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Muscle Biology and Healing

51 citationsUpdated Sep 2026
Illustration: Muscle Biology and Healing

Overview

Skeletal muscle possesses a distinct regenerative capacity driven by the origin, gene expression, and coordinated regulation of stem cell populations [11]. Satellite cells play a central role in muscle fiber repair and remodeling, a function integrated from in vivo human studies within the wider context of satellite cell biology [4]. Understanding these basic principles of regeneration and healing processes helps clinicians avoid potential dangers and accelerate the return to competition [1]. Emerging findings further describe the mechanisms that underlie normal versus aberrant muscle-tissue repair [7], while type 2 innate immunity is required for regeneration after injury [15]. Genetic determinants governing the formation or repair of various muscles during different stages of myogenesis are unexpectedly diverse [18].

Current clinical strategies for muscle tissue loss include novel methods for muscle regeneration, which face challenges for future clinical translation [3]. Adipose-derived mesenchymal stem cells have demonstrated efficacy in muscle, tendon, bone, and cartilage regeneration in preclinical and clinical studies [6], though their safety and side effects at long-term remain unsolved [6]. Muscle-derived stem cells are well suited for gene therapy and tissue engineering applications for the musculoskeletal system [30]. Satellite cells and other cell types also hold therapeutic potential for muscle regeneration [10]. Prevention of fibrosis could enhance muscle regeneration, thereby facilitating more efficient muscle healing [2].

Muscles may have a preferred sarcomere length operating range [48], and a compartment is defined as an enclosed space formed by fascia or by a combination of fascia and bone that contains one or more muscles [49]. Fatty infiltration should be used with caution as an indicator of chronicity or a contraindicating parameter for repair [27]. Ultrasonography offers dynamic muscle assessment and is fast, relatively inexpensive, and easier for patients [12]. It allows for detection and severity assessment of muscle injuries as well as serial evaluation to follow healing [12]. A consistent English terminology and a comprehensive classification system for athletic muscle injuries are presented in the Munich consensus statement [13].

How It Works

Cellular Mechanisms and Regeneration

Muscle fiber repair and remodeling rely on satellite cell function [4]. The inflammatory response is critical, with IL-10 playing a central role in regulating the switch of muscle macrophages from a pro-inflammatory M1 to a pro-regenerative M2 phenotype in injured muscle in vivo [16]. This phenotypic transition is necessary for normal growth and regeneration of muscle [16]. Matrix metalloproteinases (MMPs) are involved in muscle repair, with fast and slow twitch muscles exhibiting different patterns of MMP-9 and MMP-2 activity during regeneration [21].

Growth factor signaling and genetic regulation further modulate the healing environment. FGF signaling can play a role in regulation of postnatal skeletal muscle maintenance [32] and could offer potentially novel and efficient therapeutic options for attenuating muscle atrophy during aging, illness, and spaceflight [32]. MicroRNAs have myriad roles in muscle biology that pose prospects for therapeutic manipulation in muscle disease [31]. Additionally, the inherent pro-regenerative potential of tissue-specific endothelium could be used therapeutically to orchestrate fibrosis-free healing and to restore homeostasis in tissues [20].

Pathology and Aberrant Repair

Aberrant muscle-tissue repair involves distinct mechanisms compared to normal repair [7]. Sarcopenia reflects a progressive withdrawal of anabolism and an increased catabolism, along with a reduced muscle regeneration capacity [17]. Understanding sex-dependent responses to different forms of muscle atrophy and inflammation is of pivotal importance to design innovative, tailored, and efficient interventions [23]. Epigenetic mechanism observations have the potential to become useful tools in sports medicine as predictors of approaching pathophysiological alterations and injury biomarkers [19].

Clinical Interventions and Outcomes

Imaging: Ultrasonography allows for detection and severity assessment as well as serial evaluation to follow healing [12].

Biological Agents: Decorin can efficiently prevent fibrosis and enhance muscle regeneration in the lacerated muscle [24]. Simultaneous PRP and suramin use reduced fibrosis in the injured muscle and promoted healing without negatively affecting the muscle's contractile properties [22]. Customized platelet-rich plasma is a promising method to improve PRP's beneficial effect on skeletal muscle repair [14]. However, several worries about the safety and side effects of adipose-derived mesenchymal stem cells at long-term remain unsolved [6].

Mechanical and Surgical Factors: Immobilization resulted in slower muscle regeneration and the development of a large area of scar tissue [5]. Commonly used therapeutic modalities affect muscle regeneration and identify opportunities to further improve the treatment of skeletal muscle strain injuries [9]. Atrophy of the supraspinatus muscle is reversible following repair of the rotator cuff [33].

