. 2020 Jan 2;2(1):100019. doi: 10.1016/j.ocarto.2019.100019

Table 2.

Summary of recent mass spectrometry-based studies carried out on human skeletal muscle cells to identify secretome components. Classically secreted proteins contain a signal sequence, unlike non-classically secreted proteins.

ECM Proteins	Function	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
Basement membrane-specific heparan sulfate proteoglycan core protein (Perlecan)	component of basement membranes	Classical		↑	↑	↓		↓	⁶^,²⁷^,²⁶^,²⁸^,³¹^,³³^,³⁶^,³⁷^,⁴⁰
Biglycan	collagen fiber assembly	Classical		↑		↓		↓	²⁵^,⁴⁰^,²⁷^,²⁸^,²⁶^,⁶^,35, 36, 37, 38
Collagen I(α1, α2), II(α1), III(α1), IV(α1,α2,α3), V(α1,α2,α3), VI(α1,α2,α3), VII(α1), VIII(α1), IX(α3), XI(α1,α2), XII(α1), XIII(α1), XIV(α1), XV(α1), XVIII(α1)	cell adhesion	Classical		III(α1), XVIII(α1) ↓ I (α1), II(α1), V (α1, α3) VI(α1), XI (α1) ↑		IV(α2)↑	VI(α1)↑ V (α2)↓ III(α1)↓	I (α1, α2)↓ III(α1)↓ IV(α1, α2)↓ V (α1, α2)↓ VI(α1,α2)↓ VIII(α1,α2)↓ XI (α1)↓ XII(α1)↓ XV (α1)↓	³⁰^,²⁹^,³¹^,²⁷^,⁶^,³³^,³²^,34, 35, 36, 37, 38, 39^,²⁶
Decorin	affects the rate of fibril formation	Classical		↑				↓	²⁷^,²⁸^,²⁶^,⁶^,³⁵^,³⁶
Dystroglycan	laminin and basement membrane assembly, cell survival and migration	Classical		↑				↓	²⁷^,²⁸^,²⁶^,³³^,³⁵^,³⁶^,³⁸^,³⁹
Extracellular matrix protein 1, 2	Angiogenesis, Biomineralization, Mineral balance, Osteogenesis	Classical						2 ↓	²⁹^,²⁸^,⁶^,34, 35, 36^,³⁹
Fibrillin 1, 2	Structural component of microfibrils	Classical		1 ↑		1↓			²⁵^,²⁷^,²⁸^,²⁶^,⁶^,³⁵^,³⁶
Fibromodulin	affects the rate of fibril formation	Classical		↑				↓	³⁶^,²⁷
Fibronectin	Cell adhesion, cell shape	Classical			↑			↓	³²^,²⁹^,²⁷^,²⁸^,⁶^,³⁴^,36, 37, 38, 39^,²⁴
Fibulin 1, 2, 5, 7	cell–cell interaction, cell migration, ECM remodelling	Classical		1 (isoform C) ↓ 1 (isoform D), 2 (isoform B), 5↑				1, 2, 5 ↓	³¹^,²⁷^,²⁸^,²⁶^,⁶^,³³^,³²^,³⁵^,³⁶^,³⁹
Glypican 1, 6	Role in skeletal muscle differentiation	Classical		1↑				1↓	²⁹^,²⁷^,²⁸^,⁶^,³³^,³⁵^,³⁷^,³⁹^,⁴⁰
Laminin α (1, 2, 3, 4, 5), β (1, 2), γ (1,2)		Classical		α5, α2, β2 ↓				α(4,5), β1, γ1↓	⁶^,²⁷^,²⁸^,³⁰^,³³^,35, 36, 37
Latent-transforming growth factor beta-binding protein 1(S, L), 2, 3, 4	growth factor binding	Classical		3↑				1, 2, 3, 4↓	⁶^,26, 27, 28^,³⁵^,³⁶
Lumican	Collagen binding	Classical						↓	⁶^,²⁸^,²⁹^,³⁶
Matrilin 2, 3	involved in matrix assembly	Classical						2↓	26, 27, 28^,³⁵^,³⁶
Matrix Gla Protein		Classical							²⁷^,³³^,³⁵^,³⁶
Mimecan	Induces bone formation in conjunction with TGF-β-1 or TGF-β-2.	Classical		↓				↓	²⁶^,²⁷^,³¹^,³²^,³⁵^,³⁶
Moesin	connection cytoskeletal structure - plasma membrane				↑				⁶^,²⁷^,⁴⁰^,²⁸^,²⁹^,³¹^,34, 35, 36^,³⁸^,³⁹
Nidogen 1, 2		Classical						1,2↓	⁶^,26, 27, 28^,³³^,³⁵^,³⁶^,38, 39, 40
Periostin	cell adhesion	Classical		↑		↑		↓	⁶^,25, 26, 27, 28^,³⁵^,³⁶
Prolargin		Classical		↑				↓	²⁶^,²⁷^,³⁶
Protein S100 (A4, A6 (calcyclin), A11, A13, A16)		Classical					A6↓		⁶^,²⁸^,³⁰^,33, 34, 35, 36^,³⁸^,³⁹
Proteoglycan 4	Role in joint lubrication			↑					⁶^,²⁷^,³⁵
Secreted protein acidic and rich in cysteine (SPARC)	Regulates cell growth, binds calcium and copper	Classical		↑ and ↓	↑			↓	⁶^,²⁶^,36, 37, 38, 39, 40^,27, 28, 29, 30, 31, 32^,³⁴^,³⁵
Sushi, von Willebrand factor type A, EGF and pentraxin domain-containing protein 1	cell attachment process	Classical		↓				↓	⁶^,³⁵^,³⁷
Syndecan 2, 4	proteoglycan that bears heparan sulfate	Classical							²⁷^,²⁸^,31, 32, 33^,³⁶
