TYPE Review

PUBLISHED 15 November 2022

DOI 10.3389/fspor.2022.1035190

Inter-set stretch: A potential

OPEN ACCESS time-efficient strategy for
enhancing skeletal muscle
EDITED BY
Olivier Girard,
University of Western
Australia, Australia

REVIEWED BY
adaptations
Brendan Richard Scott,
Murdoch University, Australia
Zsolt Radak, Brad J. Schoenfeld1*, Henning Wackerhage2 and
Semmelweis University, Hungary
Ben Dascombe, Eduardo De Souza3
The University of Newcastle, Australia 1
Department of Exercise Science and Recreation, Lehman College, Bronx, NY, United States,
2
*CORRESPONDENCE Department of Sport and Exercise Sciences, Technical University of Munich, Munich, Germany,
Brad J. Schoenfeld 3
Department of Health Sciences and Human Performance, The University of Tampa, Tampa, FL,
[email protected] United States
SPECIALTY SECTION
This article was submitted to
Elite Sports and Performance Time is considered a primary barrier to exercise adherence. Therefore,
Enhancement,
developing time-efficient resistance training (RT) strategies that optimize
a section of the journal
Frontiers in Sports and Active Living muscular adaptations is of primary interest to practitioners. A novel approach
RECEIVED 05 September 2022 to the problem involves combining intensive stretch protocols with RT.
ACCEPTED 02 November 2022 Conceivably, integrating stretch into the inter-set period may provide an
PUBLISHED 15 November 2022
added stimulus for muscle growth without increasing session duration.
CITATION
Mechanistically, stretch can regulate anabolic signaling via both active and
Schoenfeld BJ, Wackerhage H and De
Souza E (2022) Inter-set stretch: A passive force sensors. Emerging evidence indicates that both lengthening
potential time-efficient strategy for contractions against a high load as well as passive stretch can acutely activate
enhancing skeletal muscle
adaptations.
anabolic intracellular signaling pathways involved in muscle hypertrophy.
Front. Sports Act. Living 4:1035190. Although longitudinal research investigating the effects of stretching between
doi: 10.3389/fspor.2022.1035190 RT sets is limited, some evidence suggests it may in fact enhance hypertrophic
COPYRIGHT adaptations. Accordingly, the purpose of this paper is threefold: (1) to
© 2022 Schoenfeld, Wackerhage and
De Souza. This is an open-access
review how the active force of a muscle contraction and the force of a
article distributed under the terms of passive stretched are sensed; (2) to present evidence for the effectiveness
the Creative Commons Attribution of RT with inter-set stretch for muscle hypertrophy (3) to provide practical
License (CC BY). The use, distribution
or reproduction in other forums is recommendations for application of inter-set stretch in program design as well
permitted, provided the original as directions for future research.
author(s) and the copyright owner(s)
are credited and that the original
publication in this journal is cited, in KEYWORDS
accordance with accepted academic hypertrophy, contraction, lengthening, force sensors, mechanical tension
practice. No use, distribution or
reproduction is permitted which does
not comply with these terms.
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).

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Schoenfeld et al. 10.3389/fspor.2022.1035190

