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Muscle Releases Alpha-Sarcoglycan Positive Extracellular Vesicles Carrying miRNAs in the Bloodstream
Open accessAbstract
In the past few years, skeletal muscle has emerged as an important secretory organ producing soluble factors, called myokines, that exert either autocrine, paracrine or endocrine effects. Moreover, recent studies have shown that muscle releases microRNAs into the bloodstream in response to physical exercise. These microRNAs affect target cells, such as hormones and cytokines. The mechanisms underlying microRNA secretion are poorly characterized at present. Here, we investigated whether muscle tissue releases extracellular vesicles (EVs), which carry microRNAs in the bloodstream under physiological conditions such as physical exercise. Using density gradient separation of plasma from sedentary and physically fit young men we found EVs positive for TSG101 and alpha-sarcoglycan (SGCA), and enriched for miR-206. Cytometric analysis showed that the SGCA+ EVs account for 1-5% of the total and that 60-65% of these EVs were also positive for the exosomal marker CD81. Furthermore, the SGCA-immuno captured sub-population of EVs exhibited higher levels of the miR-206/miR16 ratio compared to total plasma EVs. Finally, a significant positive correlation was found between the aerobic fitness and muscle-specific miRNAs and EV miR-133b and -181a-5p were significantly up-regulated after acute exercise. Thus, our study proposes EVs as a novel means of muscle communication potentially involved in muscle remodeling and homeostasis.
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“These ELVs were also positive for CD81, TSG101 and miR-206”
From Skeletal Muscle-Released Extracellular Vesicles: State of the Art (2019) · cites this paper as Guescini et al., 2015
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“ELVs containing alpha-sarcoglycan (SGCA) a component of the dystrophin-glycoprotein complex involved in the stability of muscle fiber membranes, were found in human plasma”
From Skeletal Muscle-Released Extracellular Vesicles: State of the Art (2019) · cites this paper as Guescini et al., 2015
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Bradford, M. M. (1976) · Analytical Biochemistry · doi:10.1016/0003-2697(76)90527-3 · Cited by 255,302 · cited 1×
“After centrifugation at 18,000 g, protein concentration was determined by the Bradford assay [35].”
MethodAnalysis of Relative Gene Expression Data Using Real-Time Quantitative PCR and the 2−ΔΔCT Method
Livak, K. J., & Schmittgen, T. D. (2001) · Methods · doi:10.1006/meth.2001.1262 · Cited by 180,860 · cited 1×
“Quantification was performed according to the ΔCq method [34], miR-16 or the exogenously added spike-in cel-miR-39 were used as a reference.”
MethodValadi, H., Ekström, K., Bossios, A., Sjöstrand, M., Lee, J. J., & Lötvall, J. O. (2007) · Nature Cell Biology · doi:10.1038/ncb1596 · Cited by 12,834 · cited 1×
“Numerous studies indicate that circulating miRNAs may be protected from degradation by several complementary mechanisms, including their incorporation into secreted membrane vesicles such as microvesicles, exosomes or apoptotic bodies [16–18], or via the formation of RNA-binding protein complexes [19].”
Background(2014) · Medical Entomology and Zoology · Cited by 9,400
Skog, J., Würdinger, T., van Rijn, S., Meijer, D. H., Gainche, L., Curry, W. T., Carter, B. S., Krichevsky, A. M., & Breakefield, X. O. (2008) · Nature Cell Biology · doi:10.1038/ncb1800 · Cited by 5,001
Mutation of the mouse klotho gene leads to a syndrome resembling ageing
Kuro-o, M., Matsumura, Y., Aizawa, H., Kawaguchi, H., Suga, T., Utsugi, T., Ohyama, Y., Kurabayashi, M., Kaname, T., Kume, E., et al. (1997) · Nature · doi:10.1038/36285 · Cited by 3,837
Arroyo, J. D., Chevillet, J. R., Kroh, E. M., Ruf, I. K., Pritchard, C. C., Gibson, D. F., Mitchell, P. S., Bennett, C. F., Pogosova-Agadjanyan, E. L., Stirewalt, D. L., et al. (2011) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1019055108 · Cited by 3,294 · cited 1×
“Numerous studies indicate that circulating miRNAs may be protected from degradation by several complementary mechanisms, including their incorporation into secreted membrane vesicles such as microvesicles, exosomes or apoptotic bodies [16–18], or via the formation of RNA-binding protein complexes [19].”
BackgroundMuscles, exercise and obesity: skeletal muscle as a secretory organ
Pedersen, B. K., & Febbraio, M. A. (2012) · Nature Reviews Endocrinology · doi:10.1038/nrendo.2012.49 · Cited by 2,909 · cited 1×
“Therefore, the discovery that contracting muscle is a myokine-producing organ has opened up a whole new field of research on skeletal muscle [6].”
BackgroundMicroRNAs are transported in plasma and delivered to recipient cells by high-density lipoproteins
Vickers, K. C., Palmisano, B. T., Shoucri, B. M., Shamburek, R. D., & Remaley, A. T. (2011) · Nature Cell Biology · doi:10.1038/ncb2210 · Cited by 2,741 · cited 2×
“In the extracellular environment, miRNAs could be stabilized by EVs (such as exosomes, microvesicles or apoptotic bodies), HDL/LDLs, or protected by RNA-binding proteins (reviewed in [14,45]).”