What the Evidence Shows

Muscle Biology and Regeneration Mechanisms

Interleukin-10 (IL-10) centrally regulates the transition of muscle macrophages from an M1 to an M2 phenotype in injured muscle in vivo, a shift necessary for normal growth and regeneration [16]. The structure and physiological function of the extracellular matrix (ECM) serve as important regulators of cell functions and skeletal muscle development [52]. Matrix metalloproteinases (MMPs) participate in muscle repair, with fast and slow twitch muscles exhibiting distinct patterns of MMP-9 and MMP-2 activity [21]. Additionally, altered expression and secretion of soluble mediators, including IL-6 and IGF-1, modulate regulatory networks involved in altered regeneration and fibrosis during aging and diseases [50].

Satellite Cells and Stem Cells

Satellite cell function in muscle fiber repair and remodeling is integrated from in vivo human studies within the wider context of satellite cell biology, which is largely based on animal and cell models [4]. Muscle-derived stem cells (MDSCs) demonstrate potential for long-term regeneration and multipotency in enhancing skeletal muscle repair [26]. In a rat injury model, single intramuscular administration of muscle precursor cells (MPCs) improved histological outcome and force recovery of the injured skeletal muscle [35]. Adipose-derived mesenchymal stem cells have demonstrated efficacy in muscle, tendon, bone, and cartilage regeneration in preclinical and clinical studies, though long-term safety and side effects remain unsolved [6].

Fibrosis and Aberrant Repair

Aberrant muscle-tissue repair is characterized by specific mechanisms that differ from normal repair [7]. Fatty infiltration is irreversible and progressive if left untreated, though slight reversal of muscle atrophy has been noted after repair in some studies [37]. Simultaneous use of platelet-rich plasma (PRP) and suramin reduced fibrosis in the injured muscle and promoted healing without negatively affecting the muscle's contractile properties [22].

Diagnostic and Classification Standards

Ultrasonography offers dynamic muscle assessment and is fast, relatively inexpensive, and easier for patients, allowing for detection and severity assessment as well as serial evaluation to follow healing [12]. A consistent English terminology and a comprehensive classification system for athletic muscle injuries have been presented that are proven in daily practice [13].

Therapeutic Modalities and Interventions

Immobilization: Immobilization resulted in slower muscle regeneration and the development of a large area of scar tissue compared to suturing [5]. Pulsed Ultrasound: Treatment with pulsed ultrasound can promote the satellite cell proliferation phase of myoregeneration, but it does not seem to have significant effects on the overall morphological manifestations of muscle regeneration [36]. Hyperbaric Oxygen: Exposure to 1 hour of hyperbaric oxygen (HBO) treatment initiated within 20 minutes after exercise for 3 to 5 days enhances muscle torque recovery [45]. Sports Massage: Sports massage treatment did not affect the level or duration of pain or the loss of strength or function following eccentric exercise [43]. Platelet-Rich Plasma: Customized platelet-rich plasma (PRP) is a promising method to improve PRP's beneficial effect on skeletal muscle repair [14]. Proximal Nerve Injuries: Strategies to minimize muscle damage in proximal nerve injuries include physical exercises, electrical stimulation, medications, regenerative medicine, and surgical procedures, noting that most data comes from experimental animal studies while clinical data remains limited [8]. Surgical Intervention: Early surgical intervention with fasciotomy played a substantial role in allowing optimal muscle regeneration, leading to complete functional recovery in a patient with compartment syndrome and familial rhabdomyolysis [34].

Clinical Outcomes and Surgical Techniques

Surgical repair of a clinical tear of the pectoralis major results in greater recovery of peak torque and work performed than conservative management [28]. A presented surgical technique for proximal hamstring tears provides a strong repair that can tolerate rehabilitation and give good long-term results [29]. Outcomes after repair of partial- and full-thickness rotator cuff tears using a bioinductive implant show safety and efficacy at 1-year follow-up [25]. Peak load at failure was comparable between repair constructs in double-layer rotator cuff repairs of delaminated tears [40]. Nonoperative treatment remains a viable option for certain patients with traumatic rotator cuff tears; however, results demonstrate a considerable early failure rate [42]. Randomised trials are needed to assess and compare conservative with surgical interventions and to evaluate different surgical procedures for rotator cuff injury [44]. Understanding the injury patterns, treatment, and prevention options is important for a successful recovery and return to sport for CrossFit athletes [46].

