In vitro and in vivo studies demonstrated that myostatin-deficient mice exhibited a significantly increased muscle mass characterized by an increased cross-sectional area and an elevated number of fibers. National Library of Medicine Myostatin is a member of the transforming growth factor (TGF)- superfamily and it negatively regulates SKM growth and mass [82]. One study in Duchennes patients, where only two doses were received due to epistaxis and telangiectasias showed trends to improved muscle function (Campbell et al., 2017). Some myokines that are also adipokines have also been named adipomyokines, and these molecules will be presented in the next chapter. 2016 Nov;54(2):284-305. doi: 10.1007/s12020-015-0834-0. ScienceDirect is a registered trademark of Elsevier B.V. ScienceDirect is a registered trademark of Elsevier B.V. Progress in Molecular Biology and Translational Science, The Cellular Secretome and Organ Crosstalk, Lambernd etal., 2012; Nikolic etal., 2012, Thermoregulation: From Basic Neuroscience to Clinical Neurology Part I, Inflammatory and Oxidative Stress Markers in Skeletal Muscle of Obese Subjects, Myokines: The endocrine coupling of skeletal muscle and bone, Potthoff et al., 2009; Domouzoglou and Maratos-Flier, 2011, ). A picomole (pM) is 10 mol/L. (2006) showed that IL-4 was secreted during human myoblast differentiation and is required for myotube maturation. Cytokines are signaling proteins. Jenny E. Gunton, in Encyclopedia of Endocrine Diseases (Second Edition), 2019. Myostatin was overexpressed in bone tissue of a diabetic mouse model with a tibial defect. Other studies suggest that plasma BDNF is a biomarker of impaired memory and general cognitive function in ageing women and a low circulating BDNF level was recently shown to be an independent and robust biomarker of mortality risk in old women. [10], an autoantibody-stimulating model of PE demonstrated that antibodies isolated from PE-affected mothers could stimulate IL-6 from human mesangial cells [10]. noted that over 1,000 genes are upregulated by exercise and examined how expression of FNDC5 was affected by exercise in ~200 humans. Rather, BDNF produced in skeletal muscle appears to enhance the oxidation of fat. Mol Biol Rep. 2023 Mar;50(3):2723-2734. doi: 10.1007/s11033-022-07821-3. Careers. There are several animal models of myostatin deficiency, including mice, dogs and cattle. write: "It appears consistently in the literature that IL-6, produced locally by different cell types, has a positive impact on the proliferative capacity of muscle stem cells. 2023 May;13(5):613-619. doi: 10.5455/OVJ.2023.v13.i5.14. [7], As studies have demonstrated IL-6 has pro-inflammatory functions when evaluated in regard to sepsis and obesity, it was initially hypothesized that the exercise-induced IL-6 response was related to muscle damage. In 2008, the first myokine, myostatin, was identified. Interleukin-4 (IL-4) is a complex glycoprotein produced mostly by mast cells, basophils, a subset of activated T cells, eosinophils and neutrophils (Chomarat and Banchereau, 1997). It is suggested that the term myokine is restricted to proteins secreted from skeletal muscle cells, excluding proteins that are secreted by other cell types in skeletal Myostatin is naturally antagonized by follistatin, another TGF superfamily member, and one of the most cost-effective inhibitor of myostatin [91]. Recently, it has been proposed, in an in vitro study on mouse osteocytes (Ocy454), that myostatin acts on osteocytes inducing upregulation of osteogenic factors such as SOST, DKK-1 and the receptor activator of nuclear factor kappa- ligand (RANKL), important regulators of bone remodeling [97]. If it has chosen the abnormal path, i.e. Background In recent years, several studies have revealed a new role for skeletal muscle as a secretory organ since its contraction has emerged as an important activator of the release of proteins and peptides called myokines. This last finding may explain the concomitant finding of low circulating levels of BDNF in individuals with type 2 diabetes, and the association between low plasma BDNF and the severity of insulin resistance. Association between muscle strength and mass and bone mineral density in the US general population: data from NHANES 1999-2002. The nadir in expression was 8h after exercise. These genes might be considered as upstream intermediates of differential expression of late response genes including myokines whose expressions were slightly increased by 2-h electrical pulse stimulation (Sidorenko et al., 2018) and further potentiated after prolongation of EPS to 48h (Nedachi et al., 2008; Nikoli et al., 2012; Whitham et al., 2012). Victoria Cataln, Javier Gmez-Ambrosi, in Obesity, 2018. It is still unclear whether the muscular release of myonectin is caused by exercise itself or due to intake of nutrients (carbohydrates and lipids) following the exercise program. Indeed, muscle-produced IL-6 after repeated contractions also has important autocrine and paracrine benefits, acting as a myokine, in regulating energy metabolism, controlling, for example, metabolic functions and stimulating glucose production. It was suggested that myostatin regulates the mechanosensitivity of bone marrow-derived MSCs (BMSCs) by suppressing osteogenic differentiation during mechanical stimulation and that myostatin deficiency does not prevent bone catabolism with unloading [96]. Regulation of IL-6 production has also been demonstrated to be altered in women with early onset PE. Decorin is released from contracting human myotubes, and circulating decorin levels are increased in response to acute resistance exercise in humans. They conclude that these data support the view that irisin may have a role in the modulation not only of energy balance but also the aging process. sharing sensitive information, make sure youre on a federal We applied the level of evidence proposed by the Oxford Center for Evidence-Based Medicine to determine the cause-effect relationship of exercise with circulating level of myokine and of myokine with the brain function, as well as exercise While the murine findings reported by Bostrm et al. This site needs JavaScript to work properly. 2019 Mar;471(3):491-505. doi: 10.1007/s00424-019-02253-8. In fact, many myokines are also produced by adipocytes and we propose to name them adipo-myokines. [9], "Some myokines exert their effects within the muscle itself. In addition, we have demonstrated that there is a cerebral output of BDNF and that this is inhibited during hyperglycaemic clamp conditions in humans. Bookshelf 2023 Apr 3;9(1):30. doi: 10.1038/s41526-023-00273-4. WebMyokines: The endocrine coupling of skeletal muscle and bone. The authors enquired whether the favorable impact of irisin on white adipose tissue might be associated with maintenance of telomere length, a well-established genetic marker in the aging process. MeSH eCollection 2023 Apr. IL-6 is a pleiotropic myokine expressed in endothelial cells, fibroblasts, macrophages, monocytes, and T cells as well as myocytes, hepatocytes and adipocytes. In another study, Shanaki et al. Check out the pronunciation, synonyms and grammar. We use cookies to help provide and enhance our service and tailor content and ads. In vivo overexpression of decorin in murine skeletal muscle promoted expression of the pro-myogenic factor Mighty, which is negatively regulated by myostatin. They showed that myostatin is involved in osteoclastogenesis and bone resorption in RA, and that myostatin deficiency or antibody-mediated inhibition of myostatin result in an improvement of the arthritis. Zhang et al. A myokine is a cytokine or signaling protein from the skeletal muscles. importance on measuring release from muscle, in addition to monitoring changes in gene. Taken together, the analysis of conditioned media from EPS models has generated additional knowledge on the complex scenario of skeletal muscle as an endocrine organ. Considering this, it might be proposed that along with the above-listed signaling pathways, elevation of [Na+]i/[K+]i-ratio detected in exercising skeletal muscle and EPS-treated myotubes contributes to myokine production (Kapilevich et al., 2015). Molecular Communication from Skeletal Muscle to Bone: A Review for Muscle-Derived Myokines Regulating Bone Metabolism. Unauthorized use of these marks is strictly prohibited. Osteonectin can be classed as a myokine, as it was found that even a single bout of exercise increased its expression and secretion in skeletal muscle in both mice and humans. WebTranslate Myokine. Importantly, the same genes were also found in ouabain-treated C2C12 cells (Supplementary Table S1 in the online version at https://doi.org/10.1016/bs.ctm.2019.01.006), thus suggesting an impact of Na+i/K+i-mediated, Ca2+-independent excitation-transcription coupling in myokine production. [14] demonstrated significant elevations in both IL-6 and receptor in both cohorts of women. This is considered important, because excess visceral adipose tissue in particular distorts the whole body energy homeostasis, increases the risk of cardiovascular disease and raises exposure to a milieu of adipose tissue-secreted hormones (adipokines) that promote inflammation and cellular aging. WebMyokine. In a deep penetrant injury mouse model, Elkasrawy and co-workers analyzed the space-temporal localization of myostatin expression during muscle and bone repair. In fact, irisin treatment has been shown to attenuate reactive oxygen species-induced hepatic cell damage [151]. However, the muscle secretome is still insufficiently characterized. WebLook through examples of myokine translation in sentences, listen to pronunciation and learn grammar. One