Tenascin C, XB, X	Cell adhesion	Classical		C ↑				↓	⁶^,26, 27, 28^,³⁰^,³⁵^,³⁶
Vitronectin	Cell adhesion and spreading factor	Classical							⁶^,³⁶
*Cytokines and growth factors*	Specificity	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
Anamorsin (Ciapin 1)	Apoptosis, metal binding	Non-classical		↑					²⁸^,³¹^,³²^,³⁵^,³⁶
Angiopoietin 1, 4	angiogenesis, cell proliferation, adhesion and migration	Classical		1↑				4↓	²⁷^,²⁸^,³⁵^,³⁶
Bone morphogenetic protein 1, 4	Cytokine, growth factor, protease, metalloprotease	Classical					1 ↓	1 ↓	26, 27, 28^,³⁰^,³⁵^,³⁶
Brain derived neurotrophic factor	regulates skeletal muscle metabolism	Classical							²⁸^,³⁶
C–C motif chemokine 2, 3, 5, 7, 8, 9	cytokine, inflammatory response, chemotaxis	Classical	2↑	2, 7, 8↑				2↑ 9↓	⁶^,²⁷^,²⁸^,³⁰^,³⁵^,³⁶
Cell adhesion molecule-related/down-regulated by oncogenes	Promotes differentiation of myogenic cells	Classical		↑				↓	²⁶^,²⁷^,³⁵^,³⁶
Complement C1q tumor necrosis factor-related protein 1, 3, 5		Classical		3, 5↑				1, 3 ↓	26, 27, 28^,³³^,35, 36, 37
Connective tissue growth factor	Cell adhesion, DNA synthesis, growth factor activity	Classical		↑			↓	↓	⁶^,26, 27, 28^,³⁵^,³⁶^,³⁸^,³⁹
C-X-C motif chemokine (CXCL) 1, 2, 5, 6, 10, 12, 16	Cytokine, growth factor	Classical		5↓				1, 5↑	⁶^,²⁷^,²⁸^,³²^,³⁵^,³⁶
Fibroblast growth factor 17, 21	Mitogenic and cell survival activities	Classical		21↑					²⁷^,30, 31, 32^,³⁵^,³⁶
Follistatin-related protein 1, 3	Heparin binding, modulate action of some growth factors	Classical		1 ↓, 3 ↑			1↓	1↓	⁶^,²⁶^,37, 38, 39^,27, 28, 29^,³¹^,³²^,34, 35, 36
Galectin 1, 3, 9	regulation of cell proliferation and migration + bind galactose	Classical	1↑	1, 3↑	1↑			9↑	⁶^,²⁶^,³⁹^,⁴⁰^,²⁷^,²⁸^,³⁰^,34, 35, 36, 37, 38
Glucose-6-phosphate isomerase	Cytokine, growth factor, isomerase	Classical			↑				⁶^,²⁴^,³⁹^,26, 27, 28, 29^,34, 35, 36^,³⁸
Granulins	Cytokine, role in inflammation and tissue remodeling	Classical						↓	26, 27, 28^,³⁵^,³⁶
Granulocyte colony-stimulating factor	cytokine, growth factor, role in hematopoiesis								⁶
Gremlin 1	Cytokine	Classical							²⁸^,³³^,³⁵^,³⁶
Growth/differentiation factor 8 (myostatin), 11, 15	Regulates food intake, energy expenditure and body weight	Classical						11↓	²⁷^,²⁸^,³⁰^,³⁵^,³⁶
Hepatocyte growth factor	Mitogen for hepatocytes cells, growth factor	Classical						↓	²⁷^,³⁵^,³⁶
Hepatoma-derived growth factor	Growth factor, repressor	Non-classical		↓				↑	³⁷^,³⁹
High mobility group protein B1, B2, B3	Immunity, autophagy, chemotaxis	Non-classical		B1 ↑ and ↓					²⁷^,³²^,35, 36, 37
Insulin-like growth factor 1, 1A, 2		Classical		1,2↑	2↑		2↓	1↓	²⁶^,²⁷^,³⁰^,34, 35, 36
Insulin-like growth factor binding protein(IGFBP) 2, 3, 4, 5, 6, 7	regulation of cell proliferation	Classical		2↓ 4↑ 5↑ and ↓			6↓	2, 5, 6, 7↓ 4 ↑ and ↓	⁶^,²⁶^,³⁹^,⁴⁰^,²⁷^,²⁸^,³¹^,³²^,35, 36, 37, 38
Interleukin 1β, 2, 4, 6, 7, 8, 10, 13, 17A, 25, 34		Classical							⁶^,²⁸^,³⁵^,³⁶
Macrophage colony-stimulating factor 1	Cytokine, growth factor	Classical				↓			⁶^,²⁵^,²⁶^,²⁸^,²⁹^,³⁵^,³⁶
Macrophage migration inhibitory factor	cytokine, isomerase, immunity	Classical							⁶^,²⁷^,³⁸^,³⁹^,²⁸^,²⁹^,31, 32, 33^,35, 36, 37
Osteoclast-stimulating factor 1	induces bone resorption								³⁵^,³⁶
Placenta growth factor	angiogenesis, differentiation	Classical							^28,3536
Platelet-derived growth factor A, C	Developmental protein, mitogen, growth factor	Classical		A, C↑				C ↓	²⁷^,³⁵^,³⁶