Practical Considerations

General Principles and Classification

A consistent English terminology and a comprehensive classification system for athletic muscle injuries, proven in daily practice, are presented to standardize clinical communication [13]. In the elderly population, sarcopenia represents a loss of muscle mass and function that reduces mobility and diminishes quality of life. This condition can lead to fall-related injuries, which require costly hospitalization and extended rehabilitation [47].

Imaging and Assessment

No specific evidence bullets were provided for this subsection in the source document.

Surgical Management

Objective strength testing demonstrates that surgical repair of a clinical tear of the pectoralis major results in greater recovery of peak torque and work performed than conservative management [28]. For proximal hamstring tears, the presented technique provides a strong repair that can tolerate rehabilitation and yields good long-term results [29].

Non-Operative and Adjunctive Therapies

Clinicians should base advisory guidance and management on sound medical principles, noting that while NSAIDs decrease perceived soreness, they do not impact muscle weakness [41]. In clinical practice, the authors recommend the use of CWI and CRYO for reducing exercise-induced muscle damage [39]. Blood flow restriction (BFR) is applied to enhance strength and hypertrophy while understanding associated safety issues [38]. Platelet-rich plasma (PRP) benefits on skeletal muscle repair can be improved through specific approaches [14]. Various strategies to minimize muscle damage include physical exercises, electrical stimulation, medications, regenerative medicine, and surgical procedures; however, most data comes from experimental animal studies while clinical data remains limited [8]. Commonly used therapeutic modalities affect muscle regeneration, identifying opportunities to further improve the treatment of skeletal muscle strain injuries [9].

Regenerative Medicine and Cellular Therapy

Stem cell therapies and tissue engineering offer promising avenues for enhancing skeletal muscle repair, with muscle-derived stem cells (MDSCs) showing potential for long-term regeneration and multipotency [26]. These cells are well suited for gene therapy and tissue engineering applications for the musculoskeletal system, with the potential to revolutionize the management of certain musculoskeletal diseases [30]. The therapeutic potential of satellite cells and other cell types for muscle regeneration is addressed in current literature [10]. While preclinical and clinical studies have demonstrated efficacy in muscle, tendon, bone, and cartilage regeneration, several worries about their safety and side effects at long-term remain unsolved [6]. A transformative model has opened a fresh chapter in translational vascular medicine, raising the possibility that the inherent pro-regenerative potential of tissue-specific endothelium could be used therapeutically to orchestrate fibrosis-free healing and to restore homeostasis in tissues [20]. These results support the hypothesis that prevention of fibrosis could enhance muscle regeneration, thereby facilitating more efficient muscle healing [2].

Biological Mechanisms

This article reviews the emerging findings of the mechanisms that underlie normal versus aberrant muscle-tissue repair [7]. A comprehensive overview of the epidemiology of muscle tissue loss highlights current strategies in clinical treatment and discusses novel methods for muscle regeneration and challenges for their future clinical translation [3]. The present review examines studies focusing on the origin, gene expression, and coordinated regulation of stem cell populations to highlight the regenerative capacity of skeletal muscle and emphasize the challenges for this field [11]. Recent results from in vivo human studies on satellite cell function in muscle fiber repair and remodeling are integrated within the wider context of satellite cell biology, whose literature is largely based on animal and cell models [4]. Collectively, these data show that IL-10 plays a central role in regulating the switch of muscle macrophages from a M1 to M2 phenotype in injured muscle in vivo, and this transition is necessary for normal growth and regeneration of muscle [16].