BackgroundExosomes: Extracellular organelles important in intercellular communication
Mathivanan, S., Ji, H. L., & Simpson, R. J. (2010) · Journal of Proteomics · doi:10.1016/j.jprot.2010.06.006 · Cited by 2,581 · cited 1×
“11 g/mL, as previously described [37].”
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Items 101–200 of 227
Vann, C. G., Zhang, X., Khodabukus, A., Orenduff, M. C., Chen, Y., Corcoran, D. L., Truskey, G. A., Bursac, N., & Kraus, V. B. (2022) · Frontiers in Physiology · doi:10.3389/fphys.2022.937899 · Cited by 18
Yedigaryan, L., Martínez‐Sarrà, E., Giacomazzi, G., Giarratana, N., van der Veer, B. K., Rotini, A., Querceto, S., Grosemans, H., Cortés-Calabuig, Á., Salucci, S., et al. (2022) · Frontiers in Immunology · doi:10.3389/fimmu.2022.977617 · Cited by 18
“Interestingly, it has been reported that EVs carry members of the myomiR family of miRNAs, such as miR-1, miR-133a, and miR-206, and thus they might contribute directly to conveying regenerative signalling to the skeletal muscle (19, 20).”
Extracellular Vesicles in Cancer Drug Resistance: Implications on Melanoma Therapy
Musi, A., & Bongiovanni, L. (2023) · Cancers · doi:10.3390/cancers15041074 · Cited by 18
“In the skeletal muscle, the role of EVs has also been investigated, showing that development and differentiation of the muscle after an intense exercise could also be EV-dependent [15].”
BackgroundExtracellular vesicles and Duchenne muscular dystrophy pathology: Modulators of disease progression
Yedigaryan, L., & Sampaolesi, M. (2023) · Frontiers in Physiology · doi:10.3389/fphys.2023.1130063 · Cited by 19
“As an example, miR-206 is one of the most abundant miRNAs in muscle-derived EVs which aids in muscle injury repair by promoting development and differentiation of tissues (Guescini et al., 2015).”
BackgroundBean, A., Sahu, A., Piechocki, C., Gualerzi, A., Picciolini, S., Bedoni, M., & Ambrosio, F. (2023) · Experimental Gerontology · doi:10.1016/j.exger.2023.112179 · Cited by 19
Pinto, A. C., Tavares, P., Neves, B. M., Oliveira, P. F., Vitorino, R. M. P., Moreira‐Gonçalves, D., & Ferreira, R. M. (2024) · Journal of Molecular Medicine · doi:10.1007/s00109-024-02427-7 · Cited by 19
“In addition to variations in the levels of circulating EVs, it remains uncertain whether the proportion of SkM-derived EVs increases beyond the baseline of 5% after an exercise bout [10], whether in physically active or sedentary individuals.”
InfluentialGoldschmidt‐Clermont, P. J., Khan, A., Jimsheleishvili, G., Graham, P., Brooks, A. E., Silvera, R., Goldschmidt, A. J. P., Pearse, D. D., Dietrich, W. D., Levi, A. D., et al. (2024) · Neural Regeneration Research · doi:10.4103/nrr.nrr-d-23-01815 · Cited by 18
Skeletal muscle-derived exosomes prevent osteoporosis by promoting osteogenesis
Xing, Z., Guo, L., Li, S., Huang, W., Su, J., Chen, X., Li, Y., & Zhang, J. (2024) · Life Sciences · doi:10.1016/j.lfs.2024.123079 · Cited by 20
Sutton, E. R., & Pekovic‐Vaughan, V. (2025) · Antioxidants · doi:10.3390/antiox14091132 · Cited by 20
“A further study by Guescini and colleagues reported that circulation of miR-133b and miR181a-5p in exosomes was increased following acute exercise, with a positive correlation between aerobic fitness and circulating skeletal muscle-abundant miRNAs [25].”
InfluentialExtracellular vesicles secreted from mouse muscle cells improve delayed bone repair in diabetic mice
Takafuji, Y., Kawao, N., Ohira, T., Mizukami, Y., Okada, K., Jo, J., Tabata, Y., & Kaji, H. (2022) · Endocrine Journal · doi:10.1507/endocrj.ej22-0340 · Cited by 18
Kargl, C. K., Jia, Z., Shera, D. A., Sullivan, B. P., Burton, L. C., Kim, K. H., Nie, Y., Hubal, M. J., Shannahan, J. H., Kuang, S., et al. (2023) · Scientific Reports · doi:10.1038/s41598-023-45787-9 · Cited by 18
Hart, D. A. (2021) · Frontiers in Sports and Active Living · doi:10.3389/fspor.2021.685845 · Cited by 18
“Understanding the mechanisms associated with alterations induced by sedentary behavior is an active area of research (Pedersen and Febbraio, 2012; Guescini et al., 2015; Wu et al., 2015; Smits et al., 2018; Ghosh et al., 2019; Greco et al., 2019; Olanrewaju et al., 2020; Padilha et al., 2021; Hou et al., 2022).”
