Muscles, exercise and obesity: skeletal muscle as a secretory organ
Abstract
During the past decade, skeletal muscle has been identified as a secretory organ. Accordingly, we have suggested that cytokines and other peptides that are produced, expressed and released by muscle fibres and exert either autocrine, paracrine or endocrine effects should be classified as myokines. The finding that the muscle secretome consists of several hundred secreted peptides provides a conceptual basis and a whole new paradigm for understanding how muscles communicate with other organs, such as adipose tissue, liver, pancreas, bones and brain. However, some myokines exert their effects within the muscle itself. Thus, myostatin, LIF, IL-6 and IL-7 are involved in muscle hypertrophy and myogenesis, whereas BDNF and IL-6 are involved in AMPK-mediated fat oxidation. IL-6 also appears to have systemic effects on the liver, adipose tissue and the immune system, and mediates crosstalk between intestinal L cells and pancreatic islets. Other myokines include the osteogenic factors IGF-1 and FGF-2; FSTL-1, which improves the endothelial function of the vascular system; and the PGC-1α-dependent myokine irisin, which drives brown-fat-like development. Studies in the past few years suggest the existence of yet unidentified factors, secreted from muscle cells, which may influence cancer cell growth and pancreas function. Many proteins produced by skeletal muscle are dependent upon contraction; therefore, physical inactivity probably leads to an altered myokine response, which could provide a potential mechanism for the association between sedentary behaviour and many chronic diseases.
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BDNF can be induced by exercise and released from skeletal muscle, contributing to metabolic regulation.
“Additionally, factors like BDNF can be induced by exercise and released from skeletal muscle, contributing to metabolic regulation [75].”
From Exercise-Induced Myokines in Obesity-Related Metabolic Disorders and Cardiovascular Protection: A Narrative Review (2026) · cites this paper as [75]
No review has provided a comprehensive integrative overview of these novel molecular players and the mechanisms involved in metabolic fuel redistribution, weight and fat mass loss, and reduced inflammation.
“To our knowledge, no review has provided a comprehensive integrative overview of these novel molecular players and the mechanisms involved in the redistribution of metabolic fuel during and after exercise, the loss of weight and fat mass, and reduced inflammation.”
These myokines exert beneficial physiologic and metabolic effects in skeletal muscle and distant tissues such as white adipose tissue, bone, liver, central nervous system, and immune cells to drive a systemic anti-inflammatory and insulin-sensitive state permissive for optimization of total-body energy expenditure.
“These exert beneficial physiologic and metabolic effects not only in SkM itself but also in distant tissues such as white adipose tissue (WAT), bone, liver, the central nervous system (CNS), and cells of the immune system to drive a systemic anti-inflammatory and insulin-sensitive state, permissive for optimization of total-body energy expenditure [5,7,26,27].”
Literature on this topic has been reviewed by others on a separate basis, by describing either individual molecules or groups of factors, proteins, hormones, or molecules.
“Literature on this topic is vast and has been reviewed by others but on a separate basis, by describing either individual molecules or groups of factors, proteins, hormones, or molecules [2,3,4,5,6,7,8,9,29].”
Power training (PT) may amplify the myokine response, contributing to the observed reductions in body fat and improvements in body composition.
“PT, by involving intense and repeated contractions of large muscle groups, may amplify this myokine response, contributing to the observed reductions in body fat and improvements in body composition [37,38].”
From Efficacy of Power Training on Sarcopenic Obesity in Community-Dwelling Older Women: A 32-Week Randomized Clinical Trial (2025) · cites this paper as [37]
Skeletal muscles function as an endocrine organ by releasing myokines in response to contraction, which can influence fat metabolism, reduce inflammation, and improve insulin sensitivity.
“Skeletal muscles function as an endocrine organ by releasing myokines in response to contraction, which can influence fat metabolism, reduce inflammation, and improve insulin sensitivity.”
From Efficacy of Power Training on Sarcopenic Obesity in Community-Dwelling Older Women: A 32-Week Randomized Clinical Trial (2025) · cites this paper as [37]
During physical exercise, skeletal muscle and adipose tissue engage in complex and dynamic interorgan crosstalk essential for maintaining metabolic homeostasis, mediated by myokines and adipokines.