Prosaposin (sulfated glycoprotein 1)	Lipid metabolism, growth factor activity			↑			↑		⁶^,26, 27, 28^,³¹^,35, 36, 37
Secreted frizzled-related protein 2, 4	Developmental protein, regulation of cell growth and differentiation	Classical						2 ↓	⁶^,²⁷^,³⁵^,³⁶
Stromal cell-derived factor 1, 2	cytokine, growth factor, chemotaxis	Classical							²⁷^,²⁸^,³⁵^,³⁶
Transforming growth factor β (1, 2, 3)	Multifunctional proteins	Classical		1, 2, 3↑				2↓	26, 27, 28^,³⁵^,³⁶
Tumor necrosis factor α (TNF-α)	Cytokine								⁶^,²⁸
Vascular endothelial growth factor A, C, D	growth factor, angiogenesis, mitogen, differentiation	Classical						A, D↓	³⁵^,³⁶
*Enzymes*	Specificity	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
6-phosphogluconate dehydrogenase, decarboxylating	Oxydoreductase				↑				⁶^,²⁶^,²⁹^,³²^,34, 35, 36^,³⁸^,³⁹
6-phosphogluconolactonase	Hydrolase	Classical			↑				⁶^,²⁹^,³¹^,³²^,34, 35, 36^,³⁸
A disintegrin and metalloproteinase with thrombospondin motifs (ADAMTS) 1, 2, 4, 5, 6, 7, 9, 10, 12, 19	cell migration, inflammation, development	Classical		1↑				1, 2, 5, 7, 12↓	²⁷^,²⁸^,³²^,³⁵^,³⁶
Adenosylhomocysteinase	Hydrolase							↓	⁶^,²⁹^,33, 34, 35, 36^,³⁹
Alcohol dehydrogenase [NADP( + )]	Oxidoreductase				↑	↑			⁶^,²⁵^,²⁶^,²⁹^,34, 35, 36
Aldo-keto reductase family 1 member A4, C1, C2, C3	Oxidoreductase	Non-classical		A4 ↓					⁶^,²⁹^,³²^,³⁵^,³⁶
Aldose reductase	Oxidoreductase				↑				⁶^,¹¹^,³⁹^,²⁶^,²⁸^,²⁹^,33, 34, 35, 36^,³⁸
Aminoacylase-1	Hydrolase								⁶^,26, 27, 28, 29^,³⁶
Aminopeptidase-B,-N	Aminopeptidase, hydrolase, metalloprotease, protease	Classical			B↑				⁶^,²⁶^,²⁸^,²⁹^,33, 34, 35, 36
Angiogenin	angiogenesis, differentiation, stress response	Classical							²⁸^,³⁶
Arylsulfatase A, B, K	Hydrolase	Classical						A↓	²⁸^,²⁹^,³⁶
ATP synthase subunit beta, mitochondrial	Production of ATP from ADP				↑		↑		³¹
Carboxypeptidase A4, D, E, Q	Carboxypeptidase, Hydrolase, Metalloprotease, Protease	Classical					E↑	E, Q↓	²⁶^,²⁸^,³³^,³⁵^,³⁶^,³⁸
Cathepsin A, B, D, H, L1, Z, O	Hydrolase, Protease, Thiol protease	Classical		D, L1, O, Z, ↑	D↑ B, Z, L1↓			L1, O ↓	⁶^,²⁶^,³⁸^,³⁹^,27, 28, 29^,³¹^,³²^,34, 35, 36
Chymotrypsinogen B	Hydrolase, Protease, Serine protease	Classical		↑		↑			²⁵^,²⁷^,³⁵^,³⁶
Creatine kinase B-type, M-type	Kinase, Transferase			B↑	B↑				⁶^,¹¹^,26, 27, 28, 29^,34, 35, 36
Delta-aminolevulinic acid dehydratase	Allosteric enzyme, Lyase	Non-classical			↑	↓			⁶^,²⁵^,²⁹^,³¹^,³²^,34, 35, 36
Dipeptidyl peptidase 1, 2, 3, 4, 8, 9	Aminopeptidase, Hydrolase, Protease	Classical		2↑					⁶^,26, 27, 28, 29^,³⁵^,³⁶^,³⁸^,³⁹
Disintegrin and metalloproteinase domain-containing protein (ADAM) 9, 10, 12, 15 17, 19	Role in tumorigenesis and angiogenesis	Classical						10, 12, 19 ↓	²⁸^,³⁵^,³⁶^,³⁹
Extracellular superoxide dismutase [Cu–Zn]	Antioxidant, Oxidoreductase	Classical		↑				↓	⁶^,²⁶^,²⁷^,35, 36, 37
Farnesyl pyrophosphate synthetase	Transferase				↑				⁶^,²⁹^,32, 33, 34, 35, 36
Ferritin heavy chain, light chain	Stores iron in a soluble and non-toxic form	Non-classical			LC↓				⁶^,¹¹^,³⁶^,²⁶^,²⁸^,²⁹^,31, 32, 33, 34, 35
Fibroblast growth factor receptor 1, 4	Kinase, receptor, transferase	Classical		4↑					²⁸^,³⁶
Fructose-biphosphate aldolase A, B, C	Lyase involved in glycolysis			A↑	A↑			A↑	⁶^,¹¹^,²⁷^,²⁹^,32, 33, 34^,37, 38, 39
Glutathione S-transferase A4, MU1, MU2, ω-1, P1, alpha 3	Oxidoreductase, transferase	Classical			P1, ω-1↑				⁶^,¹¹^,³⁶^,³⁸^,³⁹^,²⁶^,²⁸^,²⁹^,31, 32, 33, 34, 35
Glutathione synthetase	Ligase	Non-classical			↓				⁶^,²⁶^,²⁹^,³⁴^,³⁶