recently published approach (Catoire etal., 2014b) is based on the exvivo analysis of human skeletal muscle biopsies taken before and after acute exercise and exercise training, followed by microarray-based analysis of secreted proteins. Although a variety of EPS protocols are currently used that may explain a number of divergent findings, it is generally accepted that EPS of human muscle cells is a valid model to study contraction-mediated processes and adaptation to exercise training. Interestingly, the exercising limb was not the source of follistatin net release, suggesting an alternative source of follistatin other than contracting SKM [7]. Sample translated sentence: myokines Myokine {pl} [noun] [biochem.] As will be discussed in the next chapter, many of these myokines are also adipokines and are produced by adipose tissue. BDNF has been shown to regulate neuronal development and to modulate synaptic plasticity. Calculating femoral arterial-venous Na+ and K+ concentration differences, it was found that in humans intense knee extensor exercise rapidly increased intracellular Na+ concentration ([Na+]i) in skeletal muscle from 13 to 23mM and reduced [K+]i from 162 to 129mM (Sjgaard, Adams, & Saltin, 1985). An additional limitation of the EPS model is certainly the lack of work load in this system and inability to fully mimic the complex scenario of invivo muscle contraction. These cells show adaptation to the continuous contraction by reorganization of the cytoskeleton, de novo formation of sarcomeric structures, increased glucose uptake and insulin sensitivity, and increased mitochondrial content. Moreover, decorin expression in skeletal muscle is increased in humans and mice after chronic training. indicates that TNF increase in response to sepsis is inhibited by the exercise-induced production of, IL-6, among an increasing number of other recently identified, B. K. Pedersen and her colleagues have conducted research showing that skeletal muscle functions as an endocrine organ by secreting cytokines and other peptides, now referred to as, In particular, the increase in levels of IL-6 (interleukin 6), a, That is, contracting muscles release multiple substances known as, Adipose tissue and skeletal muscle produce adipokines and. [16] However, a recent study suggests that eccentric exercise is not associated with a larger increase in plasma IL-6 than exercise involving concentric nondamaging muscle contractions. showed no differences in irisin concentration in patients with obesity, simple steatosis and NASH, whereas irisin was increased in lean subjects [154]. Different muscle fiber types slow twitch muscle fibers, oxidative muscle fibers, intermediate twitch muscle fibers, and fast twitch muscle fibers release different clusters of myokines during contraction[citation needed]. Many adipo-myokines produced by skeletal muscle or adipose tissue are influenced by exercise. Myokines work on fat cells, liver, pancreas, bone marrow, heart, white blood cells, immune cells, brain cells, and even stem cells. Kim S, Choi JY, Moon S, Park DH, Kwak HB, Kang JH. demonstrated that increased IL-6 and CRP were significantly associated with presence of fatty liver as assessed by computed tomography (CT) scan after adjusting for other CVD risk factors in a population study composed of four ethnic groups [150]. A beneficial effect of BDNF on neuronal function has been noted in multiple studies. Toward A Clinical Definition. 2020 Elsevier Inc. All rights reserved. This observation remains somewhat controversial with respect to the extent to which irisin concentration increases after exercise. PMC The https:// ensures that you are connecting to the In contrast, heavy endurance efforts, such as ultramarathon, were associated with a drastic decrease of follistatin levels and consequently with increased levels of myostatin [4]. These findings highlight the therapeutic potential of myostatin inhibition for the treatment of RA-associated bone resorption and joint destruction and potentially other diseases characterized by an increased myostatin expression. Bethesda, MD 20894, Web Policies [15], Aerobic exercise provokes a systemic cytokine response, including, for example, IL-6, IL-1 receptor antagonist (IL-1ra), and IL-10 (Interleukin 10) and the concentrations of chemokines, IL-8, macrophage inflammatory protein (MIP-1), MIP-1, and MCP-1 rise after vigorous exercise. government site. Accessibility and transmitted securely. en.wiktionary.2016 Pronunciation of Myokines with 2 audio pronunciations 1 rating 0 rating Record the pronunciation of this word in your own voice and play it to listen to how you Yu C, Du Y, Peng Z, Ma C, Fang J, Ma L, Chen F, Zhang C, Geng R, Zhang Y, Han X, Li J, Lv Y, Lu S. Exp Ther Med. This hypothesis is supported by several observations. [7][8], There is an emerging understanding of skeletal