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From Tissue to System: What Constitutes an Appropriate Response to

Loading?

Article in Sports Medicine · November 2024

DOI: 10.1007/s40279-024-02126-w

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https://doi.org/10.1007/s40279-024-02126-w

REVIEW ARTICLE

From Tissue to System: What Constitutes an Appropriate Response

to Loading?
Tim J. Gabbett1 · Eric Oetter2

Accepted: 20 September 2024

© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2024

Abstract
Optimal loading involves the prescription of an exercise stimulus that promotes positive tissue adaptation, restoring function
in patients undergoing rehabilitation and improving performance in healthy athletes. Implicit in optimal loading is the need
to monitor the response to load, but what constitutes a normal response to loading? And does it differ among tissues (e.g.,
muscle, tendon, bone, cartilage) and systems? In this paper, we discuss the “normal” tissue response to loading schema and
demonstrate the complex interaction among training intensity, volume, and frequency, as well as the impact of these training
variables on the recovery of specific tissues and systems. Although the response to training stress follows a predictable time
course, the recovery of individual tissues to training load (defined herein as the readiness to receive a similar training stimulus
without deleterious local and/or systemic effects) varies markedly, with as little as 30 min (e.g., cartilage reformation after
walking and running) or 72 h or longer (e.g., eccentric exercise-induced muscle damage) required between loading sessions
of similar magnitude. Hyperhydrated and reactive tendons that have undergone high stretch–shorten cycle activity benefit
from a 48-h refractory period before receiving a similar training dose. In contrast, bone cells desensitize quickly to repetitive
loading, with almost all mechanosensitivity lost after as few as 20 loading cycles. To optimize loading, an additional dose
(≤ 60 loading cycles) of bone-centric exercise (e.g., plyometrics) can be performed following a 4–8 h refractory period.
Low-stress (i.e., predominantly aerobic) activity can be repeated following a short (≤ 24 h) refractory period, while greater
recovery is needed (≥ 72 h) between repeated doses of high stress (i.e., predominantly anaerobic) activity. The response of
specific tissues and systems to training load is complex; at any time, it is possible that practitioners may be optimally loading
one tissue or system while suboptimally loading another. The consideration of recovery timeframes of different tissues and
systems allows practitioners to determine the “normal” response to load. Importantly, we encourage practitioners to interpret
training within an athlete monitoring framework that considers external and internal load, athlete-reported responses, and
objective markers, to contextualize load–response data.

1 Tissue Response to Load is Variable

Key Points
As early as the 1940s, researchers demonstrated that the
We provide the typical timelines of recovery when
biochemical response of a tissue depended on the type of
exposing different tissues and systems to varying loading
training that was performed [1]. In one of the few studies
intensities, volumes, and frequencies.
to document tissue-specific load injury, Orchard et al. [2]
demonstrated that acute and chronic loads had differential Muscle, tendon, and bone are highly stressed in some
effects on tendon, bone, joint, and muscles. High chronic loading activities (e.g., sprinting). Given the different
recovery timeframes, optimal loading may occur in one
tissue while suboptimal loading may occur in another.
* Tim J. Gabbett
[email protected] We emphasize the importance of targeting the vulnerable
tissue in rehabilitation programs and “training around”
Eric Oetter
[email protected] the injured tissue to maintain global load capacity and
capacity of healthy tissues.
1
Gabbett Performance Solutions, Brisbane, QLD 4011,
Australia
2
Memphis Grizzlies, Memphis, TN 38103, USA

Vol.:(0123456789)
 T. J. Gabbett, E. Oetter

loads might be protective against some injuries (e.g., muscle speed), and the time course for recovery is dependent on the
injuries) but predictive of other injuries (e.g., joint injuries), volume and intensity of training, joint range of motion, and
while tendon injuries occur most frequently in response to contraction speed and type. A minimum of 48 h recovery is
large spikes and troughs in acute load. High-intensity and recommended between consecutive resistance sessions uti-
maximal-effort running has been shown to alter tendon lizing the same muscle groups; longer recovery is required
structure for 2 days [3, 4], with tendon structure returning to following sessions involving maximal intensity lifting owing
normal within 3–4 days. In healthy bone, mechanosensitivity to the large involvement of the central nervous system. Due
is diminished after only 20 repetitive loading cycles [5], but to the high intensity of maximal-effort sprinting activities
is 90% restored within 8 h of rest; thus, further bone adap- and the associated metabolic and tissue stress, best-practice
tation can be achieved if appropriate recovery time and an recommendations involve 48–72 h recovery between high-
additional bone-centric stimulus is provided [6]. With such intensity sprinting exposures [12, 13].
diverse tissue responses to an exercise stimulus, practitioners
might wonder if “optimal” loading is even possible. If tissue 2.1 Physiological Responses to Exercise
responses to the same stimulus vary so greatly, is it possible and Training
that practitioners might be optimally loading one tissue, but
suboptimally loading another? [7] 2.1.1 Acute and Chronic Adaptations to Training Stress
In this narrative review, we discuss the “typical” tissue
response to loading schema and demonstrate the complex The acute and chronic adaptations to training stress depend
interaction among training intensity, volume, and frequency. on the type, volume, and intensity of exercise performed
Training recommendations for specific tissues and systems [14]. Neuromuscular adaptations predominantly occur in
are provided to assist practitioners in the development of response to strength and power training, while endurance
rehabilitation and performance programs. We searched the training involves both neuromuscular and cardiovascular
PubMed database, using combinations of the following key- adaptations. Classic endurance training results in enhanced
words: (i) “training”, “load”, “exercise”, “intensity”, “vol- stroke volume and cardiac output, maximal oxygen uptake
ume”, and “frequency”; (ii) “athlete” and “patient”; (iii) (V̇ O2max), and mitochondrial biogenesis [15]. In contrast,
“tissue”, “muscle”, “tendon”, “bone” and “cartilage”; (iv) strength training results in increased muscle cross-sectional
“response” and “recovery”. Individual systematic reviews area, enhanced motor unit recruitment and synchronization,
and meta-analyses were hand-checked to identify any studies and improved maximal force production [15]. Although
missed during the electronic database search, and only Eng- exercise is commonly separated into long-duration, low-
lish-language peer-reviewed publications involving human intensity exercise (involving the generation of low forces,
subjects were included. predominantly aerobic energy release, and low central nerv-
ous system activation) and short-duration, high-intensity
exercise (involving the generation of high forces, predomi-
2 Part 1: From Tissue to System—Different nantly anaerobic energy release, and high central nervous
Loads Create Different Responses system activation), energy systems function in an integrated
manner, with very few activities deriving energy from any
Similar to tissue adaptation, the development of specific single system [16]. Equally, high-intensity training (in the
systems requires targeted exercise stimuli. Training load form of strength training, high-intensity intermittent train-
that lacks sufficient intensity will be inadequate to stress ing, and sprint-interval training) has been shown to improve
the anaerobic systems and engage fast-twitch muscle fibers endurance performance [17, 18], while low-load strength
[8–10], whereas frequent, higher volumes of intense train- training that approaches failure may lead to strength adapta-
ing may compromise recovery and/or aerobic adaptations tions [19].
[11]. It is well established that the development of sprint
running performance is dependent on appropriate exposure 2.1.1.1 Endurance Training Endurance training results in
to high-intensity sprinting, although sprint training activities cardiovascular and musculoskeletal adaptations that sup-
can range from low (e.g., tempo runs at 60–70% maximal port performance during prolonged aerobic exercise. The
velocity) through high (e.g., resisted sprints, speed endur- increased mitochondrial biogenesis and capillary density
ance, acceleration and maximal velocity at 80–98% maxi- that occur with endurance training result in improved use
mal velocity) and supramaximal (e.g., assisted sprints, up to of oxygen to generate energy [20]. These adaptations, along
105% maximal velocity) efforts [12]. Further complicating with changes in maximal cardiac output (driven predomi-
the load–response relationship for system development is nantly by a higher stroke volume) [21], result in greater oxy-
that underlying physical qualities (e.g., strength and power) gen delivery to the working muscles and increases in exer-
are often targeted to enhance sport-specific qualities (e.g., cise capacity and performance [22].
Optimal Loading for Tissues and Systems