“During physical exercise, skeletal muscle and adipose tissue engage in complex and dynamic interorgan crosstalk that is essential for maintaining metabolic homeostasis. This communication is mediated by a wide range of myokines and adipokines”
From Exercise-Induced Muscle–Fat Crosstalk: Molecular Mediators and Their Pharmacological Modulation for the Maintenance of Metabolic Flexibility in Aging (2025) · cites this paper as [87]
Myokines released from contracting muscle fibers stimulate lipolysis in WAT, increasing availability of circulating free FA acids as energy substrates for active muscles.
“Myokines released from contracting muscle fibers stimulate lipolysis in WAT, increasing the availability of circulating free FA acids as energy substrates for active muscles”
From Exercise-Induced Muscle–Fat Crosstalk: Molecular Mediators and Their Pharmacological Modulation for the Maintenance of Metabolic Flexibility in Aging (2025) · cites this paper as [87]
Physical activity is necessary to lose weight and improve muscle strength, which is beneficial to ameliorate the state of sarcopenic obesity.
“Numerous studies have proven that physical activity is necessary to lose weight and improve muscle strength, which is beneficial to ameliorate the state of SO (16, 17).”
From Diabetes and Sarcopenic Obesity: Pathogenesis, Diagnosis, and Treatments (2020) · cites this paper as [17]
Myokines mediate communication between muscle and other organs, including brain, adipose tissue, bone, liver, gut, pancreas, vascular bed, and skin, as well as within the muscle itself.
“Myokines mediate communication between muscle and other organs, including brain, adipose tissue, bone, liver, gut, pancreas, vascular bed, and skin, as well as within the muscle itself”
From Muscle–Organ Crosstalk: The Emerging Roles of Myokines (2020) · cites this paper as [7]
During exercise, myokines signal within the muscle and mediate muscle-organ crosstalk to the brain, adipose tissue, bone, liver, gut, pancreas, vascular bed, and skin.
“During exercise, myokines signal within the muscle and mediate muscle–organ crosstalk to the brain, adipose tissue, bone, liver, gut, pancreas, vascular bed, and skin (7, 29, 30).”
From Muscle–Organ Crosstalk: The Emerging Roles of Myokines (2020) · cites this paper as [7]
IL-6 works in a paracrine manner exerting metabolic effects within the muscle itself.
“While IL-6 is characterized as a myokine with endocrine effects, it also works in a paracrine manner exerting metabolic effects within the muscle itself (6, 7).”
From Muscle–Organ Crosstalk: The Emerging Roles of Myokines (2020) · cites this paper as [7]
Muscle contraction is sensed by the brain, suggesting that peripheral factors induced by exercise may be involved in direct crosstalk between working muscle and brain function.
“The finding that muscle contraction is sensed by the brain suggests that peripheral factors induced by exercise may be involved in direct crosstalk between working muscle and brain function (7, 29, 30, 83) (Fig. 2).”
From Muscle–Organ Crosstalk: The Emerging Roles of Myokines (2020) · cites this paper as [7]
Myokines provide skeletal muscle with the ability to mediate whole-body metabolism via endocrine signaling to adipose tissue, liver, pancreas, heart, and brain.
“Myokines provide skeletal muscle with the ability to mediate whole-body metabolism via endocrine signaling to adipose tissue, liver, pancreas, heart, and brain (122).”
From Metabolic Flexibility as an Adaptation to Energy Resources and Requirements in Health and Disease (2018) · cites this paper as [122]
Resting circulating IL-6 levels are lower in exercise-trained people than in untrained people.
“increased after acute exercise, the baseline (or ‘resting’) circulating levels of IL-6 are lower in exercise-trained individuals than in untrained individuals78.”
From Exerkines in health, resilience and disease (2022) · cites this paper as [78]
Bone, muscle, and adipose tissues secrete numerous cytokines.
“It is well understood that the bone, 165 muscle 236 and adipose tissues 237 secrete numerous cytokines, some of which represent an unexamined source of potential novel therapeutic targets.”
From The role of the androgen receptor in the pathogenesis of obesity and its utility as a target for obesity treatments (2022) · cites this paper as [236]
During exercise skeletal muscle tightly interacts with adipose tissue, pancreas, and liver to sustain the energy demands of physical activity and mediate positive effects in whole‐body energy metabolism.
“More precisely, during exercise skeletal muscle tightly interacts with adipose tissue, pancreas, and liver to sustain the energy demands of physical activity and mediate positive effects in whole‐body energy metabolism.”
From Inter‐organ communication: a gatekeeper for metabolic health (2019) · cites this paper as [17]
IL‐6 induces an anti‐inflammatory response by reducing TNF‐α production while increasing the levels of IL‐10, IL‐1Ra, and IL‐15.