Key Evidence

  • [L5] Recognition of basic principles of skeletal muscle regeneration and healing processes can help avoid dangers and accelerate return to competition. [1] (10.1177/0363546505274714)
  • [Paper] These results support our hypothesis that prevention of fibrosis could enhance muscle regeneration, thereby facilitating more efficient muscle healing. [2] (10.1152/japplphysiol.00915.2002)
  • [L5] This review provides a comprehensive overview of the epidemiology of muscle tissue loss, highlights current strategies in clinical treatment, and discusses novel methods for muscle regeneration and challenges for their future clinical translation. [3] (10.1155/2018/1984879)
  • [L5] This review integrates recent results from in vivo human studies on satellite cell function in muscle fiber repair and remodeling within the wider context of satellite cell biology, whose literature is largely based on animal and cell models. [4] (10.3389/fphys.2015.00283)
  • [L5] Immobilization resulted in slower muscle regeneration and the development of a large area of scar tissue. [5] (10.1177/03635465990270021801)
  • [L4] While preclinical and clinical studies have demonstrated efficacy in muscle, tendon, bone, and cartilage regeneration, several worries about their safety and side effects at long-term remain unsolved. [6] (10.3390/ijms20123105)
  • [L5] This article reviews the emerging findings of the mechanisms that underlie normal versus aberrant muscle-tissue repair. [7] (10.1186/2044-5040-1-21)
  • [L5] It outlines various strategies including physical exercises, electrical stimulation, medications, regenerative medicine, and surgical procedures to minimize muscle damage, noting that most data comes from experimental animal studies while clinical data remains limited. [8] (10.1177/17531934231216646)
  • [L5] It summarizes how commonly used therapeutic modalities affect muscle regeneration and identifies opportunities to further improve the treatment of skeletal muscle strain injuries. [9] (10.1123/jsr.2016-0107)
  • [Paper] The present review examines studies focusing on the origin, gene expression, and coordinated regulation of stem cell populations to highlight the regenerative capacity of skeletal muscle and emphasize the challenges for this field. [11] (10.1101/gad.1419406)
  • [L5] Ultrasonography offers dynamic muscle assessment and is fast, relatively inexpensive, and easier for patients, allowing for detection and severity assessment as well as serial evaluation to follow healing. [12] (10.1148/radiol.2017160267)
  • [L5] A consistent English terminology as well as a comprehensive classification system for athletic muscle injuries which is proven in the daily practice are presented. [13] (10.1136/bjsports-2012-091448)
  • [L5] This approach is a promising method to improve PRP's beneficial effect on skeletal muscle repair. [14] (10.1177/2325967116s00143)
  • [L5] Type 2 innate immunity is required for regeneration of skeletal muscle after injury. [15] (10.1016/j.cell.2013.02.053)
  • [L5] Collectively, these data show that IL-10 plays a central role in regulating the switch of muscle macrophages from a M1 to M2 phenotype in injured muscle in vivo, and this transition is necessary for normal growth and regeneration of muscle. [16] (10.4049/jimmunol.1103180)
  • [L4] Sarcopenia reflects a progressive withdrawal of anabolism and an increased catabolism, along with a reduced muscle regeneration capacity. [17] (10.1093/bmb/ldq008)
  • [L5] Genetic determinants of formation or repair of various muscles during different stages of myogenesis are unexpectedly diverse. [18] (10.1097/mco.0b013e328336ea98)
  • [L4] Epigenetic mechanism observations have the potential to become useful tools in sports medicine, as predictors of approaching pathophysiological alterations and injury biomarkers that have already taken place. [19] (10.3390/genes13081471)
  • [L5] This transformative model has opened a fresh chapter in translational vascular medicine and raised the possibility that the inherent pro-regenerative potential of tissue-specific endothelium could be used therapeutically to orchestrate fibrosis-free healing and to restore homeostasis in tissues. [20] (10.1038/nature17040)
  • [L5] MMPs are involved in muscle repair, and that fast and slow twitch muscles exhibit different patterns of MMP-9 and MMP-2 activity. [21] (10.1387/ijdb.072331mz)
  • [L5] Simultaneous PRP and suramin use reduced fibrosis in the injured muscle and promoted healing without negatively affecting the muscle's contractile properties. [22] (10.1177/03635465211030295)
  • [L5] Understanding sex-dependent responses to different forms of muscle atrophy and inflammation is of pivotal importance to design innovative, tailored, and efficient interventions. [23] (10.3390/ijms24054651)
  • [L5] These results suggest that decorin can efficiently prevent fibrosis and enhance muscle regeneration in the lacerated muscle. [24] (10.1177/03635465010290040201)
  • [L4] Outcomes after repair of partial- and full-thickness rotator cuff tears using a bioinductive implant show safety and efficacy at 1-year follow-up. [25] (10.1016/j.arthro.2019.02.019)