Although prolonged low-intensity exercise has predomi- and greater fatigue and muscle soreness in the 48 h post-
nantly been used to elicit aerobic adaptations, more recently, exercise. Finally, Nuuttila et al. [31] compared physiological,
sprint-interval training (~ 30 s maximal bouts) and high- perceptual, and performance responses to a 2-week block
intensity interval training (~ 1–4 min near-maximal bouts) of either high-intensity interval training (ten sessions) or
have been employed as a time-efficient way of improving low-intensity training in recreational runners. Both training
endurance [15, 23]. High-intensity interval training has been interventions elicited similar improvements in 3000-m run-
shown to improve exercise capacity through adaptations in ning performance, although the high-intensity interval train-
mitochondrial content and respiration after only 2 weeks of ing was associated with greater muscle soreness, reduced
training [24]. Furthermore, skeletal muscle oxidative capac- parasympathetic activity, and elevated sympathetic activ-
ity and mitochondrial adaptations have been reported to be ity. Collectively, these findings suggest that (1) the fatigue
similar [25] or, in some cases, greater [26] following sprint- response to high-intensity interval training is comparable to
interval training than prolonged low-intensity exercise and that to continuous exercise, as long as total session volume
high-intensity interval training. Recent evidence suggests is low, (2) maximal effort intervals (i.e., sprint intervals) are
that high-intensity training (i.e., sprint-interval training and associated with longer recovery timeframes than high-inten-
high-intensity interval training) is mostly responsible for sity intervals performed at lower intensities, (3) perceptual
mitochondrial respiration and function adaptations, while recovery may be prolonged in older athletes, and (4) while
prolonged low-intensity exercise increases skeletal muscle regular high-intensity interval training can be tolerated in
mitochondrial content [15]. the short term, adequate recovery is required between high-
Increases in creatine kinase, reductions in countermove- intensity interval sessions to optimize adaptations.
ment jump performance, and delayed-onset muscle soreness
occur in the 24 h following high-intensity interval exercise 2.1.1.2 Strength and Power Training In the early stages of
[27]. Several researchers have investigated the effect of the a strength training program, improvements in strength are
high-intensity interval protocol on fatigue responses. Farias predominantly elicited through neural adaptations [32].
Junior et al. [28] assessed pressure-pain threshold, pressure- These adaptations include enhanced coordination, motor
pain tolerance, and perceived pain intensity in the rectus unit recruitment and synchronization [15]. Early unilateral
femoris, biceps femoris, and gastrocnemius before and 24 h limb strength training studies demonstrated the role of the
after either continuous exercise (20 min at 60% of maximal central nervous system in strength adaptations [33]; ~ 8%
aerobic speed) or low-volume high-intensity interval exer- improvements in strength were observed in the untrained
cise (10 × 60 s at 90% of maximal aerobic speed interspersed contralateral limb, despite unchanged cross-sectional area
with 60 s of active recovery), in untrained men. Both con- [34]. This cross-transfer of strength, which is thought to be
tinuous and high-intensity interval exercise elicited delayed- driven in part by localized muscle adaptations, cross-limb
onset muscle soreness, but no differences were observed cortical interaction, and adaptations in spinal cord excit-
between conditions. Wifison Alves et al. [29] compared the ability [34] is greater in eccentric (~ 47%) than concentric
fatigue response with two low-volume high-intensity inter- (~ 28%) training [35].
val sessions (10 × 60 s versus 20 × 30 s, both performed Morphological changes contribute more to longer-term
at 100% of maximal aerobic speed). No differences were improvements in strength [15, 36]. The force-generating
found between conditions for muscle soreness and pain in capacity of a single muscle fiber is directly proportional to
the 24–48 h post-exercise. Wiewelhove et al. [27] reported its cross-sectional area [36, 37]. Architectural factors such
greater fatigue (greater creatine kinase and delayed-onset as fascicle length and pennation angle also influence force-
muscle soreness, and larger reductions in countermovement generating capacity; longer fascicles allow greater force to
jump height) in the 24 h following exercise in team sport be produced, optimizing the length–tension relationship
athletes undergoing a sprint-interval protocol (4 × 6 × 5 s [38]. Employing a range of motion that emphasizes training
maximal-effort sprints) than following high-intensity inter- at long muscle lengths, and a repetition range between 2
vals (ranging from 80 to 95% of velocity achieved in the and 8 s, is recommended for muscle hypertrophy adapta-
30–15 Intermittent Fitness Test). Borges et al. [30] com- tions [39].
pared the recovery profiles of young (mean age 25.9 years) In a recent systematic review and meta-analysis, Sch-
and Masters (mean age 55.6 years) cyclists with a single oenfeld et al. [19] examined the effect of low-load (≤ 60%
high-intensity exercise session (6 × 30 s efforts at 175% peak one-repetition maximum, 1RM) and high-load (> 60%
power output with 4.5 min rest between efforts). No signifi- 1RM) training on strength and hypertrophy adaptations.
cant group × time interactions were observed for maximal Greater improvement in 1RM (effect size 1.69 versus 1.32)
voluntary contraction, 10 s sprint, 30 min time trial per- was observed with high loads, while changes in isometric
formance, or creatine kinase, although perceptual recovery strength and muscle mass were comparable between high
was reduced in the Masters athletes, with lower motivation and low loads. These findings challenge the notion that
 T. J. Gabbett, E. Oetter

