majority of the population.

Hypertrophic adaptations to inter-set stretch

Theoretically, combining passive stretch within a RT routine may enhance

hypertrophic adaptations compared to RT alone. While several practical options exist,
interspersing rest within the inter-set rest periods may be the best way to implement
such a strategy. Ideally, stretching should be performed immediately following the
final eccentric action of the RT set, which may potentiate the tension imposed on the
muscles via residual e"ects of eccentric actions (55). In this hypothetical model,
active lengthening leverages titin's role as a molecular spring by increasing its
sti"ness, and thus its force, to a greater extent than when passively lengthened (55).
Hence, stretching immediately after eccentric loading may result in greater passive
tension on the muscle following cross-bridge deactivation than what would be
experienced in traditional static stretching, conceivably mediated via heightened titin
sti"ness (55). In addition to the heightened tension in the stretch, the strategy allows
for a greater time-under-tension during a given session, which has been proposed as
a driving factor in hypertrophy (56).

There is limited longitudinal research investigating the e"ects of stretching between

RT sets, but some findings suggest it may be a viable strategy to enhance
hypertrophy. Silva et al. (57) provided preliminary evidence that inter-set stretch may
enhance hypertrophic adaptations. Resistance-trained men performed straight-leg
plantar flexion exercise either with a standard passive rest between sets or with the
inclusion of a 30-s loaded intra-set stretch. The researchers reported that the intra-
set stretch condition elicited a >2-fold increase in MT of the gastrocnemius
:
 compared to control. It should be noted that these findings were presented as a
conference abstract but never published in a refereed journal, thus preventing
scrutiny of the study's methodology.

Evangelista et al. (58) randomly assigned untrained young men to perform an 8-week
total-body RT program with either a passive between-set rest interval or a 30-s inter-
set stretch integrated into the 90 s rest period. The stretch was unloaded, but
reportedly performed to the point of temporary discomfort. Results indicated that
the inter-set stretch condition elicited superior summed increases in muscle
thickness for muscles of the upper and lower limbs vs. the passive rest condition
(10.5 vs. 6.7%, respectively).

In a within-subject design, Van Every et al. (59) randomized the lower limbs of
untrained young men to perform plantar flexion exercises with either a 2-min passive
rest period or a 20-s inter-set stretch at the same working load followed by 100 s of
passive rest. After 8-weeks, results showed greater increases in muscle thickness of
the soleus for the stretch condition compared to control; these results were
observed despite a decrease in RT volume load (5 to 12%) in the stretch condition.
Alternatively, inter-set stretch did not show an appreciable hypertrophic benefit in
the gastrocnemii. Given that the soleus is a predominantly slow-twitch muscle while
the gastrocnemius is a mixed-fiber type muscle (60), these findings suggest that
inter-set stretch may be more e"ective in hypertrophy of type I muscle fibers. This is
an intriguing finding, as evidence indicates that type I fibers have a diminished
hypertrophic potential compared to type II fibers (61). Thus, inter-set stretch may be
a unique strategy to target development of the more “stubborn” type I fibers.

Employing an eccentric-focused protocol, Nakamura et al. (62) investigated the

e"ects of adding a 30-s inter-set stretch to a flywheel squat program performed
twice weekly in untrained young men. Results showed that improvements in strength
measures tended to favor stretch compared to control; however, increases in
quadriceps muscle thickness were generally similar between conditions. It should be
noted that the stretch condition in this study involved moving from the flywheel unit
to a massage table and then assuming the designated stretch (62), whereas the
aforementioned studies performed inter-set stretch immediately after each set of RT
(57–59). This time lag from moving between the exercise to the table and getting
properly positioned for the stretch may have diminished the residual e"ects of
eccentric actions on the ensuing stretch, and thus impaired hypertrophic
enhancements.

In contrast to other studies on the topic, Wadhi et al. (63) found similar increases in
muscle thickness and strength of the pectoralis major in a group of resistance-
trained men who performed an 8-week chest-oriented training protocol consisting
of flat and incline bench press exercises with either a 30-s loaded intra-set stretch or
a passive rest interval. Discrepancies between studies conceivably may be explained
:
 by di"erences in the studies' designs. Wadhi et al. (63) employed a stretch on a
di"erent exercise (cable fly) amounting to 15% of participants working load from the
prior set on the bench-press or incline bench-press exercises. While the timer under
stretch was comparable to other studies (e.g., 30's), it is possible that the tension
provided by the cable fly machine may not have imposed a su!cient additional
stimulus to promote enhanced hypertrophic responses, particularly in the sample of
trained cohorts. In addition, as with the study by Nakamura et al. (62), Wadhi et al.
(63) employed a transition period between bench press performance and the stretch
in the cable fly machine, which may have diminished the potential residual e"ects of
eccentric actions on the ensuing stretch. Similar to the findings of Nakamura et al.
(62), the inter-set stretching did not compromise muscular strength adaptations.