muscle as a secretory organ, and of myokines as mediators of physical fitness through the practice of regular physical exercise (aerobic exercise and strength training), as well as new awareness of the anti-inflammatory and thus disease prevention aspects of exercise. Enter your email address to subscribe to this blog and receive notifications of new posts by email. Hye Soo Chung, Kyung Mook Choi, in Advances in Clinical Chemistry, 2020. eCollection 2023. [19][21] Dr. Pedersen writes, "Neurotrophins are a family of structurally related growth factors, including brain-derived neurotrophic factor (BDNF), which exert many of their effects on neurons primarily through Trk receptor tyrosine kinases. Recently, we reported that electrical pulse stimulation (EPS) of C2C12 myotubes resulted in about fivefold elevation of [Na+]i/[K+]i-ratio (Danilov et al., 2017). Editorial: Crosstalk between bone and other cells. A myokine is a cytokine or signaling protein from the skeletal muscles. After secretion into the circulation, myostatin dimerizes by forming a disulfide bond within the C-terminal domain. Mice with IL-6 deficiency or deletion of hepatic glycoprotein 130 presented with increased hepatic triglycerides, insulin resistance, liver inflammation and liver steatosis [147]. Also, patients with major depression have lower levels of serum BDNF than normal control subjects. However, a direct effect of myostatin on osteoblasts remains to be elucidated. 2023 May 9;14:1209053. doi: 10.3389/fphys.2023.1209053. Grit E. Legrd, Bente K. Pedersen, in Muscle and Exercise Physiology, 2019. 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. The skeletal muscles produce hundreds of different kinds of myokines. WebLearn the definition of 'myokine'. The expression of FNDC5 is under the control of the peroxisome proliferator-activated receptor coactivator 1-, a master regulator of genes involved in metabolism. Listen to the audio pronunciation of Myokines on pronouncekiwi Several studies have shown that irisin possesses protective properties against obesity, insulin resistance, and nonalcoholic fatty liver disease, showing in some of these metabolic alterations a correlation with inflammatory markers, which suggests that irisin may regulate the inflammatory response. Myostatin activates also the p38/MAPK signaling, probably through interaction with ActRIIB, the Erk1/2 MAPK pathway via Ras, the PI3K/Akt/GSK-3 pathway and the JNK signaling pathway, altogether indicating that myostatin can act on different intracellular mediators [83,84]. Indeed, it was further demonstrated that the primary source of follistatin is the liver, as mice subjected to a bout of swimming exercise revealed increased levels of follistatin mRNA and protein in this organ, correlated to elevated plasma levels. Early onset PE (<32weeks of gestation) showed higher elevations of IL-6 compared with both late onset PE and normal pregnancy (40.803.51 early PE vs 33.561.63 late PE vs. 25.971.07 control; units in pg/mL). [24] FNDC5 was initially discovered in 2002 by two independent groups of researchers. The signals may be to themselves or the same cell (autocrine), nearby cells (paracrine), and distant organs (endocrine). However, Polyzos et al. Research advances in crosstalk between muscle and bone in osteosarcopenia (Review). WebIrisin is a myokine released into the bloodstream by cleavage of the fibronectin type III domain-containing protein 5 (FNDC5) triggered by muscle contraction. Implications of exercise-induced adipo-myokines in bone metabolism. IL-6s role in mouse and cellular models of PE has been recapitulated in human clinical data. In addition to differential expression in PE patients and levels of IL-6 correlating with disease severity in PE, it plays a pivotal role in placentally mediated vasoconstriction. WebLearn the definition of 'myokines'. In humans, a BDNF release from the brain was observed at rest and increased 2- to 3-fold during exercise. They found that it was upregulated only in highly active elderly humans, casting doubt on the conclusions of Bostrm et al. Wieckowska et al. [2] The location of these receptors reflects the fact that myokines have multiple functions. To further identify the putative cerebral region(s) responsible for BDNF release, mouse brains were dissected and analysed for BDNF mRNA expression following treadmill exercise. One explanation for the increased irisin expression with exercise in mouse and man may have evolved as a consequence of muscle contraction during shivering. Myokines are implicated in the autocrine regulation of metabolism in muscles as well as in the para/endocrine regulation of other tissues and organs including the adipose tissue, liver, and brain ( Carson, 2017) through their receptors. Disclaimer. [22], IL-6 levels in PE serum had an inverse correlation with NO levels with a decrease