  • [L5] Stem cell therapies and tissue engineering offer promising avenues for enhancing skeletal muscle repair, with muscle-derived stem cells (MDSCs) showing potential for long-term regeneration and multipotency. [26] (10.1016/j.csm.2008.08.009)
  • [L4] Therefore, fatty infiltration should be used with caution as an indicator of chronicity or a contraindicating parameter for repair. [27] (10.1177/17585732211024504)
  • [L3] Objective strength testing shows that surgical repair of a clinical tear of the pectoralis major results in greater recovery of peak torque and work performed than conservative management. [28] (10.1136/bjsm.35.3.202)
  • [Paper] The presented technique provides a strong repair that can tolerate rehabilitation and give good long-term results. [29] (10.1016/j.eats.2016.10.004)
  • [L5] Muscle-derived stem cells are well suited for gene therapy and tissue engineering applications for the musculoskeletal system, with the potential to revolutionize the management of certain musculoskeletal diseases. [30] (10.5435/00124635-200802000-00004)
  • [Paper] The myriad roles of microRNAs in muscle biology pose interesting prospects for their therapeutic manipulation in muscle disease. [31] (10.1016/j.tig.2008.01.007)
  • [Paper] These results support the theory that FGF signaling can play a role in regulation of postnatal skeletal muscle maintenance, and could offer potentially novel and efficient therapeutic options for attenuating muscle atrophy during aging, illness and spaceflight. [32] (10.1186/1471-2474-8-32)
  • [L4] This is the first study to provide evidence that atrophy of the supraspinatus muscle is reversible. [33] (10.1302/0301-620x.98b10.37231)
  • [L5] Early surgical intervention with fasciotomy played a substantial role in allowing optimal muscle regeneration, leading to complete functional recovery. [34] (10.2106/00004623-200211000-00021)
  • [Paper] Single intramuscular administration of MPCs improved histological outcome and force recovery of the injured skeletal muscle in a rat injury model. [35] (10.1177/0363546521989235)
  • [L5] Treatment with pulsed ultrasound can promote the satellite cell proliferation phase of myoregeneration, but it does not seem to have significant effects on the overall morphological manifestations of muscle regeneration. [36] (10.1177/03635465990270011701)
  • [L5] Fatty infiltration is irreversible and progressive if left untreated, though slight reversal of muscle atrophy has been noted after repair in some studies. [37] (10.5435/jaaos-21-10-613)
  • [L2] It aims to clarify how to apply BFR effectively for enhancing strength and hypertrophy while understanding associated safety issues. [38] (10.3389/fphys.2019.00533)
  • [L1] In clinical practice, the authors recommend the use of CWI and CRYO for reducing exercise-induced muscle damage. [39] (10.1186/s12891-024-07315-2)
  • [L5] Peak load at failure was comparable between repair constructs. [40] (10.1177/0363546518796818)
  • [L5] Clinicians should base advisory guidance and management on sound medical principles, noting that while NSAIDs decrease perceived soreness, they do not impact muscle weakness. [41] (10.1016/j.csm.2011.09.009)
  • [L4] Nonoperative treatment remains a viable option for certain patients with traumatic rotator cuff tears; however, the results of our study demonstrate a considerable early failure rate. [42] (10.1016/j.jse.2023.11.012)
  • [L1] The treatment did not affect the level or duration of pain or the loss of strength or function following exercise. [43] (10.1177/0363546503262196)
  • [L5] Randomised trials are needed to assess and compare conservative with surgical interventions and to evaluate different surgical procedures. [44] (10.1007/s00167-009-0735-y)
  • [L1] Exposure to 1 hour of HBO treatment initiated within 20 minutes after exercise for 3 to 5 days enhances muscle torque recovery. [45] (10.1177/03635465990270050901)
  • [L4] Understanding the injury patterns, treatment, and prevention options is important for a successful recovery and return to sport. [46] (10.5435/jaaos-d-22-01219)
  • [L5] Sarcopenia is a loss of muscle mass and function in the elderly that reduces mobility, diminishes quality of life, and can lead to fall-related injuries, which require costly hospitalization and extended rehabilitation. [47] (10.1152/physrev.00061.2017)
  • [L5] The paper introduces the concept that muscles may have a preferred sarcomere length operating range. [48] (10.1098/rstb.2010.0316)
  • [Paper] A compartment is defined as an enclosed space formed by fascia or by a combination of fascia and bone that contains one or more muscles. [49] (10.1016/s0749-0712(21)00394-2)
  • [L5] This review discusses the cellular mediators of fibrosis and how the altered expression and secretion of soluble mediators, such as IL-6 and IGF-1, can modulate regulatory networks involved in the altered regeneration and fibrosis during aging and diseases. [50] (10.3390/cells8030232)
  • [L5] This review provides a comprehensive overview of the structure, physiological function, and application of the extracellular matrix (ECM) in skeletal muscle tissue. [52] (10.1186/s13578-021-00579-4)