Glyceraldehyde-3-phosphate dehydrogenase	inhibit cell spreading, glycolysis	Non-classical		↓	↑				⁶^,¹¹^,30, 31, 32^,34, 35, 36^,⁴⁰
Glycerol 3 phosphate dehydrogenase									¹¹^,³⁶
Glycogen phosphorylase, liver form, muscle form	Allosteric enzyme, Glycosyltransferase, Transferase				Muscle↑				⁶^,¹¹^,²⁶^,²⁹^,³⁴^,³⁶
Histidine triad nucleotide-binding protein 1	Hydrolase, role in apoptosis and transcription			↓					²⁸^,³⁵^,³⁶^,³⁸
Hypoxanthine-guanine phosphoribosyltransferase	Glycosyltransferase, transferase			↓	↑				²⁹^,³¹^,³²^,³⁵^,³⁶
Inosine triphosphate pyrophosphatase	hydrolase, magnesium binding	Non-classical							²⁶^,³¹^,³⁵^,³⁶
Inositol monophosphatase 1, 2	Hydrolase	Classical			↑				⁶^,²⁶^,²⁹^,34, 35, 36
Inositol-3-phosphate synthase 1	Involved in myo-inositol biosynthesis				↑				²⁶^,³⁴^,³⁶
Lipoprotein lipase	Lipid metabolism, hydrolase	Classical		↓					²⁷^,³⁵^,³⁶
Mannan-binding lectin serine protease 1	Role in immunity	Classical				↑			²⁵^,²⁶^,³⁵^,³⁶
Matrix metalloproteinase 2, 9, 14, 19	Regulation of cell migration, metalloproteinase, breakdown of ECM	Classical					2↑	2, 9, 19↓	²⁷^,²⁸^,³¹^,³⁵^,³⁶^,³⁹^,⁴⁰
NEDD8-conjugating enzyme Ubc 12	Transferase, involved in cell proliferation	Non-classical							⁶^,³¹^,³⁵^,³⁶
Obg-like ATPase 1	ATP hydrolysis				↑	↓			25, 26, 27, 28^,³⁴^,³⁶
Peptidyl-prolyl cis–trans isomerase A, B, C, D, H, like 1, CWC27 homolog, FKBP (1, 3, 4, 9)	accelerate the folding of proteins	Classical			A, B, C↑			FKBP9 ↓	⁶^,¹¹^,36, 37, 38, 39^,26, 27, 28^,31, 32, 33, 34, 35
Peroxidasin homolog	Peroxidase activity and role in ECM formation	Classical				↓			⁶^,25, 26, 27^,³⁵^,³⁶
Peroxiredoxin 1, 2, 4, 5, 6	Protecting cells from free-radical damage	Classical		1, 2 ↓	1, 5, 6↑			4 ↑ and ↓ 6↑	⁶^,¹¹^,³⁸^,³⁹^,²⁷^,²⁹^,31, 32, 33, 34, 35, 36
Phosphoglycerate mutase 1, 2	Hydrolase, isomerase		2↑		2↑				⁶^,¹¹^,³⁹^,28, 29, 30^,³²^,34, 35, 36^,³⁸
Phospholipase A1 member A, A2 activating protein, D3	Hydrolase, Protease, Serine protease	Classical						A1A↓	²⁶^,²⁷^,³³^,³⁵^,³⁶
Procollagen C-endopeptidase enhancer 1	collagenase enhancer	Classical						↓	⁶^,²⁷^,³¹^,³²^,³⁵^,38, 39, 40
Prolyl endopeptidase	Role in tissue remodeling, fibrosis, inflammation								²⁶^,²⁹^,³³^,³⁵^,³⁶
Proteasome subunit β (type 1, 2, 3, 4, 5, 6, 8), α (1, 2, 3, 4, 5, 6, 7)	Hydrolase, protease, threonine protease	Non-classical	β type-6 ↑	β type (4, 5) ↓	β6, β8↑				⁶^,¹¹^,²⁷^,³¹^,³²^,34, 35, 36^,³⁸
Protein disulfide isomerase		Classical			↑			↓	¹¹^,³⁵^,³⁶^,³⁸^,³⁹
Protein disulfide-isomerase A (3, 4, 5, 6)	chaperone, isomerase,	Classical			A3, A6↑	A3↑	A3↑	A6↓, A3, A4↑	⁶^,¹¹^,37, 38, 39^,²⁵^,²⁷^,²⁸^,³¹^,³²^,34, 35, 36
Protein/nucleic acid deglycase DJ-1	Chaperone, hydrolase, protease, RNA binding	Non-classical			↑			↓	⁶^,¹¹^,³¹^,³²^,34, 35, 36^,³⁸^,³⁹
Puromycin-sensitive aminopeptidase	involved in cell growth and viability				↑				⁶^,³⁴^,³⁶^,³⁸^,³⁹
Pyridoxal phosphate phosphatase	involved in mitosis and cytokinesis								²⁷^,³⁵^,³⁶
Pyruvate kinase (isozymes R/L, M1/M2, PKM)	Plays a key role in glycolysis, marker of myonecrotic conditions	PKM↑			M1/M2↑				¹¹^,²⁸^,²⁹^,33, 34, 35, 36^,³⁸^,³⁹
Secretory leucocyte protease inhibitor									³⁶
Serine protease HTRA1, 2, 23	growth factor binding, hydrolase, protease, serine protease	Classical						2↑ 1↓	⁶^,26, 27, 28^,³⁵^,³⁶
Spermidine synthase	Transferase	Non-classical			↑				³¹^,³²^,34, 35, 36
Sulfhydryl oxidase 1, 2		Classical							⁶^,26, 27, 28^,³⁵^,³⁶