Conclusion and practical applications

In summary, emerging evidence suggests that inter-set stretch may enhance

muscular adaptations compared to traditional RT programs without increasing the
time spent exercising. Its e"ectiveness appears to be predicated on performing the
stretch immediately after the final repetition of a set, which conceivably takes
advantage of the residual e"ects of previous eccentric actions. This research should
be considered somewhat preliminary and requires further study to draw stronger
inferences on its implications. However, given that several studies have observed
beneficial hypertrophic e"ects with no evidence of a detriment, the strategy would
seem to have a good cost-benefit profile. Intriguingly, despite some research
showing acute strength impairments subsequent to static stretching protocols (64),
current research does not indicate deleterious e"ects across di"erent populations on
long-term strength outcomes and in some instances the strategy shows positive
adaptations (59, 62).

In addition to a general need for further exploration on the topic, several questions
remain to be answered as to best practice guidelines. First, while the preliminary
evidence to date suggests the need to stretch to some level of discomfort to induce
additive benefits to RT, no attempts have been made to quantify the degree of
intensity required to optimize results. Conceivably there is a threshold for level of
tension beyond which no further increases are observed in anabolic signaling. In vitro
research indicates that a high magnitude of strain is required to maximally stimulate
p70S6K (65), but it is not clear how these findings translate in vivo. Based on the
limited current research to date, it would seem that inter-set stretch should be
carried out to a discomfort level of at least an ‘8' on the rating of perceived exertion
scale (range of 1 to 10). Further study is warranted with protocols employing varying
levels of stretch intensity.
:
 Second, how long should the stretch be held between sets? Research to date has
employed intra-set stretch durations of 20 to 30 s; would longer durations promote
a superior benefit or perhaps blunt results by negatively impacting the volume load
of the RT session? Given that 20 second bouts of high-intensity inter-set stretch
have been shown to enhance hypertrophy, this should be the minimum duration
employed until research indicates otherwise. Further study is warranted with di"ering
stretch durations during the intra-set rest period.

Third, are the e"ects of stretching between RT sets population-specific? Age, sex and
training status are known to influence exercise-induced adaptations and thus
conceivably may play a role in the response to the strategy. Studies to date have
focused exclusively on young men. Therefore, further studies are warranted to
evaluate muscular adaptations following inter-set stretch protocols in alternative
populations.

Finally, does inter-set stretch induce fiber type-specific adaptations and/or is it

specific to certain muscle groups? As noted, avian models of loaded stretch show
greater hypertrophy in muscles comprised predominantly of type I compared to type
II myofibers (46). The aforementioned study by Van Every et al. (59) found that inter-
set stretch elicited greater hypertrophy in the soleus, a type I dominant muscle,
compared to the gastrocnemii, a mixed-fiber muscle. Replication of these findings
and further research into the mechanisms of inter-set stretch is needed to better
understand this phenomenon and its potential practical implications to program
design.

Author contributions

BJS conceived of the idea for the paper. All authors contributed to the writing and
revision of the paper, and approved the final draft.

Conflict of interest

Author BJS serves on the scientific advisory board for Tonal Corporation, a
manufacturer of fitness equipment.

The remaining authors declare that the research was conducted in the absence of
any commercial or financial relationships that could be construed as a potential
conflict of interest.
:
 Publisher's note

All claims expressed in this article are solely those of the authors and do not
necessarily represent those of their a!liated organizations, or those of the publisher,
the editors and the reviewers. Any product that may be evaluated in this article, or
claim that may be made by its manufacturer, is not guaranteed or endorsed by the
publisher.

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Keywords: hypertrophy, contraction, lengthening, force sensors, mechanical tension

Citation: Schoenfeld BJ, Wackerhage H and De Souza E (2022) Inter-set stretch: A potential time-e!cient
strategy for enhancing skeletal muscle adaptations. Front. Sports Act. Living 4:1035190 (tel:1035190). doi:
10.3389/fspor.2022.1035190
:
 Received: 05 September 2022; Accepted: 02 November 2022;
Published: 15 November 2022.

Edited by:
Olivier Girard (https://loop.frontiersin.org/people/44793/overview), University of Western Australia, Australia

Reviewed by:
Brendan Richard Scott (https://loop.frontiersin.org/people/176955/overview), Murdoch University, Australia
Zsolt Radak (https://loop.frontiersin.org/people/46523/overview), Semmelweis University, Hungary
Ben Dascombe (https://loop.frontiersin.org/people/244808/overview), The University of Newcastle, Australia

Copyright © 2022 Schoenfeld, Wackerhage and De Souza. This is an open-access article distributed under the
terms of the Creative Commons Attribution License (CC BY) (http://creativecommons.org/licenses/by/4.0/). The
use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright
owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted
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*Correspondence: Brad J. Schoenfeld, brad.schoenfeld@lehman.cuny.edu

(mailto:brad.schoenfeld@lehman.cuny.edu)

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