“Moreover, IL‐6 has the ability to induce an anti‐inflammatory response by reducing tumor necrosis factor‐α (TNF‐α) production while increasing the levels of interleukin 10 (IL‐10), interleukin 1 receptor antagonist (IL‐1Ra), and interleukin 15 (IL‐15) 17, 114.”
From Inter‐organ communication: a gatekeeper for metabolic health (2019) · cites this paper as [17]
IL‐6 mediates muscle glucose uptake, increased insulin sensitivity, and increased fatty acid oxidation.
“Exercise‐induced IL‐6 production was initially believed to be linked to muscle damage, but has now been proven to directly depend on contraction and to mediate muscle glucose uptake, increased insulin sensitivity, and increased fatty acid oxidation.”
From Inter‐organ communication: a gatekeeper for metabolic health (2019) · cites this paper as [17]
Exercise-induced IL‐6 production directly depends on contraction.
“Exercise‐induced IL‐6 production was initially believed to be linked to muscle damage, but has now been proven to directly depend on contraction and to mediate muscle glucose uptake, increased insulin sensitivity, and increased fatty acid oxidation.”
From Inter‐organ communication: a gatekeeper for metabolic health (2019) · cites this paper as [17]
IL‐6 triggers insulin production in pancreas, hepatic glucose production, and adipose tissue lipolysis, thus ensuring the availability of energy substrates for exercising muscle.
“IL‐6 also triggers insulin production in pancreas, hepatic glucose production, and adipose tissue lipolysis, thus ensuring the availability of energy substrates for exercising muscle 11, 17.”
From Inter‐organ communication: a gatekeeper for metabolic health (2019) · cites this paper as [17]
Metabolic issues associated with low healthy behaviors are reversed with exposure to physical activity (e.g., exercise).
“And are reversed with exposure to physical activity (e.g., exercise) with speculation that resistance exercise may provide the greatest impact on reversing such issues [28,47-50]”
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Self-citationAssociation between Interleukin-15 and Obesity: Interleukin-15 as a Potential Regulator of Fat Mass
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Self-citationInterleukin-6 Regulation of AMP-Activated Protein Kinase
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Self-citationMuscle Cytokine mRNA Changes after 2.5 h of Cycling: Influence of Carbohydrate
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Self-citationHumoral Nature of the Hypoglycemic Factor of Muscular Work
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Leukaemia inhibitory factor - an exercise-induced myokine
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Self-citationComparative proteomic analysis of the insulin‐induced L6 myotube secretome
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Self-citationHepatosplanchnic clearance of interleukin-6 in humans during exercise
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Self-citationCalprotectin is released from human skeletal muscle tissue during exercise
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Self-citationAutocrine IL-6 regulates GRO-alpha production in thymic epithelial cells.
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Exercise Metabolism and the Molecular Regulation of Skeletal Muscle Adaptation
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Inflammation in obesity, diabetes, and related disorders
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Homeostasis, Inflammation, and Disease Susceptibility
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Muscle–Organ Crosstalk: The Emerging Roles of Myokines
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“The role of myokines has previously been reviewed (7, 11-37), identifying more than 650 myokines (38).”
BackgroundInfluentialSelf-citationAdipose tissue browning and metabolic health
Bartelt, A., & Heeren, J. (2014) · Nature Reviews Endocrinology · doi:10.1038/nrendo.2013.204 · Cited by 1,177 · see citing works
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Exerkines in health, resilience and disease
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Self-citationIL-6 pathway in the liver: From physiopathology to therapy
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Interleukin‐6 myokine signaling in skeletal muscle: a double‐edged sword?
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Rethinking IL-6 and CRP: Why they are more than inflammatory biomarkers, and why it matters
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Metabolic Flexibility as an Adaptation to Energy Resources and Requirements in Health and Disease
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Physical activity and muscle-brain crosstalk
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Self-citationRole of Inactivity in Chronic Diseases: Evolutionary Insight and Pathophysiological Mechanisms
Booth, F. W., Roberts, C. K., Thyfault, J. P., Ruegsegger, G. N., & Toedebusch, R. G. (2017) · Physiological Reviews · doi:10.1152/physrev.00019.2016 · Cited by 696 · see citing works
Skeletal muscle inflammation and insulin resistance in obesity
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Targeting inflammation in the treatment of type 2 diabetes: time to start
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Irisin Mediates Effects on Bone and Fat via αV Integrin Receptors
Kim, H., Wrann, C. D., Jedrychowski, M., Vidoni, S., Kitase, Y., Nagano, K., Zhou, C., Chou, J., Parkman, V.-J. A., Novick, S. J., et al. (2018) · Cell · doi:10.1016/j.cell.2018.10.025 · Cited by 638 · see citing works
“Physical activity doesn’t only target osteocytes but also stimulates the production of several hormone-like molecules from skeletal muscle termed “myokines” (Pedersen and Febbraio, 2012).”