Superoxide dismutase [Cu–Zn]	Antioxidant, Oxidoreductase	Non-classical	↑	↓	↑				⁶^,¹¹^,²⁷^,²⁸^,³¹^,³²^,³⁴^,³⁵^,³⁸^,³⁹
Tissue-type plasminogen activator	Tissue remodeling and degradation	Classical		↑					²⁶^,²⁷^,³⁵^,³⁶
Transketolase	Transferase, calcium and magnesium binding								⁶^,²⁸^,²⁹^,34, 35, 36^,³⁸^,³⁹
Triosephosphate isomerase	Isomerase involved in gluconeogenesis and glycolysis				↑				⁶^,¹¹^,³⁶^,³⁸^,³⁹^,²⁶^,28, 29, 30^,32, 33, 34, 35
Ubiquitin carboxyl-terminal hydrolase 17-like protein D	Protease, hydrolase, thiol protease					↓			²⁵
Ubiquitin-conjugating enzyme E1-like 2 (isoform CRA_a), E2 (D1, D2, L3, K, N, Z, variant 1 et 2)	Transferase	Non-classical		E1-like 2 ↓	E2 variant 1, K, N↑				²⁶^,²⁷^,³¹^,³²^,34, 35, 36
α, β, γ enolase	lyase		α, γ ↑		α, β, γ↑		α↑	α ↑	⁶^,¹¹^,³⁸^,³⁹^,²⁶^,²⁸^,²⁹^,32, 33, 34, 35, 36
α, β-mannosidase		Classical						β↓	⁶^,²⁶^,³⁶
*Enzymatic inhibitors*	Specificity	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
Amyloid-β A4 protein	Apoptosis, cell adhesion, endocytosis, Notch signaling pathways	Classical						↓	²⁶
Cystatin B, C	cysteine protease inhibitor	Classical		C↑				C↓	⁶^,²⁶^,³⁹^,⁴⁰^,²⁷^,²⁸^,³¹^,³²^,35, 36, 37, 38
Metalloproteinase inhibitor (TIMP) 1, 2	Cytokine, growth factor, metalloendopeptidase inhibitor	Classical		1, 2↑	1↓	1↓	2↓	2↓	⁶^,²⁵^,35, 36, 37, 38, 39^,26, 27, 28, 29, 30, 31, 32^,³⁴
Phosphatidylethanolamine-binding protein 1	Protease inhibitor, serine protease inhibitor	Non-classical	↑	↑ and ↓	↑			↑	⁶^,¹¹^,²⁸^,²⁹^,35, 36, 37, 38, 39
Serpin A6, A8 (angiotensinogen), A11, A3N	Regulator of blood pressure, body fluid and electrolytes homeostasis	Classical				A3N↑		A6↓	²⁵^,²⁸^,³⁶
Serpin B1 (Leukocyte elastase inhibitor), B2 (Plasminogen activator inhibitor 2), B6, B9	Inhibitor of serine protease, protease inhibitor	Non-classical		B2↓	B1, B6↑				⁶^,27, 28, 29^,32, 33, 34, 35, 36^,³⁹
Serpin C1 (Antithrombin-III)	Inhibitor of serine protease, regulation of cell migration	Classical	↑	↓					⁶^,²⁷^,²⁹^,³¹^,³²^,³⁵
Serpin E1 (Plasminogen activator inhibitor 1), E2	Inhibitor of serine protease, regulation of cell migration	Classical		E1, E2↑			E1↑	E1↑ E2↓	⁶^,26, 27, 28, 29^,³⁵^,³⁶^,⁴⁰
Serpin F1 (pigment epithelium derived factor (PEDF))	Inhibitor of serine protease, regulation of cell migration	Classical						↓	⁶^,26, 27, 28, 29^,35, 36, 37^,⁴⁰
Serpin G1 (Plasma protease C1 inhibitor)	Inhibitor of serine protease, regulation of cell migration				↓				⁶^,²⁶^,²⁷^,³⁰^,³⁴
Serpin H1	Inhibitor of serine protease, regulation of cell migration	Classical					↑	↑ and ↓	⁶^,²⁸^,³¹^,³²^,³⁵^,³⁶^,³⁸^,³⁹
Tissue factor pathway inhibitor	Anti-thrombic action and associates lipoproteins in plasma	Classical		↑				↓	²⁷^,³⁵^,³⁶
α(1,2)-macroglobulin	Protease inhibitor, serine protease inhibitor	Classical	1, 2↑	↓					⁶^,27, 28, 29^,³⁴^,³⁵
*Cytoskeletal proteins*	Specificity	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
Actin, aortic smooth muscle									⁶^,¹¹^,²⁸^,²⁹^,³⁵^,³⁶
Actin, cytoplasmic 1, 2									⁶^,²⁸^,²⁹^,³²^,³⁶^,³⁹
Actin, α cardiac muscle 1			↑						⁶^,¹¹^,²⁸^,²⁹
Actin, α skeletal muscle			↑		↑				⁶^,¹¹^,²⁸^,²⁹^,34, 35, 36^,³⁹
Actin-related protein 2, 3					2↑				⁶^,²⁸^,²⁹^,34, 35, 36
Calponin 2, 3	regulation and modulation of muscle contraction	non-classical			3↑				²⁸^,³¹^,³²^,34, 35, 36
Cell division control protein 42 homolog	Differentiation, neurogenesis	non-classical							³¹^,³²^,³⁵^,³⁶