an optimum repetition range exists for adaptations in lean 20-m sprint, countermovement jump, and isometric mid-
muscle mass [40], and that both hypertrophy and isometric thigh pull performance returned to baseline values within
strength can occur across a range of loads and repetition 24 h in the subjects who performed isometric exercise, per-
ranges. formance was still depressed following 24 h of recovery fol-
lowing heavy resistance training. Following an acute bout,
2.1.2 Muscle Adaptations eccentric exercise is associated with greater strength loss
[47], muscle damage, and soreness [48] than concentric-only
Eccentric strength training is commonly used to elicit mus- exercise. Although the muscle soreness following eccentric
cle architectural adaptations in athletes that are thought to be and isometric exercise is comparable, the reductions in
beneficial for both performance and injury prevention [41]. strength persist for longer following eccentric exercise [47].
For example, 12 sessions of eccentric hamstring training Following severe bouts of eccentric exercise, muscle damage
(using the Nordic hamstring exercise) resulted in increased biomarkers (e.g., creatine kinase, myoglobin) are elevated
fascicle length of the biceps femoris long head, along with for up to 8 days [49], while (depending on the protocol)
improvements in increased concentric and eccentric knee athlete-reported perceptions of soreness can remain elevated
flexion peak torque at 60°/s, concentric peak torque at above baseline for 3 days [50, 51]. Responses to eccentric
180°/s, and increased isometric knee flexion peak torque exercise follow a dose–response relationship, with low
[41]. The effects of different strength and power training volumes associated with less pain [52], less muscle dam-
interventions (e.g., isometric, isolated eccentric, or sprint- age [49], and poorer strength adaptations [53] than higher
ing) on muscle architecture and performance have been volumes.
studied, although results are equivocal. Sancese et al. [42] Rapid eccentric activity, typical of high-volume sprint-
reported improvements in isokinetic strength with both Nor- ing sessions, is associated with reductions in sprint perfor-
dic hamstring exercise and sprint training performed across mance and increases in hamstring strain injury risk factors
eight sessions, although no changes in sprint performance [50, 54, 55]. Carmona et al. [54] showed that an acute,
or mechanics were observed in either group. Timmins et al. high-volume sprint dose (10 × 40-m maximal efforts, per-
[43] reported increases in biceps femoris long head fascicle formed every 3 min) induced decrements in subsequent
length, thickness, and eccentric strength in response to either maximum sprinting performance (acceleration, maximum
isometric or eccentric (Nordic hamstring exercise) training velocity, and horizontal and vertical force) that persisted
across a 38-week football season in semi-professional ath- for 48–72 h. Reductions in posterior chain muscle strength,
letes. Improvements in acceleration and maximum velocity and lumbo-pelvic control, coupled with increases in muscle
qualities occurred in the eccentric training group but not soreness and creatine kinase also occurred across this time
in the isometric training group [43]. Freeman et al. [44] period [54, 55]. Others have shown that residual fatigue (i.e.,
reported improvements in eccentric hamstring strength in increased muscle damage and delayed-onset muscle soreness
athletes who underwent either Nordic hamstring exercise and reduced well-being, hamstring force production capac-
or sprint training, but improvements in sprint performance ity, and physical performance) persists for > 72 h following
were greater in the sprint training group. Equally, 6 weeks team sport competition [56–59].
of sprint training was associated with greater increases in
biceps femoris long head fascicle length, sprint performance,
and mechanics than isolated hamstring training using the 2.1.3 Tendon Adaptations
Nordic hamstring exercise [45]. Although there is conflicting
evidence, these results demonstrate the muscle architectural Tendons are sensitive to their mechanical environment [60].
adaptations that occur through both slow (e.g., Nordic ham- Mechanical loading of tendon tissue results in upregulation
string exercise) and rapid (e.g., sprinting) eccentric activity; of collagen expression and increased synthesis of collagen
these changes are beneficial for mitigating hamstring injury protein; the extent of protein synthesis is likely a function
risk and improving performance. of the strain experienced by tenocytes [61, 62]. Collagen
Lum and Howatson [46] studied the acute effects of a sin- synthesis peaks ~ 24 h after exercise. Collagen degradation
gle session of either isometric strength training (back squat also occurs post-exercise, but peaks earlier than collagen
at four different knee angles, and Romanian deadlift and synthesis [61].
split squat at three different knee angles) or heavy resistance Following a period of mechanical loading, tendon stiff-
training (back squat at 85% 1RM, split squat at 40% 1RM ness increases owing to changes in both tendon material and
squat, and Romanian deadlift at 95% 1RM squat) on subse- morphological properties [60, 63]. These changes in tendon
quent 20-m sprint, countermovement jump, and isometric properties help the storage and return of strain energy dur-
mid-thigh pull performance. Performance was reduced in ing locomotor (e.g., walking) and athletic (e.g., sprinting,
both groups immediately following training. However, while jumping) tasks. A wide range of training interventions have
Optimal Loading for Tissues and Systems