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[14] Customized Platelet-Rich Plasma for Skeletal Muscle Injuries. Orthopaedic Journal of Sports Medicine. 2016. DOI: 10.1177/2325967116s00143

[15] Type 2 Innate Signals Stimulate Fibro/Adipogenic Progenitors to Facilitate Muscle Regeneration. Cell. 2013. DOI: 10.1016/j.cell.2013.02.053

[16] IL-10 Triggers Changes in Macrophage Phenotype That Promote Muscle Growth and Regeneration. The Journal of Immunology. 2012. DOI: 10.4049/jimmunol.1103180

[17] Sarcopenia: characteristics, mechanisms and functional significance. British Medical Bulletin. 2010. DOI: 10.1093/bmb/ldq008

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[40] Double-Layer Rotator Cuff Repair: Anatomic Reconstruction of the Superior Capsule and Rotator Cuff Improves Biomechanical Properties in Repairs of Delaminated Rotator Cuff Tears. The American Journal of Sports Medicine. 2018. DOI: 10.1177/0363546518796818

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[45] Effects of Hyperbaric Oxygen on a Human Model of Injury. The American Journal of Sports Medicine. 1999. DOI: 10.1177/03635465990270050901

[46] Common Orthopaedic Injuries in CrossFit Athletes. Journal of the American Academy of Orthopaedic Surgeons. 2023. DOI: 10.5435/jaaos-d-22-01219

[47] Sarcopenia: Aging-Related Loss of Muscle Mass and Function. Physiological Reviews. 2019. DOI: 10.1152/physrev.00061.2017

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[52] Extracellular matrix: an important regulator of cell functions and skeletal muscle development. Cell & Bioscience. 2021. DOI: 10.1186/s13578-021-00579-4

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Considerations for the public: By using one of our public licenses, a licensor grants the public permission to use the licensed material under specified terms and conditions. If the licensor's permission is not necessary for any reason--for example, because of any applicable exception or limitation to copyright--then that use is not regulated by the license. Our licenses grant only permissions under copyright and certain other rights that a licensor has authority to grant. Use of the licensed material may still be restricted for other reasons, including because others have copyright or other rights in the material. A licensor may make special requests, such as asking that all changes be marked or described. Although not required by our licenses, you are encouraged to respect those requests where reasonable. More considerations for the public: wiki.creativecommons.org/Considerations_for_licensees


Creative Commons Attribution-NonCommercial 4.0 International Public License

By exercising the Licensed Rights (defined below), You accept and agree to be bound by the terms and conditions of this Creative Commons Attribution-NonCommercial 4.0 International Public License ("Public License"). To the extent this Public License may be interpreted as a contract, You are granted the Licensed Rights in consideration of Your acceptance of these terms and conditions, and the Licensor grants You such rights in consideration of benefits the Licensor receives from making the Licensed Material available under these terms and conditions.

Section 1 -- Definitions.

a. Adapted Material means material subject to Copyright and Similar Rights that is derived from or based upon the Licensed Material and in which the Licensed Material is translated, altered, arranged, transformed, or otherwise modified in a manner requiring permission under the Copyright and Similar Rights held by the Licensor. For purposes of this Public License, where the Licensed Material is a musical work, performance, or sound recording, Adapted Material is always produced where the Licensed Material is synched in timed relation with a moving image.

b. Adapter's License means the license You apply to Your Copyright and Similar Rights in Your contributions to Adapted Material in accordance with the terms and conditions of this Public License.

c. Copyright and Similar Rights means copyright and/or similar rights closely related to copyright including, without limitation, performance, broadcast, sound recording, and Sui Generis Database Rights, without regard to how the rights are labeled or categorized. For purposes of this Public License, the rights specified in Section 2(b)(1)-(2) are not Copyright and Similar Rights.

d. Effective Technological Measures means those measures that, in the absence of proper authority, may not be circumvented under laws fulfilling obligations under Article 11 of the WIPO Copyright Treaty adopted on December 20, 1996, and/or similar international agreements.

e. Exceptions and Limitations means fair use, fair dealing, and/or any other exception or limitation to Copyright and Similar Rights that applies to Your use of the Licensed Material.

f. Licensed Material means the artistic or literary work, database, or other material to which the Licensor applied this Public License.

g. Licensed Rights means the rights granted to You subject to the terms and conditions of this Public License, which are limited to all Copyright and Similar Rights that apply to Your use of the Licensed Material and that the Licensor has authority to license.

h. Licensor means the individual(s) or entity(ies) granting rights under this Public License.

i. NonCommercial means not primarily intended for or directed towards commercial advantage or monetary compensation. For purposes of this Public License, the exchange of the Licensed Material for other material subject to Copyright and Similar Rights by digital file-sharing or similar means is NonCommercial provided there is no payment of monetary compensation in connection with the exchange.

j. Share means to provide material to the public by any means or process that requires permission under the Licensed Rights, such as reproduction, public display, public performance, distribution, dissemination, communication, or importation, and to make material available to the public including in ways that members of the public may access the material from a place and at a time individually chosen by them.

k. Sui Generis Database Rights means rights other than copyright resulting from Directive 96/9/EC of the European Parliament and of the Council of 11 March 1996 on the legal protection of databases, as amended and/or succeeded, as well as other essentially equivalent rights anywhere in the world.

l. You means the individual or entity exercising the Licensed Rights under this Public License. Your has a corresponding meaning.