Cofilin 1, 2	Regulates actin cytoskeleton dynamics	non-classical		1 ↓	2↑			1 ↑	⁶^,¹¹^,³⁸^,³⁹^,²⁶^,²⁸^,31, 32, 33, 34, 35, 36
Desmin	Muscle-specific type III intermediate filament	non-classical	↑		↑				⁶^,¹¹^,³⁶^,²⁶^,28, 29, 30, 31, 32^,³⁴^,³⁵
Destrin	Actin-depolymerizing protein	non-classical							⁶^,³¹^,³²^,³⁵^,³⁶
Ezrin	Cell shape								⁶^,²⁸^,²⁹^,³³^,³⁵^,³⁶
F-actin-capping protein subunit α (1, 2), β	actin capping, actin binding								⁶^,²⁹^,³⁵^,³⁶
Filamin A, B, C	Actin binding				C↑				⁶^,¹¹^,²⁸^,²⁹^,33, 34, 35, 36^,³⁸^,³⁹
Gelsolin	actin capping, actin binding	Classical			↑				⁶^,²⁴^,³⁸^,26, 27, 28, 29^,33, 34, 35, 36
Myomesin 1, 3	Major component of the vertebrate myofibrillar M band								¹¹^,³⁶
Myosin 1, 3, 4, 6, 7, 8, 9, 10	Role in cytokinesis, cell shape, secretion and capping				3↑				⁶^,¹¹^,³⁹^,28, 29, 30^,33, 34, 35, 36^,³⁸
Myosin binding protein C, fast type	cell adhesion								¹¹^,³⁰
Myosin light chain 1, 2, 3, 4, 6B, 12B, 1/3	Muscle contraction, motor activity, calcium binding				1, 4, 6B, 12B↑				⁶^,¹³^,²⁸^,²⁹^,³¹^,35, 36, 37^,⁴⁰^,²⁹
Myosin regulatory light chain 2, skeletal muscle isoform, 4, 12B	Contractile protein				2↑				⁶^,¹¹^,²⁸^,34, 35, 36^,³⁸^,³⁹
Myotilin	Involved in a complex of actin cross-linking proteins								¹¹
Palladin	Role in cell morphology, motility, cell adhesion								²⁸^,²⁹^,³⁵^,³⁶
Peripherin	Neuronal intermediate filament protein	Non-classical							⁶^,³¹^,³²
Plectin	actin binding		↑						⁶^,¹¹^,²⁸^,³³^,³⁵^,³⁶
Profilin 1, 2	Actin binding	Non-classical		1 ↓	1↑		1↑		⁶^,¹¹^,²⁸^,³¹^,³²^,³⁴^,³⁶^,³⁸^,³⁹
Titin	Key component in the functioning of striated muscle				↑				¹¹^,²⁹^,³⁰^,33, 34, 35, 36
Transgelin 1, 2		Non-classical		2 ↓	1, 2↑				⁶^,²⁸^,31, 32, 33, 34, 35^,³⁸^,³⁹
Tropomyosin α (1, 3, 4), β (1, 2)	actin-binding	Non-classical	α1, β ↑	β ↓	β↑			β 2 ↑	⁶^,¹¹^,30, 31, 32^,³⁴^,³⁵^,³⁸^,³⁹
Troponin I (slow skeletal muscle), C (slow, skeletal and cardiac muscle), T	inhibitory subunit of troponin, muscle protein, actin binding				I, C, T↑				⁶^,¹¹^,34, 35, 36^,³⁹
Tubulin α (A,4A, B), β (2A, 3, 4B, 5, 6)		Non-classical	α4A↑		α1A↑				⁶^,²⁶^,32, 33, 34, 35, 36^,³⁹
Vimentin	cytoskeletal protein	Non-classical	↑		↑			↑	⁶^,¹¹^,³⁶^,38, 39, 40^,28, 29, 30, 31, 32, 33, 34, 35
Vinculin	Actin binding, cell adhesion		↑		↑				⁶^,28, 29, 30^,34, 35, 36^,³⁸^,³⁹
α-actinin 1, 2, 3, 4	actin binding				3, 4↑				⁶^,²⁴^,²⁸^,²⁹^,34, 35, 36^,³⁸^,³⁹
*Miscellaneous*	Specificity	Secretion	Exercise	Myogenesis	Dystrophin deficiency	Atrophy	Insulin stimulation	Insulin resistant cells	References
14-3-3 protein β/α, γ, ζ/δ, ε, η, θ		Non-classical			ε↑				⁶^,²⁸^,²⁹^,32, 33, 34, 35, 36^,³⁸^,³⁹
ADAMTS-like protein 1, 2, 3, 4	Hydrolase	Classical		3↑				2↓	²⁷^,²⁸^,³⁵^,³⁶
Adapter molecule crk	cell adhesion, spreading, migration								⁶^,²⁶^,²⁸^,³⁵^,³⁶
Adiponectin	hormone involved in fat metabolism and insulin sensitivity								²⁶
Agrin	Developmental protein	Non-classical						↓	²⁷^,²⁸^,³⁵^,³⁶
Alpha-1-acid glycoprotein 1	Transport	Classical				↑			²⁵
Angiopoietin-related protein 1, 2, 4	Induces sprouting in endothelial cells	Classical		2↑				2↓ 4↑	²⁶^,²⁸^,³¹^,³²^,³⁵^,³⁶
Annexin A (1, 2, 4, 5, 6)	involved in exocytosis, regulation of cell migration and fusion	1, 5, Non-classical 2 Classical		1, 2 ↓				A1, A2↓	⁶^,²⁶^,38, 39, 40^,²⁷^,²⁸^,31, 32, 33, 34, 35, 36
AP-1 complex subunit β1, γ1, σ (1, 2), μ1	Transport, protein transport	Non-classical							²⁸^,³¹^,³⁵^,³⁶
Apolipoprotein B-100, D, E	Lipid metabolism and transport	Classical	B-100↑	E↑					⁶^,²⁷^,²⁸^,³⁵^,³⁶