been employed to increase the mechanical load on tendons, relationship reported between the magnitude of tissue stress/
including varying training intensity, sets, repetitions, train- strain and bone fatigue. Small increases in stress and strain
ing frequency per week, duration of a single loading cycle, result in large decreases in the number of bone cycles that
and duration of the training intervention [60]. Of these, con- can be performed before failure [7]. Weaker bones have
traction intensity and the duration of the training interven- lower stress and strain tolerance. Enhancing tolerance to
tion appear to be most important. In their 2015 systematic bone stress requires the introduction of mechanical stress to
review and meta-analysis, Bohm et al. [60] showed that the promote skeletal adaptation and ensuring that the microda-
effect sizes for changes in tendon stiffness were significantly mage that occurs as a result of cumulative loading does not
greater (effect size 0.90 versus 0.04) when training with outweigh the ability of the bone to repair itself.
higher muscle contraction intensities (> 70% of maximal The adaptations that occur in bone differ between endur-
voluntary contraction or 1RM) than with lower contraction ance and strength training. For example, endurance runners
intensities. Longer intervention periods (≥ 12 weeks) are who participated only in their primary sport had lower bone
considered more effective at eliciting tendon adaptations density than those who also engaged in heavy strength train-
than shorter interventions, although there is evidence that ing [72]. Equally, retrospective studies have shown that run-
tendons respond positively to mechanical loading within ners [73] and military cadets [74] who did not participate
2 months [60]. The type of muscle contraction (i.e., iso- in regular strength training were at increased risk of bone
metric, concentric–eccentric, or eccentric only) has minimal stress injury than habitually strength-trained participants.
effect on tendon adaptations [60]. Multidirectional jumping, landing, and change-of-direction
It should be noted that, although contraction intensity and activities are recommended to build a robust skeleton [7].
intervention duration appear to be the most important factors Conversely, the unidirectional nature of running is not effec-
in changing the mechanical, material, and morphological tive for building bone strength [7]. Bone cells desensitize to
properties of tendons, loading duration [64], frequency [65] repetitive loading, with 95% of mechanosensitivity lost after
and rate [64], joint angle [66], and repetitive versus static as few as 20 loading cycles [5]. These findings have impor-
loading [64] are also important loading conditions that have tant implications for practitioners as they suggest that, after
been shown to influence tendon adaptations. this mechanosensitivity is lost, further bone loading does
Healthy tendons respond well to consistent (i.e., daily) not produce proportional increases in bone adaptation. How-
loading as long as stretch–shorten cycle activity and tendon ever, a 4–8 h refractory period restores mechanosensitivity
strain are minimized [67]. Net loss of collagen may occur if and allows for a second small dose (≤ 60 loading cycles)
less than 24 h recovery is provided between exercise bouts, of bone-centric exercise (e.g., plyometrics), potentially
leaving the tendon vulnerable to injury [61]. For patients maximizing osteogenesis and adaptive potential in a given
with tendinopathy, less frequent loading is recommended timeframe [7]. Although rehabilitation can be accelerated
[68]. For example, hyperhydrated and reactive tendons that for some specific tissue injuries (e.g., muscle strains) [75],
have undergone high stretch–shorten cycle activity benefit rehabilitation from bone stress injuries requires practitioners
from a 48-h refractory period before receiving a similar to respect a cautious and gradual recovery timeframe. For
training dose. Heavy loads, including the use of isometric example, athletes with bone stress injuries are advised to
exercise, have been shown to induce beneficial changes in be pain-free during daily activities for 5 consecutive days
tendon architecture, while also reducing pain in patients with before commencing return-to-run programs [76].
tendinopathy [68–71].
2.1.5 Cartilage Adaptations
2.1.4 Bone Adaptations
Although cartilage has the ability to deform and regain
Microdamage that occurs through repetitive loading is a shape under the application and removal of forces [37],
necessary and normal component of bone adaptation. This the effects of mechanical loading on cartilage regulation
microdamage results in bone-resorbing osteoclasts remov- are unclear. Mechanical loading and cartilage thickness
ing damaged regions of bone and osteoblasts forming new, dose–response studies are limited, but it is generally rec-
undamaged bone [7]. In cortical bone, it may take up to ognized that adaptations of human cartilage to exercise are
4 weeks for osteoclast activation and resorption to occur, not linear [77]. Immobilization is associated with reduced
while full mineralization of new bone may take between 3 cartilage thickness [78] while lower thigh muscle strength
and 12 months. This process is longer in trabecular bone [7]. [79] and cross-sectional area [80] are associated with poorer
Skeletal microdamage is dependent on both the number cartilage morphology and more severe osteoarthritis after
of times a bone is loaded and the magnitude (i.e., inten- partial meniscectomy. In addition, in patients with osteoar-
sity) and the rate at which the load is applied [7]. Bone thritis, lower pain and disability symptoms were reported in
fatigue is most influenced by load magnitude, with a strong those undergoing quadriceps-specific exercise than in those
 T. J. Gabbett, E. Oetter

engaging in general lower limb exercises [81]. Cartilage has 3 Part 2: Programming Considerations
been shown to rapidly recover following mechanical load- for Practitioners
ing [82–84]. Van Ginckel and Witvrouw [85] assessed the
deformation of knee cartilage in response to squatting exer- 3.1 A Conceptual Framework for Determining
cise in individuals with mild tibiofemoral osteoarthritis and the Appropriate Response to Loading
healthy controls. Reductions in cartilage volume occurred
in both groups but returned to baseline within 15 min. No Given the wide range of tissue responses to a given load,
significant between-group differences were found for volume how can practitioners provide practical, evidence-based
changes. Similar findings have been shown for talar cartilage training interventions to their patients and athletes? Based
volumes following squatting exercise; cartilage deformation on the underpinning physiological responses and adapta-
occurred in response to loading but was restored to baseline tions that occur during acute and chronic exercise, and
within 30 min [86]. Beneficial adaptations that may occur the expected load–response, several recommendations for
in response to mechanical loading of cartilage tissue include practitioners responsible for rehabilitation and performance
stiffening of the pericellular and interterritorial matrix [87], training programs are provided below.
increased cartilage volume and concentration of glycosa-
minoglycans [88], and reduced cartilage loss [89]. Collec- 3.2 Prioritizing Tissue and System Loading
tively, these findings suggest that strength training is a safe for Healthy and Injured Athletes
method of loading articular cartilage and plays a critical role
in maintaining joint integrity and health. We have recently presented a training model that describes
Cartilage is an extremely resilient tissue that can with- the relative emphasis of local-tissue and sport-specific
stand large loads. Studies investigating the influence of loading for healthy and injured athletes [97]. The loading
exercise intensity on joint changes are equivocal, with some emphasis for injured athletes is on restoring local-tissue
studies showing no effect [90] and others reporting greater capacity, while in healthy athletes, resources are predomi-
deformation [91] with higher intensities. Harkey et al. [90, nantly directed toward sport-specific loading. However,
91] investigated deformation of the medial femoral carti- an overemphasis on either approach can result in athletes
lage in response to walking, running, and drop landing in with poor local tissue health (in athletes who overempha-
healthy individuals. No differences in deformation were size sport-specific loading) or poor sport-specific capacity
reported between walking (− 6.7%) and running (− 8.9%) (in athletes who overemphasize local-specific loading). It is
[90], although cartilage deformation after the drop-landing widely suggested that successful rehabilitation and perfor-
task required longer time to recover compared with the walk- mance programs employ a combination of local-tissue and
ing condition [91]. For example, ultra-endurance marathon sport-specific loading [97, 98].
runners experienced collagen disorientation and a decrease A challenge for practitioners is maintaining the function
in the concentration of glycosaminoglycans in the femoroti- of healthy tissues in the presence of an injury to a specific
bial joint within the first 1100 km of the race, but no further tissue type. In injured athletes, practitioners should prior-
changes in the subsequent 3500 km [92]. Equally, no new itize reloading of the vulnerable tissue while considering
osteochondral lesions were found and no further softening adaptive, supportive loading strategies for noncompromised
of the cartilage occurred following completion of the race tissues. When returning an athlete from bone stress injury, a
[92]. Although some studies have shown a greater preva- more conservative loading approach may be required, even
lence of hip and knee osteoarthritis in professional runners in the presence of healthy muscle or connective tissue [76].
(13.3%) and sedentary individuals (10.2%) than in recrea- However, there are evidence-based approaches that permit
tional runners (3.5%), it was not possible to determine if practitioners to “train around” the load-compromised bone.
these associations were causative or confounded by other For example, the liberal use of blood flow restriction train-
risk factors, such as previous injury [93]. There is currently ing offers an ideal modality to maintain muscle adaptations
no strong evidence that strenuous exercise increases the risk [99] while minimizing joint and bone stress [100, 101]. The
of osteoarthritis in healthy joints [94–96]. effects of blood flow restriction training on the mechanical
properties of tendon (e.g., tendon stiffness) are equivocal,
with some studies demonstrating positive connective tissue
adaptations with low-load resistance training [102, 103] and
others [104] showing no effect. In the presence of muscle
injury, the health of connective tissue can be maintained
through the use of short-to-long muscle length isometric
exercise and eccentric training that employs progressive
Optimal Loading for Tissues and Systems