BackgroundBydak, B., Pierdoná, T. M., Seif, S., Sidhom, K., Obi, P. O., Labouta, H. I., Gordon, J. W., & Saleem, A. (2022) · Membranes · doi:10.3390/membranes12050464 · Cited by 15
“, alpha-sarcoglycan [50], while abundant in muscle, cannot be guaranteed to be expressed in all Skm-EVs, (2) the difficulty in confirming that the Skm-EVs originated from the muscle undergoing contractile activity”
BackgroundConkright, W. R., Kargl, C. K., Hubal, M. J., Tiede, D., Beckner, M. E., Sterczala, A. J., Krajewski, K. T., Martin, B. J., Flanagan, S. D., Greeves, J. P., et al. (2024) · Medicine & Science in Sports & Exercise · doi:10.1249/mss.0000000000003408 · Cited by 16
“%) of circulating EVs originate from skeletal muscle (38), and there is little overlap between circulating EV miRNAs and those derived from muscle (39).”
BackgroundAnabolic and Catabolic Signaling Pathways That Regulate Skeletal Muscle Mass
McCarthy, J. J., & Murach, K. A. (2018) · Elsevier eBooks · doi:10.1016/b978-0-12-813922-6.00024-2 · Cited by 17
The Role of miRNAs and Extracellular Vesicles in Adaptation After Resistance Exercise: A Review
Csala, D., Ádám, Z., & Wilhelm, M. (2025) · Current Issues in Molecular Biology · doi:10.3390/cimb47080583 · Cited by 16
“Therefore, Annibalini et al. [65] suggested that skeletal muscle also participates in the expression of circulating EVs, since the post-RE EV-packaged miR-206 amount increased [121].”
Potential Role of Exercise Induced Extracellular Vesicles in Prostate Cancer Suppression
Zhang, Y., Kim, J., Wang, T.-Z., Newton, R. U., Galvao, D. A., Gardiner, R. A., Hill, M. M. C., & Taaffe, D. R. (2021) · Frontiers in Oncology · doi:10.3389/fonc.2021.746040 · Cited by 15
“Not only moderate-intensity aerobic exercise but high and low-intensity exercise also have been shown to increase serum EVs concentration and EVs protein content (45, 47).”
BackgroundKargl, C. K., Sullivan, B. P., Middleton, D., York, A. P. E., Burton, L. C., Brault, J. J., Kuang, S., & Gavin, T. P. (2022) · Experimental Physiology · doi:10.1113/ep090874 · Cited by 14
“Acute exercise increases SkM-EV release (Gao et al., 2021; Guescini et al., 2015) and it appears that EV release is regulated by intrinsic metabolic properties of muscle fibres and an oxidative phenotype (Estrada et al., 2021; Nie et al., 2019).”
BackgroundLee, H., He, X., Ni, K., Carnino, J. M., & Jin, Y. (2021) · American Journal of Physiology-Lung Cellular and Molecular Physiology · doi:10.1152/ajplung.00318.2020 · Cited by 15
Lipinski, S., & Tiemann, K. (2021) · International Journal of Molecular Sciences · doi:10.3390/ijms22073374 · Cited by 15
Bone-Derived Extracellular Vesicles: Novel Players of Interorgan Crosstalk
Li, Y., Yin, P., Guo, Z., Lv, H., Deng, Y., Chen, M., Gu, Y., Tang, P., & Zhang, L. (2019) · Frontiers in Endocrinology · doi:10.3389/fendo.2019.00846 · Cited by 15
“found that muscle tissue could release EVs carrying muscle-specific miR-133b and miR-181a-5p under regular physical exercise (22).”
BackgroundSharma, A. R., Chatterjee, S., Lee, Y., & Lee, S. (2023) · Aging and Disease · doi:10.14336/ad.2023.00903 · Cited by 15
“It has been observed that approximately 60-65% of these muscle-specific EVs are positive for CD81 [228].”
BackgroundDomańska-Senderowska, D., Jastrzębski, Z., Kiszałkiewicz, J., Brzeziański, M., Pastuszak‐Lewandoska, D., Radzimińki, Ł., Brzeziańska‐Lasota, E., & Jegier, A. (2017) · Biology of Sport · doi:10.5114/biolsport.2017.69820 · Cited by 13
Bourgeois, B. L., Levitt, D. E., Molina, P. E., & Simon, L. (2022) · Alcohol · doi:10.1016/j.alcohol.2022.11.001 · Cited by 13
Circulating microRNAs: let’s not waste the potential
Hiam, D., & Lamon, S. (2020) · American Journal of Physiology-Cell Physiology · doi:10.1152/ajpcell.00175.2020 · Cited by 12
Shen, P., Qiu, Y., Sun, Y.-Y., Jiang, Y.-Y., Guan, X., Cheng, M., & Wang, Y. (2024) · Reviews in Cardiovascular Medicine · doi:10.31083/j.rcm2511392 · Cited by 13
Intracellular and extracellular redox signals during exercise and aging
Caporossi, D., Jackson, M. J., & Henríquez‐Olguín, C. (2025) · Free Radical Biology and Medicine · doi:10.1016/j.freeradbiomed.2025.10.283 · Cited by 15
Mastrototaro, L., & Roden, M. (2023) · PROTEOMICS · doi:10.1002/pmic.202300078 · Cited by 13
“Furthermore, SGCA + EV are characterized by high levels of muscle-specific miRNAs (myomiRNA), which positively correlate with VO 2max , but only miR-133b and − 181a-5p are upregulated after acute endurance exercise in healthy individuals [185].”