in NO production by ~33% in PE serum versus normal pregnant patients. The .gov means its official. In NAFLD with histologic confirmation, irisin expression in serum and liver was positively associated with disease severity, i.e., steatosis grade, fibrosis severity and progression to NASH [152]. In HepG2 human liver carcinoma cells, irisin enhanced cell survival via AMPK activation [155]. In this context, an interesting study using mass spectrometrybased proteomics showed that lipid-induced insulin resistance affected nearly one-half of the secreted proteins (Deshmukh etal., 2015). Selected Myokines With Proven Function in Humans, Jan Nedergaard, Barbara Cannon, in Handbook of Clinical Neurology, 2018. They reported on July 1, 2014: "The small leucine-rich proteoglycan decorin has been described as a myokine for some time. Stimulated by both T-cells and macrophages, IL-6 is elevated in response to microbial pathogens and TNF stimulation. How to say myokine in English? Myokines exert an autocrine function in regulating muscle metabolism as well as a paracrine/endocrine regulatory function on distant organs and tissues, such as bone, adipose tissue, brain and liver. Think of cytokines as text messages from one organ to another organ. Low levels of circulating BDNF are also found in obese individuals and those with type 2 diabetes. This implies that variation of exercise types, particularly aerobic training/endurance training and muscle contraction against resistance (strength training) may offer differing myokine-induced benefits. Within the liver, irisin has been identified in hepatocytes, sinusoidal endothelial cells and Kupffer cells by immunohistochemical detection [396]. The therapeutic potential of irisin is obvious. A model of PE was recapitulated in pregnant mice. Skeletal muscle encompasses one of the major Multiple studies have demonstrated a potential role for IL-6 in placental ischemia and vascular damage. This hypothesis is consistent with profound accumulation of IRGs in skeletal muscle biopsies obtained from humans after intensive resistance exercises (Broholm et al., 2011). Picomoles are enough. Kanzleiter et al have established that this myokine is secreted during muscular contraction against resistance, and plays a role in muscle growth. FOIA Healthy subjects performing acute bicycle exercise showed up to sevenfold increase in follistatin plasma levels. The myokine BDNF has only auto- and paracrine functions, as far as is currently known . These findings suggest that pharmacological inhibition of myostatin early after musculoskeletal injury might enhance the muscle and bone regenerative potential [90]. To study the control of myokine regulation of -cell function, SkM biopsies were taken from non-diabetic (ND) and Type 2 diabetic (T2D) subjects and satellite cells cultured to myotubes (MT). The skeletal WebMyokine is anti-inflammation cytokine and peptide produced by striated muscles. Research about myokines is just beginning to describe their functions and interactions. From: Progress in Molecular Biology and 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-1alpha-dependent myokine irisin, which drives brown fat-like development. Third, we recently reported that in C2C12 cells short-term electrical pulse stimulation triggers expression of potent transcription regulators including immediate response genes such as Fos, Egr1, Myc, Egr2, Atf3, Fosl1, Fosb, 12 histone clusters, and 29 microRNAs (Sidorenko et al., 2018). Whether longer treatments with irisin and/or higher doses would cause more weight loss remains to be determined. Myostatin enhances RANKL-mediated osteoclastogenesis by activating the nuclear factor of activated T-cells (NFATc1). Webmyokine translation in English - English Reverso dictionary, see also 'moki, myogenic, moniker, moline', examples, definition, conjugation "Most recently, we have shown that BDNF appears to be a major player not only in central metabolic pathways but also as a regulator of metabolism in skeletal muscle. The trials of inhibitors have mostly been of short duration, and or small in size, and there are no reports in people regarding whether there are or are not bone effects. Finally, MCP-1 and fractalkine were found to be increased at both the mRNA and the plasma level, and these myokines could play a yet-to-be-defined role in the communication between skeletal muscle and other organs. A novel myokine osteonectin, or SPARC (secreted protein acidic and rich in cysteine), plays a vital role in bone mineralization, cell-matrix interactions, and collagen binding. Recently, we reported that electrical pulse stimulation (EPS) of C2C12 myotubes resulted in about fivefold elevation of [Na+]i/[K+]i-ratio (Danilov et al., 2017). Being recognized as an endocrine organ itself, skeletal muscle secretes a panel of cytokines and proteins named myokines, synthesized