increases in force, speed, and time under tension. This pro- insulin-like growth factor-1 (IGF-1), walking speed, and
gression in eccentric activity incrementally prepares both the strength in patients with multiple sclerosis compared with
muscles and tendons for the higher-intensity stretch–shorten control subjects [117]. Case studies of elite soccer [120]
cycle activity demands (e.g., sprinting, jumping, changes of and rugby [121] players have also shown the importance of
direction) required on return to competition. maintaining systemic loading to prevent reductions in lean
muscle mass, particularly during the early stages of injury.
3.3 If Unloading is Necessary, Maintain Global Load This maintenance of strength and skeletal muscle [122] is
Capacity driven in part through activation of the IGF-1 signaling
pathway, which promotes myogenesis and inhibits protein
Although load capacity is improved through graded exposure degradation and cell death [119]. Furthermore, exercise has
to training stress [105–107], there may be occasions where been associated with increases in brain-derived neurotrophic
training loads need to be regressed before sensibly progress- factor (BDNF), a neurotransmitter modulator that plays an
ing [108]. In these instances, practitioners are encouraged important role in neuroplasticity, learning, and memory
to maintain global loading to minimize the systemic effects [123] as well as minimizing sleep and mood disturbances
of detraining and moderate the risk of reinjury upon rein- [124]. Given the importance of sleep to recovery processes
troducing sport-specific activities. The protective effect of [125, 126], and the need to reintegrate injured athletes into
high global capacity has recently been demonstrated in soc- technical and tactical training post-injury [127], the increase
cer players; players with higher chronic internal loads (as in BDNF through exercise appears to play a critical role in
estimated using the session rating of perceived exertion) had the return-to-play process.
reduced risk of injury at any given high-speed running vol-
ume compared with players with low chronic loads [109]. 3.4 Consider Recovery Timeframes of Different
The protective effect of training has also been shown by Tissues and Systems When Determining
other researchers. In men’s professional soccer, players were the “Normal” Response to Load
87% more likely to be injured in the first match after return
to play than the seasonal average injury rate. However, for Inter- and within-individual variability notwithstanding, the
each additional training session performed before returning “typical” timeframes of recovery following training for dif-
to play, the risk of muscle injury was decreased by 13% ferent tissues and systems are relatively predictable and are
[110]. Similar findings have been documented in Australian shown as a conceptual framework in Fig. 1. After the initial
Rules football players; higher training loads (in terms of training dose, performance is reduced, followed by a return
both more sessions and accumulation of greater high-speed to performance at or above (i.e., supercompensation) the
running chronic loads) were associated with lower risk of initial levels [128, 129]. While it is well established that the
reinjury on return to competition [111]. intensity of load prolongs these recovery times, an under-
The majority of research has employed laboratory-based standing of the normal response to load allows practitioners
experimental designs using limb immobilization to study to assess if the athlete has recovered from the training dose
the effects of disuse. These studies have shown that healthy, and is ready to receive a subsequent training dose. For exam-
inactive muscle tissue atrophies at ~ 0.5% per day [112, ple, maximum-force isometric muscle contractions are asso-
113] with the greatest loss of muscle mass occurring in the ciated with low fatigue and rapid recovery [46]. Compared
initial 1–2 weeks of inactivity [114]. Systemic loading is with heavy strength training, maximum-force isometric
commonly used during return-to-play protocols to comple- muscle training results in smaller decrements in speed and
ment local tissue loading. Along with the maintenance of isometric mid-thigh pull performance, and greater perceived
tissue architecture (e.g., collagen turnover in tendons and recovery in the 24-h post-training window [46]. Given the
muscles) [62, 115] and neurophysiological performance postactivation performance-enhancing effects [130], and the
determinants, the metabolic stress associated with systemic rapid recovery following maximum-force isometric muscle
loading promotes adaptive hormonal secretions that can contractions [46], this type of loading should be encour-
support tissue healing [116]. The benefits of systemic load- aged when training for sport-specific training adaptations.
ing to prevent atrophy and strength losses associated with Conversely, sprinting elicits neuromuscular fatigue that is
disuse can be found in studies encompassing a wide range accompanied by a prolonged decrease in hamstring strength
of conditions and populations [117–119]. For example, in [55]. These reductions in hamstring strength are thought to
comparison with control patients, a combination of aerobic be driven by both central and peripheral mechanisms (e.g.,
and strength training improved grip strength and walking neuromuscular fatigue, muscle damage), which impair
speed in patients following hospitalization with myocardial contractile function, but are restored within 48–72 h [12].
infarction [118]. Similarly, a combination of aerobic and Importantly, for both muscles and tendons, low-intensity
light resistance training resulted in significant increases in activity can be performed, and is even recommended, during
 T. J. Gabbett, E. Oetter

Fig. 1  Normal time course of recovery from loading bouts for differ- enced by health and training status, with many health conditions asso-
ent tissues and systems. The response of specific tissues and systems ciated with compromised tissue responses to training load. The time
to training load is complex; at any time, it is possible that practition- course for adaptations of specific tissues and systems is based on the
ers may be optimally loading one tissue or system while suboptimally following references [4, 6, 7, 12, 27–29, 46–51, 54–58, 76, 90, 91].
loading another. Note that the response to exercise stress will be influ- CNS central nervous system

the recovery window. Further evidence for the influence of reductions in well-being [56, 138–140] and neuromuscular
intensity on recovery can be found from cartilage deforma- function [56, 139]. Notwithstanding the wide range of perio-
tion [91], and tendon [60] and bone-loading [7] studies. dization strategies employed by coaches [141], due to the
Although cartilage recovery occurs rapidly (i.e., within fatiguing nature of high-intensity exercise, relatively small
30 min) following walking and running, longer recovery amounts of training time are devoted to these activities, even
periods (i.e., ≥ 45 min for cartilage thickness and cross- in well-designed programs. For example, in track sprinters,
sectional area) may be required following higher-intensity athletes typically only perform two high-intensity sprint ses-
drop-landing tasks [91]. Equally, even small increases in sions per week and keep high-intensity training volume to
bone stress and strain significantly reduce the number of a minimum (e.g., 50–150 m of maximum velocity sprinting
bone cycles that can be tolerated before failure [7], indicat- per session) [12]. Equally, despite the need to perform at
ing the need for greater recovery following high-intensity high intensities during competition, endurance runners and
bone loading. cyclists (for events lasting approximately ≥ 2 min) devote
most of their training time toward low-intensity training,
3.5 Consider Recovery Timeframes When with comparatively lower proportions (≤ 20%) performed
Implementing High‑Intensity Training at high intensities [11, 142]. This ensures (1) the allocation
of sufficient training time to develop an aerobic capacity
Research has shown that high-intensity activity occurs at capable of supporting high-intensity activity, (2) the con-
critical moments of competition (e.g., goals scored or con- solidation of “high stress” training activities, (3) adequate
ceded, shots on goal) [131–133]. These findings provide sup- recovery time between high-intensity sessions to promote
port for the importance of high-intensity training for most training adaptations, and (4) that any high-intensity exercise
sports. Equally, performance in individual sports requires performed is at an adequate intensity to prepare the tissues
athletes to maintain high speeds and power outputs over a for the demanding nature of competition. This polarized
prolonged period of time. The production and maintenance approach to training is not limited to individual or endurance
of these high-intensity efforts is extremely fatiguing, result- sport athletes; team sports (such as American Football and
ing in both transient (60 min post-exercise) and longer-term elite soccer) will also use polarized training models to ade-
(24–48 h) increases in muscle damage [56, 134–137], and quately prepare athletes for match play [143, 144], although
Optimal Loading for Tissues and Systems