BackgroundExercise-induced extracellular vesicles in reprogramming energy metabolism in cancer
Puurand, M., Llorente, A., Linē, A., & Käämbre, T. (2025) · Frontiers in Oncology · doi:10.3389/fonc.2024.1480074 · Cited by 14
Divergent multiomic acute exercise responses reveal the impact of sex as a biological variable
Lavin, K. M., O’Bryan, S. M., Pathak, K. V., Garcia‐Mansfield, K., Graham, Z., McAdam, J. S., Drummer, D. J., Bell, M. B., Kelley, C. J., Lixandrão, M. E., et al. (2025) · Physiological Genomics · doi:10.1152/physiolgenomics.00055.2024 · Cited by 14
Atheroprotective roles of exercise-regulated microRNAs
Ke, S., Cao, X.-Q., Lu, X., Xu, J., Zhang, C.-Y., Xu, L., & Jiang, X. (2025) · Atherosclerosis · doi:10.1016/j.atherosclerosis.2025.119229 · Cited by 12
Circulating microRNAs: advances in exercise physiology
Sapp, R. M., & Hagberg, J. M. (2019) · Current Opinion in Physiology · doi:10.1016/j.cophys.2019.03.004 · Cited by 12
Malnutrition and Biomarkers: A Journey through Extracellular Vesicles
Mendivil-Alvarado, H., Sosa-León, L. A., Carvajal‐Millán, E., & Astiazarán‐García, H. (2022) · Nutrients · doi:10.3390/nu14051002 · Cited by 12
“Others have suggested that most of the circulating EVs during exercise are released by the muscle tissue, as it is the organ with the highest secretory activity [93,96,97].”
BackgroundWu, Z., Gao, Y., Chen, Q., & Li, Y. (2025) · Journal of Cellular and Molecular Medicine · doi:10.1111/jcmm.70896 · Cited by 13
“…shown that exosomes from mice performing HIIT exhibit enriched levels of muscle-specific miRNAs (myomiRs), such as miR-133a and miR-133b, which are associated with VO 2 max in humans, while showing reduced expression of obesity-related miR-192, compared to those from sedentary controls [54, 65].”
Qiu, X., Zhou, J., Xu, Y., Liao, L., Yang, H., Xiang, Y., Zhou, Z., Sun, Q. X., Chen, M., Zhang, J., et al. (2022) · Frontiers in Cardiovascular Medicine · doi:10.3389/fcvm.2022.881526 · Cited by 11
“Circulating exosomes packaged with some muscle-specific miRNAs (miR-133b, miR181a-5p, miR-146a and miR-206) dramatically increased after acute exercise and exerted a versatile role in muscle repair and maintenance (55, 56).”
BackgroundRNAkines are secreted messengers shaping health and disease
Li, J., Fang, J., Jiang, X., Zhang, Y., Vidal‐Puig, A., & Zhang, C. (2023) · Trends in Endocrinology and Metabolism · doi:10.1016/j.tem.2023.12.004 · Cited by 11
“Sequence motifs are recognised by speci fi c RNA-binding proteins (RBPs) that act as the machinery for localising ncRNAs into EVs.”
Role of exosomes and exosomal microRNA in muscle–Kidney crosstalk in chronic kidney disease
Zhou, S., Cheing, G. L. Y., & Cheung, K. (2022) · Frontiers in Cell and Developmental Biology · doi:10.3389/fcell.2022.951837 · Cited by 12
“reported that the expression level of miR-181a-5p and miR-133b in musclederived circulating exosomes was elevated after acute exercise, and they found a positive correlation between aerobic fitness and muscle-specific miRNAs (Guescini et al., 2015).”
BackgroundFaraldi, M., Sansoni, V., Perego, S., Gomarasca, M., Gerosa, L., Ponzetti, M., Rucci, N., Banfi, G., & Lombardi, G. (2022) · Frontiers in Molecular Biosciences · doi:10.3389/fmolb.2022.915080 · Cited by 10
“been a growing interest in exercise-induced EV release (Fruhbeis et al., 2015; Guescini et al., 2015).”
BackgroundFoudi, N., & Legeay, S. (2021) · Fundamental and Clinical Pharmacology · doi:10.1111/fcp.12665 · Cited by 10
Circulating myo-miRs in physical exercise
Malvandi, A. M., Faraldi, M., Sansoni, V., Gerosa, L., Jaworska, J., & Lombardi, G. (2024) · Advanced Exercise and Health Science · doi:10.1016/j.aehs.2024.05.005 · Cited by 9
Exploring the Impact of Exercise-Derived Extracellular Vesicles in Cancer Biology
Silvestri, M., Grazioli, E., Duranti, G., Sgrò, P., & Dimauro, I. (2024) · Biology · doi:10.3390/biology13090701 · Cited by 9
“%) of muscle-derived EVs enter the blood-stream [39–41].”
BackgroundMilczek-Haduch, D., Żmigrodzka, M., & Witkowska‐Piłaszewicz, O. (2025) · International Journal of Molecular Sciences · doi:10.3390/ijms26094359 · Cited by 9
Li, A., Zhou, Z., Li, D., Sha, P.-R., Hu, H., Lin, Y.-Q., Yue, B.-L., Li, J., & Xiong, Y. (2025) · Cells · doi:10.3390/cells14241954 · Cited by 10
“…tissue-specific exosomes form a sophisticated inter-organ communication network and influence processes such as satellite cell differentiation, mitochondrial thermogenesis, and lipid metabolism through their complex bioactive cargo (including proteins, lipids, various nucleic acids, and RNAs) [32].”