In humans, muscle hypertrophy is primarily achieved by force (i.e., mechanosensing) as hypertrophy stimuli. Two types
the regimented performance of resistance training (RT), where of force sensors are distinguished.
myofibers are dynamically shortened and lengthened under Active force sensors sense the force of a muscle contraction.
load. Traditional RT programs involve performing multiple sets Active force sensors either localize to the Z-disc that anchors
of a variety of exercises that target the body’s major muscle actin, or to protein complexes called costameres that help to
groups. In this scenario, a set is performed for a given exercise direct the force generated by contracting sarcomeres to the
and then the individual rests for a given amount of time (usually extracellular matrix surrounding each myofiber (6).
1 to 3 mins) before performing another set. Depending on the Passive force sensors are activated by the passive stretch
RT goal, the amount of rest between sets can influence time- of a muscle. Passive force sensors generally lie in parallel to
efficiency, either increasing or decreasing the length of the RT the force generating actin and myosin and actually “go slack”
sessions duration. during a force-generating concentric action that pulls on active
Time is considered a primary barrier to exercise adherence force sensors (6). Passive force sensors generally experience force
(3, 4). Therefore, developing time-efficient RT strategies that when muscles are lengthened (e.g., during a stretch).
optimize muscular adaptations is of primary interest to What follows is an overview of candidate force sensors in
practitioners. Several popular strategies have been devised to skeletal muscle. We first discuss active force sensors and then
achieve this objective, including the use of supersets and drop consider passive force sensors.
sets. However, while these methods can reduce the duration of There is some evidence of a filamin C-BAG3-mTORC1-
training sessions, their efficacy for enhancing muscular gains protein synthesis pathway, which to date is the only mechanism
remains equivocal (2). that links the active force produced during a muscle contraction
A novel approach to the problem involves combining via an uninterrupted chain of events to muscle protein synthesis
intensive stretch protocols with RT. Conceivably, integrating (MPS) (6). Filamin C is a Z-disc bound protein and potential
stretch into the inter-set period may provide an added mechanosensor that is coupled to BAG3. BAG3 not only
stimulus for muscle growth without increasing session duration. regulates the degradation of filamin C by autophagy but is
Accordingly, the purpose of this paper is 3-fold: (1) to review also linked to two mTORC1-activating mechanisms (6). The
how the active force of a muscle contraction and the force of first of these mTORC1-activating mechanisms involves the
a passive stretch are sensed; (2) to present evidence for the binding of BAG3 to the mTORC1 inhibitors TSC1 and TSC2
effectiveness of RT with inter-set stretch for muscle hypertrophy via its so-called WW domain (the “WW” signifies the two
(3) to provide practical recommendations for application of tryptophans). Indeed, TSC1 appears to move near BAG3 after
inter-set stretch in program design as well as directions for maximal eccentric exercise of a human muscle and at the same
future research. time activation-related mTOR phosphorylation increases (9).
The second mTORC1-activating mechanism is that mechano-
activated BAG3 activates the Hippo effector YAP via an
Theoretical basis for force and intermediate step. Active YAP then increases the expression
stretch-induced hypertrophy of the genes that encode the leucine transporter LAT1 (10).
Consistent with this process, resistance exercise increases the
The dry mass of a muscle is ≈70% protein (5); hence, the expression of the leucine transporter LAT1 in human muscle
accretion of skeletal muscle tissue is largely a function of net (10). While this mechanism is far from fully characterized,
protein balance (i.e., protein synthesis > protein breakdown). it suggests that mechanical stress causes BAG3 to bind to
In this regard, muscles primarily hypertrophy when they mTOR and Hippo inhibitors, resulting in the activation of
synthesize more protein than they break down over a given mTORC1-dependent protein synthesis and a greater uptake of
time frame. Mechanical force, exercise-induced muscle damage, leucine, which would further activate mTORC1 and the protein
and metabolic stress are proposed candidate hypertrophy stimuli synthetic machinery.
(6). Of these putative mechanisms, mechanical force is generally Another candidate active force sensor is focal adhesion
considered the most relevant to muscle growth. When a muscle kinase (FAK). FAK is a component of costameres that convey
is subjected to mechanical stimuli, the body initiates a response the forces generated by the cytoskeleton and by sarcomeres
via a process called mechanotransduction, whereby these forces out of the muscle fiber to the surrounding extracellular
are transduced to chemical signals that regulate anabolic and matrix (11). FAK is involved in IGF-1-induced muscle
catabolic processes through a variety of intracellular enzymatic hypertrophy via TSC2 and mTORC1 (12). However, high load
pathways (7). Forces can be transduced longitudinally (from z- contractions of rat muscles do not appear to increase FAK
disk to z-disk within each myofibril) and/or laterally via a series Tyr397 phosphorylation (13). Thus, while FAK is a potential
of radially oriented elastic elements including the endomysium, active force sensor capable of mTORC1 activation, there is
epimysium and perimysium (8). For the purpose of this review, currently no evidence of a functional role during hypertrophy-
we will focus on the sensing of contraction and stretch-induced inducing contractions.