Section 2 -- Scope.

a. License grant.

1. Subject to the terms and conditions of this Public License, the Licensor hereby grants You a worldwide, royalty-free, non-sublicensable, non-exclusive, irrevocable license to exercise the Licensed Rights in the Licensed Material to:

a. reproduce and Share the Licensed Material, in whole or in part, for NonCommercial purposes only; and

b. produce, reproduce, and Share Adapted Material for NonCommercial purposes only.

2. Exceptions and Limitations. For the avoidance of doubt, where Exceptions and Limitations apply to Your use, this Public License does not apply, and You do not need to comply with its terms and conditions.

3. Term. The term of this Public License is specified in Section 6(a).

4. Media and formats; technical modifications allowed. The Licensor authorizes You to exercise the Licensed Rights in all media and formats whether now known or hereafter created, and to make technical modifications necessary to do so. The Licensor waives and/or agrees not to assert any right or authority to forbid You from making technical modifications necessary to exercise the Licensed Rights, including technical modifications necessary to circumvent Effective Technological Measures. For purposes of this Public License, simply making modifications authorized by this Section 2(a) (4) never produces Adapted Material.

5. Downstream recipients.

a. Offer from the Licensor -- Licensed Material. Every recipient of the Licensed Material automatically receives an offer from the Licensor to exercise the Licensed Rights under the terms and conditions of this Public License.

b. No downstream restrictions. You may not offer or impose any additional or different terms or conditions on, or apply any Effective Technological Measures to, the Licensed Material if doing so restricts exercise of the Licensed Rights by any recipient of the Licensed Material.

6. No endorsement. Nothing in this Public License constitutes or may be construed as permission to assert or imply that You are, or that Your use of the Licensed Material is, connected with, or sponsored, endorsed, or granted official status by, the Licensor or others designated to receive attribution as provided in Section 3(a)(1)(A)(i).

b. Other rights.

1. Moral rights, such as the right of integrity, are not licensed under this Public License, nor are publicity, privacy, and/or other similar personality rights; however, to the extent possible, the Licensor waives and/or agrees not to assert any such rights held by the Licensor to the limited extent necessary to allow You to exercise the Licensed Rights, but not otherwise.

2. Patent and trademark rights are not licensed under this Public License.

3. To the extent possible, the Licensor waives any right to collect royalties from You for the exercise of the Licensed Rights, whether directly or through a collecting society under any voluntary or waivable statutory or compulsory licensing scheme. In all other cases the Licensor expressly reserves any right to collect such royalties, including when the Licensed Material is used other than for NonCommercial purposes.

Section 3 -- License Conditions.

Your exercise of the Licensed Rights is expressly made subject to the following conditions.

a. Attribution.

1. If You Share the Licensed Material (including in modified form), You must:

a. retain the following if it is supplied by the Licensor with the Licensed Material:

i. identification of the creator(s) of the Licensed Material and any others designated to receive attribution, in any reasonable manner requested by the Licensor (including by pseudonym if designated);

ii. a copyright notice;

iii. a notice that refers to this Public License;

iv. a notice that refers to the disclaimer of warranties;

v. a URI or hyperlink to the Licensed Material to the extent reasonably practicable;

b. indicate if You modified the Licensed Material and retain an indication of any previous modifications; and

c. indicate the Licensed Material is licensed under this Public License, and include the text of, or the URI or hyperlink to, this Public License.

2. You may satisfy the conditions in Section 3(a)(1) in any reasonable manner based on the medium, means, and context in which You Share the Licensed Material. For example, it may be reasonable to satisfy the conditions by providing a URI or hyperlink to a resource that includes the required information.

3. If requested by the Licensor, You must remove any of the information required by Section 3(a)(1)(A) to the extent reasonably practicable.

4. If You Share Adapted Material You produce, the Adapter's License You apply must not prevent recipients of the Adapted Material from complying with this Public License.

Section 4 -- Sui Generis Database Rights.

Where the Licensed Rights include Sui Generis Database Rights that apply to Your use of the Licensed Material:

a. for the avoidance of doubt, Section 2(a)(1) grants You the right to extract, reuse, reproduce, and Share all or a substantial portion of the contents of the database for NonCommercial purposes only;

b. if You include all or a substantial portion of the database contents in a database in which You have Sui Generis Database Rights, then the database in which You have Sui Generis Database Rights (but not its individual contents) is Adapted Material; and

c. You must comply with the conditions in Section 3(a) if You Share all or a substantial portion of the contents of the database.