it is acknowledged that micro-dosing may be the preferred is some evidence from trained mountain bike cyclists that
programming strategy for sports with congested competition reducing the recovery period between high-intensity ses-
schedules [145]. Given that match day requires team sport sions has minimal effect on various performance measures.
athletes to perform highly demanding stretch–shorten cycle Hebisz et al. [150] compared 8 weeks of block training con-
activities, practitioners must provide adequate recovery time sisting of 17-day blocks of low-intensity training and 11-day
between high-intensity training activities and those of com- blocks of sprint-interval and high-intensity interval train-
petition. In this respect, the “lead-in” and “lead-out” time ing with polarized training that simultaneously consisted of
for match play are considered, with high-intensity training low- and high-intensity and sprint-interval training. While
exposures typically occurring in the 48–72 h before and after the polarized training program elicited greater improvements
competition [13, 146]. Examples of a polarized approach to in V̇ O2max, no differences were observed between the block
training for individual sport (e.g., 10,000-m endurance run- and polarized training programs for changes in maximal
ner) and team sport athletes (e.g., American Football) are aerobic power, or the power achieved at the first and second
provided in Tables 1 and 2, respectively. ventilatory thresholds.
Further evidence for the judicious use of high-intensity There is a large within- and inter-individual variation in
exercise within the training microcycle can be found in stud- response to loading [151, 152], with adaptation of most tis-
ies that have compared the inflammatory response to low-, sues (e.g., bone, tendon, and muscle) dependent on a range
moderate-, and high-intensity exercise [147, 148]. In a sys- of factors, including loading conditions (e.g., intensity, dura-
tematic review, Cerqueira et al. [147] found that the proin- tion of single loading cycle, training frequency, repetitions,
flammatory cytokine tumor necrosis factor alpha (TNF-α) sets, duration of training intervention), exercise conditions
only increased after high-intensity exercise, while interleu- (e.g., type of muscle contraction, joint angle, and whether
kin-6 (IL-6) and interleukin-1β (IL-1β) increased more with the stimulus is applied statically or dynamically) [60], and
high-intensity than with moderate-intensity exercise. Simi- even diurnal variation [153]. Of these factors, the intensity
lar findings have been found in patients undergoing regular of loading is a significant determinant of the time course
physical activity; physically active patients with gout had of recovery and degree of adaptation in tendons [60]. For
fewer flares per year, decreased C-reactive protein levels, example, tendon stiffness adaptations have been shown to
and less pain than patients who were physically inactive be greater in exercise protocols employing high intensities
[149]. While high-intensity training is important for per- (≥ 70% maximum voluntary contraction or 1RM) than in
formance outcomes, collectively these findings suggest that those using lower intensities [154, 155]. However, exces-
adequate time is required between sessions to allow tissue sive mechanical loading can contribute to tendinopathy [62],
recovery and training adaptations to occur. which is associated with activity-related pain, focal tendon
Despite most practitioners recommending at least 48 h tenderness, and decreased strength and flexibility [156].
between high-intensity sessions or competition [146], there

Table 1  An example of a polarized training microcycle for an elite individual sport athlete (e.g., 10,000-m runner)
Mon Tue Wed Thu Fri Sat Sun
AM Extensive Aerobic Intensive Anaerobic Mobility Tempo Run Mobility VO
̇ ₂ Intervals

e.g. 1-3 hour of e.g. (2x) 10x200m e.g. 1 hour of e.g. 8x1km
zone 2 running (holding <30s on 3 zone 3 running (holding ~2:30-2:40
(~75-85% of min cycle) (~80-90% of min on 7:30 min cycle)
maximum heart maximum heart
rate) rate)
PM LB Strength & Power Recovery Run FB Strength Extensive Aerobic LB Strength & Power Recovery Run
(optional)

e.g. high force, e.g. 1-5 hour of e.g. high force e.g. 1-3 hour of e.g. high force, e.g. 1-5 hour of
stretch-shorten cycle, zone 1 running activities zone 2 running stretch-shorten cycle, zone 1 running
and bone-centric (~55-75% of (~75-85% of and bone-centric (~55-75% of
activities maximum heart maximum heart activities maximum heart
rate) rate) rate)
Low Stress High Stress Low Stress Moderate Stress Low Stress High Stress Low Stress

NB: (1) accumulation of large volumes of low-intensity aerobic activity and small volumes of high-intensity anaerobic activity; (2) consolidation
of physiologically demanding activities on “high-stress” training days; (3) lower body strength and power sessions scheduled 4–8 h after running
activities to allow bone-centric activities (e.g., hopping, jumping, and landing) to be performed; (4) at least 72 h between “high-stress” training
days
LB lower body, UB upper body, FB full body
This training microcycle is an example only; several other effective programming strategies could be used as alternatives to this model
 T. J. Gabbett, E. Oetter

Table 2  An example of an in-season polarized training microcycle for an elite team sport athlete (e.g., American Football player)
Mon Tue Wed Thu Fri Sat Sun
AM Off UB Strength & Power LB Strength & Power FB Priming

e.g. incorporating high e.g. incorporating e.g. low-volume,

force (including high force (including high-intensity
isometric and isometric and resistance
eccentric exercises) eccentric exercises) exercise or low
and stretch-shorten and stretch-shorten load ballistic
cycle activities cycle activities exercise
PM Active Recovery Practice Practice Practice Walk Through Game Day

e.g. low volume & e.g. involving high e.g. involving high e.g. moderate i.e. final team
intensity, acceleration and velocity running intensity skill preparations and
movement focus deceleration activities, activities activities, no high “rehearsal”
physical contact, and velocity running before Game
minimal high velocity or physical
running contact
Low Stress Low Stress Moderate Stress High Stress Moderate Stress Low Stress High Stress
NB: (1) consolidation of physiologically demanding activities on “high-stress” training days; (2) at least 72 h between the “high-stress” activity
of game day and subsequent “high-stress” training activities; (3) at least 72 h between “high-stress” training activities and game day. An optional
low-intensity, high-volume hypertrophy session could be implemented toward the latter part of the week (likely 36 h before game day) in players
with lower lean muscle mass. Blood flow restriction exercise can also be used in joint-compromised players. Video reviews and team meetings
are not shown in the training microcycle
LB lower body, UB upper body, FB full body
This training microcycle is an example only; several other effective programming strategies could be used as alternatives to this model