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Regarding passive force sensors, titin is a candidate sensor as activate mTORC1 and MPS or other events that contribute to
it has been proposed to respond to mechanical stretch (14). Titin, muscle hypertrophy via several mechanisms (see Figure 1).
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 The effects of passive stretch on
lies parallel to the force-generating actin and myosin filaments. acute anabolic factors
This suggests that titin should go slack during a shortening
contraction but generate tension during passive stretch of a Given that there are several candidate force sensors
muscle (14). Titin has a force-activated kinase domain, so that respond to passive stretch, the question arises: Can
it possesses the ability to phosphorylate other proteins (15). passive stretching between sets be employed as a strategy to
However, there is no known signaling link to the master MPS further enhance the protein synthetic response achieved with
regulator mTORC1 and consequently there is so far no evidence traditional RT protocols? Emerging evidence indicates that both
that the activation of titin by passive stretch increases MPS. lengthening contractions against a high load as well as passive
The nuclei of muscle fibers “flatten” during passive stretch stretch can activate anabolic intracellular signaling pathways
of a muscle fiber (16). Research in non-muscle cells indicates involved in muscle hypertrophy. In particular, an extensive body
that such nuclear flattening can activate the Hippo effector YAP, of research in rodents demonstrates increased mTOR signaling
which drives the expression of the genes that encode the leucine in response to passive tension. These studies include models
LAT1 transporter (17). Although there are no experimental employing stretched cultured myoblasts and myotubes (24–26)
data on this as yet, we can speculate that passive stretch may as well as ex vivo stretch of isolated muscles (27–29).
cause nuclei to flatten, followed by YAP influx and activity, Evidence strongly implicates mitogen-activated protein
increased LAT1 expression (18), greater leucine influx, mTORC1 kinase (MAPK) as an upstream mediator of stretch-induced
activation and ultimately greater MPS. This hypothesis remains intracellular signaling. Increased phosphorylation of MAPK has
to be tested. consistently been observed following passive stretch, although
Finally, when a membrane is stretched, mechanosensitive the degrees of involvement of specific kinases in the MAPK
channels or stretch-activated channels (SAC) open and become family (i.e., ERK 1/2, p38 and JNK) have been inconsistent
permeable to ions such as Ca2+ and Na+ (19). The main (30–33). Martineau and Gardiner (32) demonstrated that p54-
function of stretch-activated channels is to “inform” the JNK signaling was responsive to both peak tension and the
nervous system about mechanical stimuli such as the touch time/tension interval following in situ passive stretch of the rat
of the skin. It has been proposed that stretching of muscle medial gastrocnemius; the application of a contraction stimulus
fibers opens SACs, initiating an intracellular signaling cascade to the stretched muscle amplifies JNK signaling, with the greatest
that induces calcium-dependent anabolic signaling via mTOR effects observed following eccentric actions (33).
and its downstream target enzymes (20). The role of SACs The PI3K/Akt pathway also appears to be involved in
in mechanostransduction has been demonstrated in multiple muscle stretch. In vitro research by Sasai et al. (26) found
studies. For example, Spangenburg and McBride (21) showed that primary cultures of chick skeletal myotubes subjected to
that inhibition of SACs blunted activation of p70S6K, a key passive cyclic stretching for 72 h activates the PI3K/Akt pathway,
regulator of protein translation, after performance of eccentric resulting in muscle hypertrophy. Similarly, Sakamoto et al.
actions. Moreover, Mirozev et al. (19) established a link between (34) demonstrated that passively stretching the incubated fast-
SAC activity and downstream anabolic signaling pathways in twitch dominant extensor digitorum longus muscle for 10 min
rat skeletal muscle during acute recovery following a period elicited a 2-fold increase in Akt activity. Intriguingly, stretch did
of mechanical unloading. These findings are consistent with not significantly increase Akt activation in the predominantly
evidence that blockage of SACs correlates with a blunted slow-twitch soleus muscle despite a marked phosphorylation
hypertrophic response to training in rodents (22). Although a of MAPK p38. Adding further context to the evidence, Russ
mechanistic rationale for SAC-induced anabolic effects remains (35) electrically stimulated the tibialis anterior muscles of rats
undetermined, possible mediators include enhanced cytoskeletal in situ at different frequencies across different lengths. Results
remodeling, calmodulin/calcineurin interactions, and elevated showed that active contractile force had a more pronounced
levels of heat shock protein-70 (23). effect on Akt phosphorylation than passive tension; however,
In summary, myofibers contain active force sensors and increasing passive tension during active muscle contraction
passive force sensors. Thus, the force generated by a muscle had a synergistic effect on the degree of Akt activation. These
contraction and the force that results from a passive stretch findings suggest a potential additive role for passive tension,
are likely sensed by different mechanosensors. While evidence whereby the combination of stretch and contractile forces
is lacking for the role of passive sensors mediating anabolic heighten Akt signaling and thus potentially produce a greater
responses, it is plausible to assume that their stimulation might hypertrophic response.