Basal cell adhesion molecule	Blood group antigen, receptor								⁶^,²⁶^,²⁹
Brain acid soluble protein					↑				²⁸^,³³^,³⁸^,³⁹
Cadherin 2, 5, 11, 13, 15 (M-cadherin), 23	cell adhesion	Classical	23 ↑	11, 13, 15↑	2, 15↑		15↑	2, 11, 13, 15↓	⁶^,²⁶^,³⁹^,27, 28, 29^,33, 34, 35, 36^,³⁸
Calcium/calmodulin-dependent protein kinase type 1	Cell cycle, differentiation, neurogenesis								²⁷^,³⁵^,³⁶
Calmodulin	mediates a large number of enzymes				↑				⁶^,¹¹^,²⁸^,³³^,³⁴^,³⁶^,³⁹
Calpastatin	inhibition of calpain, may be involved in protein degradation in muscle								²⁸^,³⁵^,³⁶^,³⁹
Calreticulin	calcium binding chaperone	Classical		↓					⁶^,¹¹^,²⁷^,²⁸^,34, 35, 36, 37, 38
Calsequestrin 1, 2	calcium storage	Classical		2↑					⁶^,²⁶^,²⁷^,³³^,³⁶^,³⁸
Calsyntenin 1, 2		Classical						1↓	⁶^,26, 27, 28^,³⁵^,³⁶^,³⁸
Calumenin	Metal-binding, calcium binding	Classical							⁶^,27, 28, 29^,34, 35, 36^,³⁸^,³⁹
Cartilage intermediate layer protein 1, 2		Classical		1, 2↑				↓	²⁶^,²⁷^,³⁶
Catenin α1	Association with cadherin					↑			²⁵^,²⁸^,³⁵^,³⁶
Cellular nucleic acid-binding protein	Single-stranded DNA-binding protein						↓		³¹
Chloride intracellular channel protein 1, 4	transport, chloride transport	Non-classical		4 <->	4↑				⁶^,²⁶^,31, 32, 33, 34, 35, 36
Chordin	Developmental protein	Classical						↓	²⁷^,³⁵^,³⁶
Clusterin	apoptosis, immunity	Classical					↑	↑ and ↓	³⁸^,³⁹
Coatomer subunit α, β, δ, ε	Transport	Classical							⁶^,²⁸^,³⁰^,³²^,34, 35, 36
Coiled-coil domain-containing protein 80, 126	Promotes cell adhesion and matrix assembly	Classical		80↑					⁶^,²⁷^,²⁸^,³⁶^,³⁸^,³⁹
Complement component 1, C3, C4 (A, B)	Immune response		C3,C4B↑						⁶^,²⁸^,³⁰^,³⁵^,³⁹
Complement factor B, D, H	Immune response	Classical		H ↑		↑		H↓	⁶^,25, 26, 27, 28^,³⁰^,³⁵^,³⁶
Cytochrome C									⁶^,¹¹^,²⁷^,²⁸^,³⁵^,³⁶
Drebrin	Actin binding, differentiation, neurogenesis								⁶^,²⁸^,³⁵^,³⁶^,³⁹
Elongation factor 1 (α1, β, δ, γ) 2	Factor elongation		2↑		1α1, 1γ, 1,β, 2↑				⁶^,¹¹^,³⁶^,³⁸^,³⁹^,26, 27, 28, 29^,32, 33, 34, 35
Endoplasmin	Chaperone	Classical			↑				⁶^,²⁷^,²⁸^,³¹^,34, 35, 36^,³⁸
Ephrin type-A receptor 1, 2, 4, 7	cell adhesion, neurogenesis	Classical						2↓	⁶^,²⁷^,²⁸^,³³^,³⁶
Exostosin 1, 2	protein glycosylation								26, 27, 28^,³⁵^,³⁶
Fascin	actin binding				↑				⁶^,³²^,34, 35, 36^,³⁸
Fatty acid-binding protein, heart	Transport of lipids		↑						¹¹^,²⁸^,³⁵^,³⁶
Follistatin	activin antagonist, inhibitor of FSH biosynthesis	Classical		↑		↑		↓	²⁵^,²⁶^,³⁵^,³⁶
Four and a half LIM domains protein 1, 2, 3	Metal binding	Non-classical							⁶^,¹¹^,³¹^,³²^,³⁵^,³⁶^,³⁸
Galectin-3-binding protein	Cell adhesion	Classical				↓			25, 26, 27^,³⁵^,³⁶
Glucosidase 2 subunit β	Metal binding, calcium binding	Classical			↑				⁶^,²⁸^,²⁹^,³⁴^,³⁶
Growth factor receptor-bound protein 2	link between growth factor and RAS signaling pathway								⁶^,²⁶^,³⁵^,³⁶
Heat shock 70 kDa protein 1A/1B, 1 like,2, 4	ATP binding, nucleotide binding		2↑						⁶^,¹¹^,²⁶^,²⁹^,34, 35, 36^,³⁸^,³⁹
Heat shock protein beta(1, 2, 3, 6, 7)	Chaperone				2↑			1 ↑ and ↓	⁶^,¹¹^,²⁸^,²⁹^,33, 34, 35, 36^,³⁸^,³⁹
Heat shock protein HSP 90 (α, β)	Chaperone			α↑	α↑				⁶^,36, 37, 38
Heme-binding protein 1, 2									⁶^,²⁸^,²⁹^,³⁵^,³⁶
High mobility group protein HMGI-C	Cell cycle, cell division, growth regulation								²⁸^,³⁵^,³⁶
Hypoxia up-regulated protein 1	role in cytoprotection triggered by oxygen deprivation	Classical							²⁷^,²⁸^,³⁶
Importin 5, 7, 9	transport nucléaire								⁶^,³³^,³⁵^,³⁶