Hepatokines: Linking nonalcoholic fatty liver disease and insulin resistance
Meex, R. C. R., & Watt, M. J. (2017) · Nature Reviews Endocrinology · doi:10.1038/nrendo.2017.56 · Cited by 631 · see citing works
Intermittent metabolic switching, neuroplasticity and brain health
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FNDC5/irisin is not only a myokine but also an adipokine
Roca-Rivada, A., Castelao, C., Senin, L. L., Landrove, M. O., Baltar, J., Crujeiras, A. B., Seoane, L. M., Casanueva, F. F., & Pardo, M. (2013) · PLoS ONE · doi:10.1371/journal.pone.0060563 · Cited by 612 · see citing works
Anti-inflammatory effects of exercise: role in diabetes and cardiovascular disease
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Self-citationMolecular mechanisms linking exercise to cancer prevention and treatment
Hojman, P., Gehl, J., Christensen, J. F., & Pedersen, B. K. (2018) · Cell Metabolism · doi:10.1016/j.cmet.2017.09.015 · Cited by 600 · see citing works
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An official European Respiratory Society statement on physical activity in COPD
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Self-citationExercise-stimulated glucose uptake — regulation and implications for glycaemic control
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The role of hepatokines in metabolism
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Exercise as an anti-inflammatory therapy for rheumatic diseases—Myokine regulation
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Self-citationCan physical activity ameliorate immunosenescence and thereby reduce age-related multi-morbidity?
Duggal, N. A., Niemiro, G., Harridge, S. D. R., Simpson, R. J., & Lord, J. M. (2019) · Nature Reviews Immunology · doi:10.1038/s41577-019-0177-9 · Cited by 436 · see citing works
The role of adipokines in health and disease
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The potential of endurance exercise-derived exosomes to treat metabolic diseases
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Blaming the brain for obesity: integration of hedonic and homeostatic mechanisms
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Energy intake and exercise as determinants of brain health and vulnerability to injury and disease
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- Authors
- Pedersen, B. K., & Febbraio, M. A.
- Affiliations
- University of Copenhagen (DK)
- Baker Heart and Diabetes Institute (AU)
- Corresponding author
- Pedersen, Bente K.
- Venue
- Nature Reviews Endocrinology
- Published
- 2012-04-03
- Volume
- 8
- Issue
- 8
- Pages
- 457-465
- ISSN
- 1759-5029 (ISSN-L)
- 1759-5037
- OpenAlex
- W2117457995
- Semantic Scholar
- 7053d2f849635807b22a4df75d203dcd876ce49b
- Cited by
- 2,909
- References
- 123
- Citations by year
- 2026: 307 · 2025: 252 · 2024: 208 · 2023: 218 · 2022: 247 · 2021: 240 · 2020: 231 · 2019: 214 · 2018: 181 · 2017: 194 · 2016: 147 · 2015: 168 · 2014: 147 · 2013: 128 · 2012: 22
- FWCI
- 19.03
- Type
- journal article
- Language
- en
- Open access
- closed
- Licence
- www.springer.com/tdm (tdm)
- Locations
- Nature Reviews Endocrinology (publishedVersion)
- PubMed (publishedVersion)
- Research at the University of Copenhagen (University of Copenhagen) (publishedVersion)
- Topics
- Health Sciences › Medicine › Rehabilitation › Exercise and Physiological Responses
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Muscle Physiology and Disorders
- Health Sciences › Medicine › Epidemiology › Adipokines, Inflammation, and Metabolic Diseases
- Keywords
- myokines, physical activity, obesity, interleukin-6, irisin, myostatin, brain-derived neurotrophic factor, AMPK, fat oxidation, muscle hypertrophy, insulin-like growth factor I, FGF2, PGC-1α, brown adipose tissue, sedentary behavior, physical inactivity, cancer, pancreatic islets, intestinal L cells
- MeSH terms
- Adipose Tissue, Bone and Bones, Brain, Humans, Liver, Obesity, Pancreas, Exercise, Cytokines, Muscle, Skeletal
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