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Schoenfeld et al. 10.3389/fspor.2022.1035190

FIGURE 1
Putative mechanisms of passive sensors mediating anabolic responses.

Other intracellular signaling molecules have been implicated the effects of varied stretching protocols on anabolic signaling
with stretch protocols, as well. Wang et al. (36) found that and MPS in human skeletal muscle.
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
Hypertrophic adaptations to
the catabolic factors MuRF1, MAFbx, myostatin, and 4E- longitudinal isolated stretch
BP1. Moreover, Czerwinski et al. (37) showed that IGF-I protocols
mRNA abundance of the avian patagialis muscle increased
3-fold after 11 days of load stretch of the birds’ wings. Compelling evidence shows that when animals have a limb
Upregulation of IGF-1 mRNA expression following passive immobilized in a lengthened position it results in muscle
stretch by plaster cast immobilization of the lower limb muscles hypertrophy while immobilizing the limb in a shortened
in a lengthened position have been reported elsewhere in position induces atrophy (44). A unique finding of these
rabbits (38). studies is that at least some of the hypertrophy occurs by
While intracellular signaling activity provides insights into adding sarcomeres in series (i.e., along the longitudinal axis)
anabolic and catabolic processes, skeletal muscle hypertrophy (44), as opposed to traditional RT protocols where a majority
is ultimately a function of protein balance, whereby the rate of hypertrophy occurs from the addition of sarcomeres in
of MPS exceeds that of proteolysis over time. Animal models parallel (45).
consistently show marked increases in MPS following passive Avian models consistently show marked hypertrophy when
stretch protocols (39–41). These trials demonstrate that tension the wings of birds are stretched and then subjected to an
alone, in the absence of contractile activity, can stimulate a external load (46–49). These protocols involve attaching a tube
protein synthetic response. A notable exception to these findings filled with lead pellets to the birds’ stretched wings either on
by Atherton et al. (42) reported a reduction in MPS in cultured a continuous or intermittent basis over the course of several
L6 skeletal muscle cells for up to 120 min after a bout of cyclic weeks. The magnitude of hypertrophy associated with such
stretch. Paradoxically, results occurred despite an increased interventions appears to be greater in type I compared to type II
phosphorylation of intracellular signaling kinases associated myofibers (46). Moreover, murine research showed that manual,
with anabolism. In contrast to animal models, Fowles et al. passive dorsiflexion stretch performed for 15 min, 5 days per
(43) failed to demonstrate an increase in soleus MPS following week significantly increased cross-sectional area CSA of the
∼27 min intermittent passive plantar flexor stretch in a cohort gastrocnemii across a 2-week study period (36).
of 8 healthy men. Given the discrepancies between human and Although animal research provides interesting insights into
animal models, further research is required to better understand the role of passive tension in hypertrophic adaptations, the