Pierri, B., Eitan, E., Witwer, K. W., Ré, D. B., Baccarelli, A., & Wu, H. (2026) · Journal of Extracellular Biology · doi:10.1002/jex2.70106 · Cited by 10
Role of Extracellular Vesicles in the Pathogenesis of Brain Metastasis
Izhar, M., & Lesniak, M. S. (2025) · Journal of Extracellular Biology · doi:10.1002/jex2.70051 · Cited by 9
“…Bearden, 2015), musculoskeletal health (Yates et al. 2022; Choi et al. 2016; Forterre et al. 2014; Le Bihan et al. 2012; Coenen-Stass et al. 2016; Guescini et al. 2015; Deng et al. 2015; Huynh et al. 2016), and immune (Hong 2018, Zhou et al. 2020; Timár et al. 2013; Lim et al. 2015; Nolan et al.…”
Circulating microRNAs fluctuations in exercise-induced cardiac remodeling: A systematic review.
Sanchis‐Gomar, F., Arnau-Moyano, M., Daimiel, L., Lippi, G., Leischik, R., Vallecillo, N., Yvert, T. P., Sáiz, S. L. J., Santiago, C., & Pareja-Galeano, H. (2021) · PubMed Central · Cited by 8
Myogenic microRNAs as Therapeutic Targets for Skeletal Muscle Mass Wasting in Breast Cancer Models
Artigas-Arias, M., Curi, R., & Marzuca-Nassr, G. N. (2024) · International Journal of Molecular Sciences · doi:10.3390/ijms25126714 · Cited by 8
“…decreased in patients with breast cancer compared to healthy individuals is because the breast cancer tumour itself induces a negative regulation of the expression of these microRNAs through exosomes or extracellular vesicles that allow for intercellular communication between tissues [79,83–85].”
BackgroundMechanism of exercise-derived circulating exosomes as a target for sarcopenia management
Wang, X., Huang, H., Chen, J., Zhang, Q., Yuan, Z., Yin, M., Peng, C., & liu, S. (2026) · Frontiers in Physiology · doi:10.3389/fphys.2025.1680485 · Cited by 7
Building‐up fit muscles for the future
Beleza, J., Stevanović-Silva, J., Coxito, P., Costa, R., Ascensão, A., Torrella, J. R., & Magalhães, J. (2021) · European Journal of Clinical Investigation · doi:10.1111/eci.13515 · Cited by 7
Spectroscopic profiling variations in extracellular vesicle biochemistry in a model of myogenesis
Davies, O. G., Powell, S., Rickard, J. J. S., Clancy, M. T., & Oppenheimer, P. G. (2021) · Journal of Tissue Engineering · doi:10.1177/20417314211022092 · Cited by 7
Circulating MicroRNA Responses to Postprandial Lipemia with or without Prior Exercise
Springer, C. B., Sapp, R. M., Evans, W. S., Hagberg, J. M., & Prior, S. J. (2021) · International Journal of Sports Medicine · doi:10.1055/a-1480-7692 · Cited by 7
Alehossein, P., Taheri, M., Ghahremani, P. T., Dakhlallah, D. A., Brown, C. M., Ishrat, T., & Nasoohi, S. (2022) · Translational Stroke Research · doi:10.1007/s12975-022-01025-4 · Cited by 8
“Furthermore, immunoprecipitation techniques exceptionally enable researchers to identify EV characteristics and purify exer-EVs originating from endothelial cells [68], EPCs [68, 78], platelets [206], lymphocytes [41], or myocytes [53].”
BackgroundmiRNAs involved in the regulation of exercise fatigue
Niu, S., Yin, X., Cao, Q., Huang, K., Deng, Z., & Cao, J. (2025) · Frontiers in Physiology · doi:10.3389/fphys.2025.1614942 · Cited by 6
Sireno, L., Dimauro, I., & Caporossi, D. (2026) · Free Radical Biology and Medicine · doi:10.1016/j.freeradbiomed.2025.12.044 · Cited by 6
Lovett, J. A. C., McColl, R. S., Durcan, P., Vechetti, I. J., & Myburgh, K. H. (2024) · Physiological Reports · doi:10.14814/phy2.70056 · Cited by 5
“…apoptotic bodies and larger microvesicles (100–1000 nm); membranous particles that bleb from the cell surface and are not known to contain selectively-packaged cargo in the same way as smaller EVs (30–200 nm) (Frattini et al., 2017; Guescini et al., 2015; Jia et al., 2022; Pratesi et al., 2013).”
BackgroundSbarigia, C., Rome, S., Dini, L., & Tacconi, S. (2024) · Journal of Extracellular Biology · doi:10.1002/jex2.70019 · Cited by 6
Recent advances in cell‐based and cell‐free therapeutic approaches for sarcopenia
Wu, S., Lin, S., Zhang, X., Alizada, M., Wang, L., Zheng, Y., Ke, Q., & Xu, J. (2022) · The FASEB Journal · doi:10.1096/fj.202200675r · Cited by 7
Epigenetics in Exercise Science and Sports Medicine
Zimmer, P., Schenk, A., & Bloch, W. (2016) · Elsevier eBooks · doi:10.1016/b978-0-12-803239-8.00029-6 · Cited by 6
Fernández‐Rhodes, M., Buchan, E., Gagnon, S. D., Qian, J., Gethings, L. A., Lees, R., Peacock, B., Capel, A. J., Martin, N. R. W., Oppenheimer, P. G., et al. (2024) · Journal of Extracellular Biology · doi:10.1002/jex2.171 · Cited by 5
“For example, it is well described that lipoproteins can function as mRNA carriers (Karvinen et al., 2023; Vickers et al., 2011) and vastly outnumber EVs in biofluids such as plasma (Guescini et al., 2015).”