and secreted by myocytes in response to muscle contraction. Lafreniere et al. myokine (plural myokines) (biochemistry) A form of cytokine produced in muscle cells. It has been shown that inhibition of the active myostatin with a recombinant myostatin pro-peptide induces muscle regeneration and bone repair in a mouse model of fibula osteotomy with associated damage to lateral compartment muscles (fibularis longus and brevis) [88]. Malhan D, Yalin M, Schoenrock B, Blottner D, Relgio A. NPJ Microgravity. Myostatin also reduces the miR-218 content in osteocytes-derived exosomes. WebIn the last years, scientists have shown that skeletal muscle is not a pure locomotor unit or responsible for propulsion and posture. Roles of myokines in exercise-induced improvement of neuropsychiatric function. First, sustained inhibition of Na+,K+-ATPase affects expression of hundreds of genes in the presence of extracellular and intracellular Ca2+-chelators (Koltsova et al., 2012) and inhibitors of Ca2+i-mediated signaling pathways (Smolyaninova, Koltsova, Sidorenko, & Orlov, 2017). Myokin Belgian blue cow By agriflanders originally posted to Flickr as Kamp- Bambino vd ijzer copy, CC BY 2.0, https://commons.wikimedia.org/w/index.php?curid=6257487. An official website of the United States government. The authors also summarize several previously-established research findings relating to exercise and brain function: (1) Aerobic exercise training increases grey and white matter volume in the prefrontal cortex of older adults and increases the functioning of key nodes in the executive control network. Level of evidences for the causative relationship between exercise, myokine, and brain function. Would you like email updates of new search results? IL-6 is also the founding member of the myokine family of muscle-produced cytokines. Considering this, it might be proposed that along with the above-listed signaling pathways, elevation of [Na+]i/[K+]i-ratio detected in exercising skeletal muscle and EPS-treated myotubes contributes to myokine production (Kapilevich et al., 2015). Myokine definition: Collins Dictionary Definition | Meaning, pronunciation, translations and examples in American English Myokine, a key cytokine for physical exercise to alleviate sarcopenic obesity. Using flame photometry, Murphy and coworkers found that in rat soleus muscle 60min of intermittent running resulted in about twofold elevation of [Na+]i and attenuation of [K+]i by 1020% (Murphy, Nielsen, & Clausen, 2008). A direct effect of myostatin on osteoclasts has also been proposed. Therefore, it is likely that adipo-myokines may contribute in the mediation of the health benefits of exercise and physical inactivity probably leads to an altered adipo-myokine profile, which could provide a potential mechanism for the association between sedentary behavior and many chronic diseases. [3][4], Receptors for myokines are found on muscle, fat, liver, pancreas, bone, heart, immune, and brain cells. This literature appears to be contentious however, as a subsequent analysis of both a Brazilian and Chinese cohort found no association with IL-6 promoter polymorphism and PE risk [1820]. Myonectin belongs to the C1q/TNF-related protein (CTRP) family and is predominantly expressed by skeletal muscle. These data show that skeletal muscle cells have a prominent basic secretory capacity, principally not different from adipocytes. Thus, presently hundreds of putative myokines have been identified and described to be regulated at the mRNA or secretory level by differentiation, contraction, or other stimuli. expression [6]. Keywords: This study combined genomics, three different mass spectrometry methods, and multiplex immunoassays to generate a comprehensive characterization of the entire human muscle secretome. [2] For instance, aerobic exercise in humans leads to significant structural alterations in the brain, while wheel-running in rodents promotes neurogenesis and improves synaptic transmission in particular in the hippocampus. Secretomic studies to identify myokines released from skeletal muscle or isolated myocytes kept in culture have therefore gained considerable interest, and at present more than 600 myokines have been described (Gorgens etal., 2015). Myonectin has been identified as a novel myokine that links skeletal muscle to systemic lipid homeostasis (Seldin et al., 2012). On the other hand, loss-of-function mutations or deletion of the myostatin gene are associated with muscle hypertrophy and hyperplasia correlated to doubling of muscle mass in several mammals such as mice, dogs, sheep, cattle and humans [86]. Increased IL-6 within the normal physiologic range may be helpful for regulation of metabolic disease, infection defense, hepatic tissue homeostasis and liver regeneration [149,395].
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