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extreme protocols employed in these models cannot necessarily a strategy. Ideally, stretching should be performed immediately
be generalized to ecologically valid human stretch training. following the final eccentric action of the RT set, which may
Indeed, longitudinal passive stretch interventions in humans potentiate the tension imposed on the muscles via residual
generally do not show appreciable increases in muscle mass effects of eccentric actions (55). In this hypothetical model,
when performed using traditional protocols. For example, Sato active lengthening leverages titin’s role as a molecular spring by
et al. (50) subjected healthy males to 360 s static stretch of increasing its stiffness, and thus its force, to a greater extent than
the plantar flexors per week for 6 weeks. Participants were when passively lengthened (55). Hence, stretching immediately
randomized to perform the stretch protocol either as one after eccentric loading may result in greater passive tension
weekly session or to distribute the stretch training across three on the muscle following cross-bridge deactivation than what
sessions; total time under stretch was equated between groups. would be experienced in traditional static stretching, conceivably
Results showed no post-study change in muscle thickness mediated via heightened titin stiffness (55). In addition to the
measures of the medial gastrocnemius; stretch frequency had heightened tension in the stretch, the strategy allows for a
no effect on the response. A follow-up study from the same lab greater time-under-tension during a given session, which has
reported no appreciable changes in plantar flexor hypertrophy in been proposed as a driving factor in hypertrophy (56).
healthy young men after 6 weeks of two weekly 30-min stretch There is limited longitudinal research investigating the
training sessions (51). Moreover, Junior et al. (52) reported that effects of stretching between RT sets, but some findings suggest
performing two, 25-s sets of passive stretch immediately prior it may be a viable strategy to enhance hypertrophy. Silva
to leg extension exercise attenuated muscle attenuated increases et al. (57) provided preliminary evidence that inter-set stretch
in vastus lateralis CSA compared to a RT-only condition (12.7 may enhance hypertrophic adaptations. Resistance-trained men
vs. 7.4%, respectively) over a 10-week study period. This finding performed straight-leg plantar flexion exercise either with a
was associated with a decrease in the number of repetitions standard passive rest between sets or with the inclusion of a
performed, suggesting that negative effects of stretching prior to 30-s loaded intra-set stretch. The researchers reported that the
RT may be related to reductions in volume load (i.e., repetitions intra-set stretch condition elicited a >2-fold increase in MT of
x load). the gastrocnemius compared to control. It should be noted that
Alternatively, some evidence does show a hypertrophic these findings were presented as a conference abstract but never
benefit to passive stretch protocols. Simpson et al. (53) found published in a refereed journal, thus preventing scrutiny of the
that loaded stretch of the plantar flexors (performed on a leg study’s methodology.
press machine) increased gastrocnemius muscle thickness vs. Evangelista et al. (58) randomly assigned untrained
a non-stretched control after 3 weeks (∼11% versus ∼5%, young men to perform an 8-week total-body RT program
respectively); however, adaptations were similar at the end of the with either a passive between-set rest interval or a 30-s
6-week study period (∼9% for both conditions). It is curious inter-set stretch integrated into the 90 s rest period. The
that a non-training control would experience appreciable stretch was unloaded, but reportedly performed to the
hypertrophy over a relatively short time frame, raising questions point of temporary discomfort. Results indicated that
as to the validity of these findings. Recently, Warneke et al. (54) the inter-set stretch condition elicited superior summed
reported robust increases in muscle thickness (∼15%) of the increases in muscle thickness for muscles of the upper
lateral gastrocnemius following a 6-week stretch protocol for and lower limbs vs. the passive rest condition (10.5 vs.
the calf muscles using a specially designed orthosis. It should 6.7%, respectively).
be noted that participants stretched for an hour every day over In a within-subject design, Van Every et al. (59) randomized
the study period with an individual rating of discomfort of 8 the lower limbs of untrained young men to perform plantar
on a scale of 1 to 10. Although the study provides evidence flexion exercises with either a 2-min passive rest period or a 20-
that passive stretch training can in fact promote hypertrophy s inter-set stretch at the same working load followed by 100 s
in free-living humans, the high volume, frequency and intensity of passive rest. After 8-weeks, results showed greater increases
required to achieve results would be impractical for the majority in muscle thickness of the soleus for the stretch condition
of the population. 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 adaptations to hypertrophic benefit in the gastrocnemii. Given that the soleus is
inter-set stretch a predominantly slow-twitch muscle while the gastrocnemius is
a mixed-fiber type muscle (60), these findings suggest that inter-
Theoretically, combining passive stretch within a RT routine set stretch may be more effective in hypertrophy of type I muscle
may enhance hypertrophic adaptations compared to RT alone. fibers. This is an intriguing finding, as evidence indicates that
While several practical options exist, interspersing rest within type I fibers have a diminished hypertrophic potential compared
the inter-set rest periods may be the best way to implement such to type II fibers (61). Thus, inter-set stretch may be a unique