BackgroundThe effects of aerobic and resistance exercise on the lipid profile of extracellular vesicles
Carpenter, T. K., McCashland, M., Sherrick, R. A., Natarajan, S. K., Mandić, M., Kachman, S. D., von Walden, F., Vechetti, I. J., & Fernandez‐Gonzalo, R. (2025) · European Journal of Applied Physiology · doi:10.1007/s00421-025-05973-1 · Cited by 6
“This is particularly relevant for skeletal muscle-derived EVs, as it has been demonstrated that skeletal muscle accounts for 1–5% of plasma EVs (Guescini et al. 2015).”
Watanabe, S., Makino, T., Hiro, K., Sasaki, T., Takahashi, Y., Futagawa, K., Suzuki, M., Sato, R., & Yamauchi, Y. (2025) · Extracellular Vesicle · doi:10.1016/j.vesic.2025.100097 · Cited by 5
Exercise-derived exosomes: molecular mediators of systemic health and disease therapy
Kong, H., Luo, J., Zou, Z., Li, Y., Tang, X., Han, J., Meng, X., Wang, X., Fu, Y., He, M. S., et al. (2026) · Journal of Nanobiotechnology · doi:10.1186/s12951-026-04115-9 · Cited by 5
“Exosomes enriched with miR-133b/206 suppress hepatic gluconeogenesis [81], and miR-324 reduces lipid accumulation in NAFLD by inhibiting Rho-associated coiled-coil-containing protein kinase 1 (ROCK1) [82].”
Boulestreau, J., Ferguson, S. K., Nelson, L. T., & Polgár, N. (2026) · Journal of Extracellular Biology · doi:10.1002/jex2.70139 · Cited by 6
“Skeletal muscle, the primary tissue activated during physical activities, appears to be a key source of these EVs, supporting the notion of a complex interplay between muscle activity and systemic health (Guescini et al. 2015 ; Estrada et al. 2022 ; Mastrototaro and Roden 2024 ).”
InfluentialMechanism of satellite cell regulation by myokines
Furuichi, Y., & FUJII, N. L. (2017) · The Journal of Physical Fitness and Sports Medicine · doi:10.7600/jpfsm.6.311 · Cited by 5
Exercise-induced plasma-derived extracellular vesicles increase adult hippocampal neurogenesis
Connolly, M. G., Fliflet, A. M., Ravi, P., Rosu, D. I., Boppart, M. D., & Rhodes, J. S. (2025) · Brain Research · doi:10.1016/j.brainres.2025.150003 · Cited by 5
Sisia, G., Giudice, E., Attanzio, A., Briglia, M., Piccione, G. G., Trunfio, C., & Arfuso, F. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27010520 · Cited by 4
“%) of circulating EVs originate from skeletal muscle [55], and there is little overlap between circulating EV miRNAs and those derived from muscle [56].”
Self, M. S., Drasbek, K. R., & Vissing, K. (2026) · American Journal of Physiology-Heart and Circulatory Physiology · doi:10.1152/ajpheart.00954.2025 · Cited by 4
Warnier, G., de ten Ryen, S. V. D., Lannoy, C., Mahy, T., Antoine, N., Boyer, E., Kienlen‐Campard, P., Verboven, K., Copine, S., Francaux, M., et al. (2025) · Journal of Extracellular Biology · doi:10.1002/jex2.70087 · Cited by 4
“However, previous work has shown that skeletal muscle-derived EVs (SkM-EVs) account for only 1 to 5% of the total circulating pool (Estrada et al. 2022; Guescini et al. 2015).”
Circulating microRNAs and physical activity: Impact in diabetes
Li, X., & Hallajzadeh, J. (2025) · Clinica Chimica Acta · doi:10.1016/j.cca.2025.120178 · Cited by 3
Wang, J., Fan, S., & Wang, J. (2025) · Molecular and Cellular Biochemistry · doi:10.1007/s11010-025-05357-1 · Cited by 3
“These investigations have contributed to a more comprehensive understanding of EV patterns following single sessions of running or cycling [81, 96].”
InfluentialExtracellular mitochondria: a potential player involved in exercise health benefits
Pedrosa, M. B., Santos, L. L., Ferreira, R. M. S., & Magalhães, J. S. (2025) · Biochimie · doi:10.1016/j.biochi.2025.12.011 · Cited by 4
Mishra, S., Kumar, A., He, Y., Su, Y., Singh, S., Santos, M. F., Singh, R., Lee, J., Furdui, C. M., Shively, C. A., et al. (2024) · GeroScience · doi:10.1007/s11357-024-01439-y · Cited by 3
“While SGCA-positive sEV in circulation have been previously reported [23–26], a few studies using less sensitive western blot-based methods failed to detect SGCA-positive sEV in the plasma/serum total sEV pool [42, 43].”
Giulietti, M., Piva, F., Cecati, M., Maggio, S., Guescini, M., Saladino, T., Scortichini, L., Crocetti, S., Caramanti, M., Battelli, N., et al. (2024) · Cells · doi:10.3390/cells13060479 · Cited by 4
“EV isolation was carried out in accordance with our previous studies [39,40].”