Both intensity and frequency of resistance training con- 3.6 Interpreting Training Within an Athlete
tribute positively to muscular strength, whereas resistance Monitoring Framework Allows Practitioners
training volume contributes significantly to both muscular to Provide Context to Load–Response Data
strength and hypertrophy [157]. Typically, 48 h of recovery
is required between training sessions involving isotonic con- Throughout this paper, we have highlighted that “load”
tractions with similar muscle groups; however, the intensity and “response” data are more powerful when evaluated as
of training and contraction type will influence the timecourse a combined dataset; training loads can only be considered
of recovery [158]. Krentz and Farthing [158] demonstrated excessive or inadequate based on the response to that load.
that intense eccentric exercise (six sets of eight maximal Equally, the athlete’s “response” (e.g., soreness, fatigue,
repetitions) performed every 2 days resulted in incomplete mood, etc.) requires context of the desired training outcome;
repair of muscle damage and prolonged impairment of mus- fatigue and soreness in preseason when training loads are
cle strength. In a separate study, Bellosta-Lopez et al. [159] high might be considered an appropriate response. However,
investigated the time course of recovery in pain perception, a similar response might be considered undesirable leading
pressure pain sensitivity, active range of motion, maximal into the first game of a seven-game finals series. We have
isometric strength, and muscle activity of the hamstrings recently proposed an athlete monitoring model that captures
during a maximal isometric contraction following a single- load–response data, allowing practitioners to evaluate the
leg deadlift exercise (5 sets of 20 repetitions) designed to effectiveness of their training interventions and readiness
elicit delayed-onset muscle soreness. While muscle activ- of athletes to continue training [164, 165]. Although this
ity had recovered by day 4, pain perception, pressure pain model was designed with the “system” in mind, this athlete
sensitivity, maximal isometric strength, and active range of monitoring cycle, which encompasses external and internal
motion only returned to baseline on day 7. Given the het- load, subjective well-being, and objective measures of readi-
erogeneous time course of recovery to high-intensity muscle ness, can also be applied to tissue stress (Fig. 2). This ath-
contractions, practitioners should consider both subjective lete monitoring framework describes a step-by-step strategy
and objective measures when determining an athlete’s readi- for interpreting data, from the exposure to a single training
ness to perform subsequent training activities [160–163]. stimulus (or external load), through to the exposure of a sub-
sequent training stimulus [164]. When combined with each
previous step, practitioners can make informed decisions
on training prescription (e.g., whether to adjust the external
load during the session, identify if the athlete is tolerating
the prescribed training load, and whether to introduce extra
recovery or activation interventions to prepare the athlete
Optimal Loading for Tissues and Systems

External Load Monitoring Internal Load Monitoring

• Global Posi oning Systems • Heart Rate, Lactate, Oxygen
1 2
• Iner al Measurement Sensors (e.g. Consump on, Muscle Oxygen Satura on
accelerometers, gyroscopes, External Load • Crea ne Kinase, Cor sol, Testosterone,
magnetometers) Inflammatory Markers
What did the athlete do?
• Pedometers • Electromyography Ac vity
• Local Posi oning Systems (e.g., • Ra ng of Perceived Exer on
radio-frequency iden fica on, • Visual Analogue Pain Scales
ultra-wideband)
!
Training Decision
Manipulate external load during the
4 session, or in subsequent sessions, to
achieve desired internal load

Objecve Measures of Physical Readiness Subjecve Measures of Well-Being

• Res ng Heart Rate • Fa gue
• Heart Rate Variability • Sleep Quality
• Force and Power (e.g. eccentric, 3 • Muscle Soreness
isometric strength, counter-movement • Stress
jump) • Mood
• Shear Wave Elastography • Pain, Swelling, or Focal Tenderness
• Ultrasound Tissue Characteriza on

! !
Training Decision Training Decision
Con nue regular training or introduce
Con nue regular training or modify the
interven ons to increase recovery or
training program
ac va on

Fig. 2  An athlete monitoring framework that considers external and tem” in mind, this athlete monitoring cycle can also be applied to tis-
internal load, subjective athlete responses, and objective measures of sue stress. Redrawn with permission from Gabbett et al. [164].
training readiness. Although this model was designed with the “sys-

for subsequent sessions) [164]. Below, we describe some of acceptable validity for most (e.g., position, distance, and
the assessment methods and technologies available to prac- average speed), but not all (e.g., instantaneous speed), meas-
titioners to monitor external and internal training load, and ures of external load [168].
subjective and objective responses to that load. Global positioning systems also include inertial meas-
urement sensors (e.g., triaxial accelerometers, gyroscopes,
3.6.1 External Load Monitoring and magnetometers). This technology, which is worn on
the body (and in other apparatus such as mouthguards and
Predominantly employed by physical activity researchers, shoes), is now commonly used in a wide range of sports
pedometers have been used extensively to count the num- [173]. Accelerometers quantify accelerations on the body.
ber of steps an individual takes within a given time period The sum of the accelerations performed in three planes (and,
(usually a day). Traditionally worn on the hip, these step therefore, an estimate of the external forces acting on the
counters are now integrated into smartphones and watches. body) provides a measure of external biomechanical loads
Perhaps the most rudimentary of external load measures, [173]. While inertial measurement systems worn on the
research has shown strong associations between pedom- upper back or sacrum can be used to provide an estimate of
eters and accelerometers (r = 0.86) and measures of energy global external load, most practitioners working with run-
expenditure (r = 0.68), with accuracy better during walking ning-, jumping-, and landing-based activities opt for a sensor
and running speeds than during slow walking [166]. worn on the shank or the insole of the shoe [174]. These
Global positioning systems are commonly used to meas- sensors provide a closer approximation of ground reaction
ure external loads (e.g., total distance, high-speed running, forces experienced in the lower body while also providing
sprinting, accelerations, and decelerations) in outdoor important information on gait deviations, ground contact
sports. However, given that this technology relies on satel- times, and stride changes. Presently, these devices provide
lites, quantifying the locomotor demands of indoor sports the best available method of estimating external tissue loads
has predominantly been performed using labor-intensive when returning athletes from lower-limb complaints, such
video-based time–motion analysis [167]. Recently, local as bone stress injuries and Achilles tendon ruptures. An
positioning systems (e.g., radio-frequency identification added advantage of the inertial sensors is that, when the
and ultra-wideband) have been used to quantify the exter- obtained data are combined with tailored algorithms, there
nal loads of athletes from indoor sports [168–172], with is potential to quantify sport-specific external loads [175].
 T. J. Gabbett, E. Oetter

In a systematic review of athlete-mounted inertial sensors, 3.6.4 Objective Measures of Physical Readiness