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strategy to target development of the more “stubborn” type to draw stronger inferences on its implications. However,
I fibers. given that several studies have observed beneficial hypertrophic
Employing an eccentric-focused protocol, Nakamura et al. effects with no evidence of a detriment, the strategy would
(62) investigated the effects of adding a 30-s inter-set stretch to seem to have a good cost-benefit profile. Intriguingly, despite
a flywheel squat program performed twice weekly in untrained some research showing acute strength impairments subsequent
young men. Results showed that improvements in strength to static stretching protocols (64), current research does not
measures tended to favor stretch compared to control; however, indicate deleterious effects across different populations on long-
increases in quadriceps muscle thickness were generally similar term strength outcomes and in some instances the strategy
between conditions. It should be noted that the stretch condition shows positive adaptations (59, 62).
in this study involved moving from the flywheel unit to a In addition to a general need for further exploration on
massage table and then assuming the designated stretch (62), the topic, several questions remain to be answered as to best
whereas the aforementioned studies performed inter-set stretch practice guidelines. First, while the preliminary evidence to date
immediately after each set of RT (57–59). This time lag from suggests the need to stretch to some level of discomfort to
moving between the exercise to the table and getting properly induce additive benefits to RT, no attempts have been made to
positioned for the stretch may have diminished the residual quantify the degree of intensity required to optimize results.
effects of eccentric actions on the ensuing stretch, and thus Conceivably there is a threshold for level of tension beyond
impaired hypertrophic enhancements. which no further increases are observed in anabolic signaling.
In contrast to other studies on the topic, Wadhi et al. In vitro research indicates that a high magnitude of strain is
(63) found similar increases in muscle thickness and strength required to maximally stimulate p70S6K (65), but it is not clear
of the pectoralis major in a group of resistance-trained men how these findings translate in vivo. Based on the limited current
who performed an 8-week chest-oriented training protocol research to date, it would seem that inter-set stretch should
consisting of flat and incline bench press exercises with either be carried out to a discomfort level of at least an ‘8’ on the
a 30-s loaded intra-set stretch or a passive rest interval. rating of perceived exertion scale (range of 1 to 10). Further
Discrepancies between studies conceivably may be explained by study is warranted with protocols employing varying levels of
differences in the studies’ designs. Wadhi et al. (63) employed stretch intensity.
a stretch on a different exercise (cable fly) amounting to 15% Second, how long should the stretch be held between sets?
of participants working load from the prior set on the bench- Research to date has employed intra-set stretch durations of 20
press or incline bench-press exercises. While the timer under to 30 s; would longer durations promote a superior benefit or
stretch was comparable to other studies (e.g., 30’s), it is possible perhaps blunt results by negatively impacting the volume load
that the tension provided by the cable fly machine may not have of the RT session? Given that 20 second bouts of high-intensity
imposed a sufficient additional stimulus to promote enhanced inter-set stretch have been shown to enhance hypertrophy,
hypertrophic responses, particularly in the sample of trained this should be the minimum duration employed until research
cohorts. In addition, as with the study by Nakamura et al. indicates otherwise. Further study is warranted with differing
(62), Wadhi et al. (63) employed a transition period between stretch durations during the intra-set rest period.
bench press performance and the stretch in the cable fly Third, are the effects of stretching between RT sets
machine, which may have diminished the potential residual population-specific? Age, sex and training status are known
effects of eccentric actions on the ensuing stretch. Similar to the to influence exercise-induced adaptations and thus conceivably
findings of Nakamura et al. (62), the inter-set stretching did not may play a role in the response to the strategy. Studies
compromise muscular strength adaptations. 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.
Conclusion and practical Finally, does inter-set stretch induce fiber type-specific
applications adaptations and/or is it specific to certain muscle groups? As
noted, avian models of loaded stretch show greater hypertrophy
In summary, emerging evidence suggests that inter-set in muscles comprised predominantly of type I compared to type
stretch may enhance muscular adaptations compared to II myofibers (46). The aforementioned study by Van Every et al.
traditional RT programs without increasing the time spent (59) found that inter-set stretch elicited greater hypertrophy
exercising. Its effectiveness appears to be predicated on in the soleus, a type I dominant muscle, compared to the
performing the stretch immediately after the final repetition gastrocnemii, a mixed-fiber muscle. Replication of these findings
of a set, which conceivably takes advantage of the residual and further research into the mechanisms of inter-set stretch is
effects of previous eccentric actions. This research should be needed to better understand this phenomenon and its potential
considered somewhat preliminary and requires further study practical implications to program design.

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Author contributions relationships that could be construed as a potential conflict

of interest.
BJS conceived of the idea for the paper. All authors
contributed to the writing and revision of the paper, and
approved the final draft.
Publisher’s note
Conflict of interest All claims expressed in this article are solely those of the
authors and do not necessarily represent those of their affiliated
Author BJS serves on the scientific advisory board for Tonal organizations, or those of the publisher, the editors and the
Corporation, a manufacturer of fitness equipment. reviewers. Any product that may be evaluated in this article, or
The remaining authors declare that the research was claim that may be made by its manufacturer, is not guaranteed
conducted in the absence of any commercial or financial or endorsed by the publisher.

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