Diagnosis of Extracellular Vesicles in Cardiovascular and Metabolic Diseases
Eichner-Seitz, N. (2023) · Advances in experimental medicine and biology · doi:10.1007/978-981-99-1443-2_12 · Cited by 3
Obi, P. O., Souza, T. F. G., Seif, S., Bydak, B., Klassen, N., West, A. R., Gordon, J. W., & Saleem, A. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.08.01.605916 · Cited by 3
“Whole blood contains a heterogenous mixture of vesicles derived from platelets, erythrocytes, endothelial cells, leukocytes, [24], [27], and with a comparatively small proportion of EVs from skeletal muscle [21].”
BackgroundMethodObi, P. O., de Souza, T. D. F. G., McManus, K. J., West, A. R., Gordon, J. W., & Saleem, A. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.08.29.610232 · Cited by 3
“Moreover, this work was done using EVs isolated from plasma (circulation), which comprise a heterogenous mixture of vesicles derived from platelets, erythrocytes, endothelial cells, leukocytes, [22], [24], and comparatively small proportion from skeletal muscle [25], [26].”
BackgroundMethodWang, Z., Chen, Q., & Choi, K. H. (2025) · Clinica Chimica Acta · doi:10.1016/j.cca.2025.120737 · Cited by 2
Exercise-derived extracellular vesicles in oncology: a new frontier for translational nanomedicine
Silvestri, M., Fantini, C., Duranti, G., Grazioli, E., Caporossi, D., Balbi, C., & Dimauro, I. (2026) · Journal of Translational Medicine · doi:10.1186/s12967-026-07742-w · Cited by 3
“Notably, small RNA-seq and profiling datasets have identified specific myo-miRs, including miR-486 and miR-181a, which regulate cell cycle progression, apoptosis, and immune cell activation, thereby influencing tumor metabolism and immune infiltration [25, 28, 63].”
Wang, A. (2023) · The Journal of Physiology · doi:10.1113/jp284547 · Cited by 2
“Thus, Xhuti et al. used a protein marker, α -sarcoglycan ( α -SGCA), to identify ELVs secreted by skeletal muscle (Guescini et al., 2015).”
BackgroundEffects of Exercise on Circulating Extracellular Vesicles in Cardiovascular Disease
Teixeira, M., Martins, T. S., Gouveia, M., Henriques, A. G., Santos, M., & Ribeiro, F. (2023) · Advances in experimental medicine and biology · doi:10.1007/978-981-99-1443-2_16 · Cited by 2
Obi, P. O., Bydak, B., Safdar, A., & Saleem, A. (2020) · Springer eBooks · doi:10.1007/978-3-030-35358-2_4 · Cited by 2
Epigenetics in exercise science and sports medicine
Schenk, A., Proschinger, S., & Zimmer, P. (2021) · Elsevier eBooks · doi:10.1016/b978-0-12-823928-5.00005-0 · Cited by 2
Obi, P. O., Turner-Brannen, E., West, A. R., Labouta, H. I., Gordon, J. W., & Saleem, A. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.02.25.481852 · Cited by 2
“Skeletal muscle is a secretory organ, releasing EVs that may influence both local and systemic adaptations to exercise [4], [5].”
Response of circulating miRNAs to acute exercise: A systematic Review and Meta-AnalysisPreprint
MacGregor, K. A., Moran, C. N., Broome, S. C., Owen, P. J., Lamon, S., & Hiam, D. (2022) · medRxiv · doi:10.1101/2022.12.20.22283756 · Cited by 2
“CfmiRNAs may be involved in cell-cell communication, influencing gene expression in cells other than the one that produced them (14, 15).”
BackgroundObi, P. O., Souza, T. F. G., Lao, Y., McManus, K. J., Gordon, J. W., LeDuc, R. D., Zahedi, R. P., & Saleem, A. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.09.09.612156 · Cited by 2
“…following acute exercise have isolated EVs from plasma [17], [39], which would comprise a heterogenous mixture of vesicles derived from platelets, erythrocytes, endothelial cells, leukocytes, [18], [40], and with a comparatively little proportion of EVs originating from skeletal muscle [41], [42].”
BackgroundXie, Y., Liu, X., Liu, W., Carr, L. R., Lee, L. P., Imai, N., Ortlund, E. A., & Cohen, D. E. (2024) · Journal of Biological Chemistry · doi:10.1016/j.jbc.2024.107855 · Cited by 1
Wang, B., He, P., Liu, X., Wu, S., Lu, X., & Xu, B. (2025) · ACS Omega · doi:10.1021/acsomega.5c02248 · Cited by 1
Bushman, T., Lin, T., Jin, X., Fu, Q., Zhang, S., & Chen, X. (2025) · Journal of Extracellular Biology · doi:10.1002/jex2.70062 · Cited by 1
“Similarly, EVs originating from hepatocytes, muscle, and cardiomyocytes have been characterised based on their distinct cell-specific markers (Povero et al. 2014, Garcia-Martinez et al. 2016, Guescini et al. 2015, Carpi et al. 2020, Xu et al. 2017, Estrada et al. 2022).”