For the avoidance of doubt, this Section 4 supplements and does not replace Your obligations under this Public License where the Licensed Rights include other Copyright and Similar Rights.

Section 5 -- Disclaimer of Warranties and Limitation of Liability.

a. UNLESS OTHERWISE SEPARATELY UNDERTAKEN BY THE LICENSOR, TO THE EXTENT POSSIBLE, THE LICENSOR OFFERS THE LICENSED MATERIAL AS-IS AND AS-AVAILABLE, AND MAKES NO REPRESENTATIONS OR WARRANTIES OF ANY KIND CONCERNING THE LICENSED MATERIAL, WHETHER EXPRESS, IMPLIED, STATUTORY, OR OTHER. THIS INCLUDES, WITHOUT LIMITATION, WARRANTIES OF TITLE, MERCHANTABILITY, FITNESS FOR A PARTICULAR PURPOSE, NON-INFRINGEMENT, ABSENCE OF LATENT OR OTHER DEFECTS, ACCURACY, OR THE PRESENCE OR ABSENCE OF ERRORS, WHETHER OR NOT KNOWN OR DISCOVERABLE. WHERE DISCLAIMERS OF WARRANTIES ARE NOT ALLOWED IN FULL OR IN PART, THIS DISCLAIMER MAY NOT APPLY TO YOU.

b. TO THE EXTENT POSSIBLE, IN NO EVENT WILL THE LICENSOR BE LIABLE TO YOU ON ANY LEGAL THEORY (INCLUDING, WITHOUT LIMITATION, NEGLIGENCE) OR OTHERWISE FOR ANY DIRECT, SPECIAL, INDIRECT, INCIDENTAL, CONSEQUENTIAL, PUNITIVE, EXEMPLARY, OR OTHER LOSSES, COSTS, EXPENSES, OR DAMAGES ARISING OUT OF THIS PUBLIC LICENSE OR USE OF THE LICENSED MATERIAL, EVEN IF THE LICENSOR HAS BEEN ADVISED OF THE POSSIBILITY OF SUCH LOSSES, COSTS, EXPENSES, OR DAMAGES. WHERE A LIMITATION OF LIABILITY IS NOT ALLOWED IN FULL OR IN PART, THIS LIMITATION MAY NOT APPLY TO YOU.

c. The disclaimer of warranties and limitation of liability provided above shall be interpreted in a manner that, to the extent possible, most closely approximates an absolute disclaimer and waiver of all liability.

Section 6 -- Term and Termination.

a. This Public License applies for the term of the Copyright and Similar Rights licensed here. However, if You fail to comply with this Public License, then Your rights under this Public License terminate automatically.

b. Where Your right to use the Licensed Material has terminated under Section 6(a), it reinstates:

1. automatically as of the date the violation is cured, provided it is cured within 30 days of Your discovery of the violation; or

2. upon express reinstatement by the Licensor.

For the avoidance of doubt, this Section 6(b) does not affect any right the Licensor may have to seek remedies for Your violations of this Public License.

c. For the avoidance of doubt, the Licensor may also offer the Licensed Material under separate terms or conditions or stop distributing the Licensed Material at any time; however, doing so will not terminate this Public License.

d. Sections 1, 5, 6, 7, and 8 survive termination of this Public License.

Section 7 -- Other Terms and Conditions.

a. The Licensor shall not be bound by any additional or different terms or conditions communicated by You unless expressly agreed.

b. Any arrangements, understandings, or agreements regarding the Licensed Material not stated herein are separate from and independent of the terms and conditions of this Public License.

Section 8 -- Interpretation.

a. For the avoidance of doubt, this Public License does not, and shall not be interpreted to, reduce, limit, restrict, or impose conditions on any use of the Licensed Material that could lawfully be made without permission under this Public License.

b. To the extent possible, if any provision of this Public License is deemed unenforceable, it shall be automatically reformed to the minimum extent necessary to make it enforceable. If the provision cannot be reformed, it shall be severed from this Public License without affecting the enforceability of the remaining terms and conditions.

c. No term or condition of this Public License will be waived and no failure to comply consented to unless expressly agreed to by the Licensor.

d. Nothing in this Public License constitutes or may be interpreted as a limitation upon, or waiver of, any privileges and immunities that apply to the Licensor or You, including from the legal processes of any jurisdiction or authority.


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