Chambers et al. [175] highlighted the use of this technology
to detect sport-specific movements (e.g., throwing, tackling, Finally, a myriad of different technologies can be used to
fast bowling, swimming stroke classification, tennis serves, provide objective data on the readiness of a tissue to receive
and snowboarding aerial maneuvers) from a wide range of a subsequent loading dose. For example, using isokinetic
individual, team, water, and snow sports. This is particularly dynamometers, force plates, and velocity-based training sys-
important in sports where a large proportion of the external tems, kinetic outputs can be used to probe the load tolerance
load arises from activities other than locomotion (i.e., high- and capacity of specific tissues during constrained isoki-
speed running and sprinting). netic and/or isotonic actions (e.g., leg extension, counter-
movement jump, bench throw). Shear wave elastography
[187] and ultrasound tissue characterization [4] are methods
3.6.2 Internal Load Monitoring to monitor tendon stiffness and morphology, respectively.
Although dependent on access and likely impractical for
While the measurement of external loads is relatively daily use, these assays can be employed on a longer-term
straightforward, musculoskeletal modeling, which involves (e.g., 4-week) basis to gain feedback on intervention efficacy
measurement of joint contact forces or muscle–tendon and monitor architectural changes that may be associated
forces, is required to monitor the internal load of musculo- with tendinopathy. In the near future, it will be possible to
skeletal tissues [173]. These direct measurements of internal use marker-less kinematic technologies to help impute spe-
tissue stress are difficult outside the laboratory environment, cific loads incurred by tissue during sport movements [188,
although surrogate measures can be used. For example, 189].
recent studies have proposed the use of wearable electro-
myography (EMG) technology to quantify internal loads and
fatigue during exercise [176–179] and rehabilitation [180]. 4 Advancement of Load–Response Research
In comparison with laboratory-based EMG measures, wear- and Practice
able EMG technology offers a valid method of monitoring
a wide range of activities including orthostatic challenges 4.1 Recommendations for Researchers
[177], isometric exercise [177], treadmill running [181], and
incremental cycle exercise to exhaustion [182]. This technol- Although research has contributed to the understanding
ogy, which is located in a small (30 mm) patch worn on the of recovery timeframes for specific tissues, there are sev-
skin surface or wearable shorts, is also sensitive to changes eral gaps that could be addressed to improve rehabilitation
in exercise intensity [181, 182]. Others have used muscle and training programs. Firstly, most [29, 30, 46, 48, 50,
oxygen saturation as a measure of internal load [183]. The 54–56, 58, 135, 194], but not all [49, 51, 159], research
rating of perceived exertion (RPE) scale is the most com- has quantified the load–response over a short (i.e., usually
mon method of monitoring global internal load of athletes. 24–72 h) recovery period following loading, with at least
It is likely that some modification of the RPE scale or the some variables (e.g., sprint performance, biceps femoris
use of differential RPE for leg muscle exertion is required to long head muscle fiber structure, lumbo-pelvic control) not
adequately capture internal local tissue loads [97]. fully recovered 72 h after activities involving high-volume
sprinting [54, 55], Therefore, it is not possible to state with
3.6.3 Subjective Well‑Being Measures certainty the exact duration of recovery required follow-
ing specific loading for some tissues (e.g., rapid eccentric
Athlete-reported measures of soreness (e.g., muscle), mood muscle contractions). Studying the response of specific
disturbances (e.g., vigor, fatigue, stress, energy, recovery), tissues for longer post-loading windows is warranted. Sec-
or pain (e.g., bone and tendon) are by far the most com- ondly, given that all training programs will have a pro-
mon approach to assess tissue tolerance to training load portion of “responders” and “stubborn responders,” [190]
[184]. These measures of subjective well-being are typi- and that positive training adaptations can still be made
cally impaired with an acute increase in training load, with with shorter recovery periods [145], identifying a “rap-
acute decreases in training load associated with improved idly adapting” athlete is of interest. If a subset of athletes
subjective well-being [184]. Visual analog scales have been can indeed be identified as “rapid adapters,” understand-
used to assess tendon pain [185]. Bone tissue damage can ing the biopsychosocial characteristics that predict rapid
be estimated from excessive focal tenderness, swelling, pain adaptation would allow very specific training interventions
at rest and on weight-bearing, and pain that increases with to be introduced to develop this trait. Finally, although
physical activity and fails to subside on cessation of physical technological advancements have presented opportunities
activity [186]. to measure a wide range of training loads and responses,
Optimal Loading for Tissues and Systems

many internal tissue loads cannot be directly measured to suggest that the time course of recovery may be shorter in
outside the laboratory environment [173]. Furthermore, individuals with better developed physical qualities [135, 194].
not all wearable devices are accompanied by software to
process the raw signals (thereby requiring programming 6 Conclusion
skills to process, filter, and interpret the acquired data),
and the cost associated with some technologies makes the In summary, this paper describes the “normal” tissue response
uptake of these devices beyond the reach of most clinics to training load and demonstrates the complex interaction
[179]. A challenge for researchers (and technology compa- among training intensity, volume, and frequency, as well as
nies) is to develop valid and cost-effective athlete monitor- the impact of these training variables on the recovery of spe-
ing devices, capable of capturing the load–response, while cific tissues and systems. The response of specific tissues and
also preserving the utility of the product. systems to training load is complex and is dependent on the
magnitude of the training dose. The consideration of recovery
timeframes of different tissues and systems allows practition-
4.2 Recommendations for Practitioners ers to determine the “normal” response to load. We encourage
practitioners to interpret tissue and system stress within an
Throughout this article, we have highlighted the importance athlete monitoring framework to contextualize load–response
of considering the tissue response when determining if an data.
appropriate dose of training load has been applied. In this
respect, monitoring either external or internal load in isola- Declarations
tion will be inadequate to determine training adaptations. Funding No funding or grants from any public, commercial, or not-
Indeed, the athlete monitoring framework described in Fig. 2 for-profit organizations were used in the preparation of this manuscript.
emphasizes the importance of using at least two sources of
monitoring information to contextualize load–response Conflict of Interest T.J.G. works as a consultant to several high-perfor-
data and inform subsequent training prescription. When mance organizations, including sporting teams, artistic, industry, mili-
presented with an injury, rehabilitation professionals are tary, and higher-education institutions. E.O. has no potential conflicts
of interest with the content of this article.
encouraged to first consider typical tissue recovery time-
frames before initiating training programs. Consideration of Data Availability No datasets were generated or analyzed for this nar-
the local tissue capacity required to perform sporting skills is rative review.
necessary to ensure appropriate local tissue loading is incor-
Author Contributions T.J.G. conceptualized and wrote the first draft
porated into the training schedule. Importantly, systemic of the paper. E.O. provided critical feedback on subsequent drafts of
loading is also recommended during rehabilitation to main- the paper. Both authors take responsibility for the content of the paper.
tain global capacity and prevent detraining of noninjured The authors would like to thank the reviewers for their constructive
tissues. Solid clinical reasoning should always underpin the comments on the paper.
training process for both injured and healthy athletes, but the
typical tissue and system load–response recommendations
presented in Fig. 1 can be used as a starting point to guide References
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