Gan, J., Zhong, S., Xie, J., & Zou, L. (2025) · Frontiers in Medicine · doi:10.3389/fmed.2025.1707928 · Cited by 1
Zhang, R. X., & Chen, K. (2026) · Frontiers in Immunology · doi:10.3389/fimmu.2026.1856546 · Cited by 1
“Contracting muscle releases vesicles into the circulation, and a proportion of exercise-induced circulating EVs can be traced by muscle-associated markers (170, 219 – 221).”
Is there any possible role for telocytes in the pathogenesis of musculoskeletal diseases?
Deeb, A., Hafez, A. A. E., & Abd-El-Hafez, A. A. A. (2020) · Journal of Medical Histology/Journal of Medical Histology · doi:10.21608/jmh.2021.54933.1088 · Cited by 1
“Nevertheless, these studies and those from healthy human subjects[15] suggest that monitoring EV-derived miRNAs may serve as one approach for tracking disease progression or noninvasively assessing muscle injury following exercise.”
BackgroundTowards unravelling the exercise‐induced adaptations in circulating extracellular vesicles
Turan, G., Reijnders, L. A. J. J., & de Jong, J. C. B. C. (2023) · The Journal of Physiology · doi:10.1113/jp285726 · Cited by 1
“For instance, a study by Guescini et al. (2015) found a different miRNA expression profile within EVs pre-and post-exercise stimulation in young healthy males.”
BackgroundSchubert, R., Wagner, J. B., Kämpfer, N., Oremek, G. M., Bille, J., Kosteczka, R., Holzgreve, F., & Ohlendorf, D. (2024) · Zentralblatt für Arbeitsmedizin Arbeitsschutz und Ergonomie · doi:10.1007/s40664-024-00555-y · Cited by 1
“Guescini et al. [17] fanden im Plasma von männlichen Studenten eine signifikante positive Korrelation zwischen der aeroben Fitness und den muskelspezifischen miRNAs miR-133b und -181a-5p nach akuter Laufband-belastung.”
From gains to liver pain: when exercise training goes too far
Burke, B. I., McCarthy, J. J., & Ismaeel, A. (2025) · Extracellular Vesicles and Circulating Nucleic Acids · doi:10.20517/evcna.2025.36 · Cited by 1
Profiling skeletal muscle-derived secretome with differentiation and acute contractile activityPreprint
Bydak, B., Pierdoná, T. M., Seif, S., Sidhom, K., Obi, P. O., Labouta, H. I., Gordon, J. W., & Saleem, A. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.02.20.481208 · Cited by 1
Fernández‐Rhodes, M., Adlou, B., Williams, S., Jalal, A. R., Lewis, M. P., & Davies, O. G. (2022) · Research Square · doi:10.21203/rs.3.rs-2041930/v1 · Cited by 1
Mangolini, V., Radeghieri, A., Piva, S., Cattaneo, S., Brucale, M., Valle, F., Balestri, A., Montis, C., Latronico, N., Bergese, P., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.02.19.580950 · Cited by 1
Via, J. O. (2018) · JMU Scholoraly Commons (James Madison University)
“Brainstorming”: Extracellular Vesicles in Physical Activity and Neuronal Health
Brahmer, A., & Krämer‐Albers, E. (2020) · Trillium Extracellular Vesicles · doi:10.47184/tev.2020.01.07 · Cited by 1
Dietary Protein and Muscle in Aging People
Cesari, M., & Marzetti, E. (2018) · MDPI eBooks · doi:10.3390/books978-3-03897-458-1
Metadata
- Authors
- Guescini, M., Canonico, B., Lucertini, F., Maggio, S., Annibalini, G., Barbieri, E., Luchetti, F., Papa, S., & Stocchi, V.
- Affiliations
- University of Urbino (IT)
- Corresponding author
- Guescini, Michele
- ORCID
- Venue
- PLoS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2015-05-08
- Volume
- 10
- Issue
- 5
- Pages
- e0125094
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC4425492
- OpenAlex
- W219060072
- Cited by
- 228
- References
- 62
- Citations by year
- 2026: 19 · 2025: 31 · 2024: 26 · 2023: 27 · 2022: 33 · 2021: 24 · 2020: 22 · 2019: 18 · 2018: 11 · 2017: 11 · 2016: 5 · 2015: 1
- FWCI
- 4.05
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- PLoS ONE (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Figshare (submittedVersion, cc-by)
- CINECA IRIS Institutional Research information system (University of Urbino) (submittedVersion, cc-by)
- Funders
- Regione Marche
- Università degli Studi di Urbino Carlo Bo
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Extracellular vesicles in disease
- Health Sciences › Medicine › Physiology (medical) › Exercise and Physiological Responses
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Muscle Physiology and Disorders
- Keywords
- extracellular vesicles, alpha-sarcoglycan, miR-206, microRNA secretion, myokines, skeletal muscle, physical activity, cardiorespiratory fitness, miR-133b, miR-181a-5p, TSG101, CD81, plasma vesicles, muscle remodeling, muscle homeostasis
- MeSH terms
- Extracellular Vesicles, Tsg101 Protein, Adult, Cell Communication, Centrifugation, Density Gradient, DNA-Binding Proteins, Gene Expression Regulation, Humans, Male, Transcription Factors, Signal Transduction, Exercise, Muscle, Skeletal, MicroRNAs, Sarcoglycans, Exosomes, Endosomal Sorting Complexes Required for Transport, Sedentary Behavior, Tetraspanin 28
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