Integrin α (1, 3, 5, 6, 7), β (1, 5)	cell adhesion, integrin, receptor, cell adhesion	Classical						α3, 5, 7↓ β1, 5 ↓	²⁷^,²⁸^,³⁰^,³³^,³⁵^,³⁶
Lactadherin	angiogenesis, cell adhesion	Classical							²⁷^,²⁸^,³⁵^,³⁶^,³⁸
Leukemia inhibitory factor receptor	Signal transducing molecule	Classical						↓	²⁷^,³⁶
l-lactate dehydrogenase A chain, B chain	Pyruvate fermentation to lactate	Non-classical		A ↓	A↑		A↑		⁶^,¹¹^,³⁸^,³⁹^,²⁴^,²⁶^,²⁸^,³¹^,³²^,34, 35, 36
Lipopolysaccharide-binding protein	Innate immune response	Classical				↑			²⁵^,³⁶
Low-density lipoprotein receptor	Binds and transport LDL	Classical							²⁶^,³⁶^,³⁸
Lysosome-associated membrane protein 1, 2					1↑ 2↓				²⁶^,³⁶
Lysyl oxidase homolog 1, 2, 3, 4	Active on elastin and collagen substrates	Classical		1↑		↑		1, 3, 4↓	25, 26, 27, 28^,³⁵^,³⁶
Macrophage-capping protein	actin capping and binding	Classical							⁶^,²⁷^,²⁸^,³³^,³⁴^,³⁶
Meteorin-like protein	Hormone induced by exercising and promotes energy expenditure	Classical							²⁷^,³⁵^,³⁶
Microtubule-associated proteins 1A/1B light chain 3B, 1S, 4	Mitophagy	Non-classical							²⁶^,³¹^,³²^,³⁵^,³⁶^,³⁹
Musculoskeletal embryonic nuclear protein 1	May be involved in the development of the musculoskeletal system						↓		³¹
Myc box-dependent-interacting protein 1	Differentiation, endocytosis				↑				⁶^,²⁷^,²⁸^,33, 34, 35, 36^,³⁸
Myoferlin	involved in muscle contraction				↑				33, 34, 35, 36
Myoglobin	oxygen transport	Non-classical							¹¹^,²⁶^,³⁰^,34, 35, 36
Myotrophin	regulation of the growth of actin filaments								⁶^,²⁸^,³⁵^,³⁶^,³⁸
NAC α domain containing	Protein transport	Classical		↑					³²^,⁴⁰
Nascent polypeptide-associated complex subunit alpha, muscle-specific form	Muscle -specific transcription factor				↑				34, 35, 36
Neogenin	cell adhesion in myogenesis	Classical						↓	²⁷^,²⁸^,³⁶
Nephronectin	Cell adhesion, spreading and survival	Classical						↓	²⁷^,³³^,³⁵^,³⁶
Neudesin	Promotes cell proliferation and neurogenesis	Classical							²⁷^,³⁵^,³⁶
Neural cell adhesion molecule 1	Cell adhesion	Classical			↑			↓	⁶^,²⁶^,³⁹^,27, 28, 29^,33, 34, 35, 36^,³⁸
Neuropilin 1, 2	Angiogenesis, differentiation, neurogenesis	Classical						1, 2↓	26, 27, 28^,³³^,³⁶
Nuclease-sensitive element-binding protein 1	mitogen, activator, repressor	Classical							²⁸^,35, 36, 37
Nucleobindin 1, 2	Calcium binding, may have a role in calcium homeostasis	Classical			1↑		1, 2↓	1 ↓, 2 ↑	²⁴^,26, 27, 28^,34, 35, 36^,³⁸^,³⁹
Nucleophosmin	cell proliferation, regulation of tumor suppresors		↑						¹¹^,³²^,³⁵^,³⁶
Olfactomedin-like protein 2A, 2B, 3	extracellular matrix organization	Classical		2A↑				2B, 3↓	26, 27, 28^,³⁵^,³⁶
Parvalbumin alpha	involved in relaxation after contraction								¹¹^,²⁷^,³⁵
Phospholipid transfer protein	Lipid transport	Classical						↓	⁶^,26, 27, 28^,³³^,³⁶
Plastin 3	Actin binding								²⁶^,²⁸^,³³^,³⁵^,³⁶^,³⁹
Plexin A (1, 2, 3, 4), B2		Classical						B2↓	²⁷^,³³^,³⁵^,³⁶
Poly(rC)-binding protein 1, 2, 3	DNA binding, RNA binding	Non-classical							⁶^,³¹^,³²^,³⁵^,³⁶
Programmed cell death 5, 6-interacting protein	apoptosis, cell cyle, cell division, transport						5↓		26, 27, 28^,³³^,³⁵^,³⁶^,³⁹
Proliferating cell nuclear antigen	DNA binding	Non-classical							²⁶^,²⁸^,³¹^,³²^,³⁵^,³⁶
Prostaglandin F2 receptor negative regulator	Regulation of myoblast fusion					↓			²⁵^,²⁷^,²⁸^,³³^,³⁶^,³⁸
Protein jagged-1	Inhibits myoblast differenciation	Classical							²⁸^,³⁶
Protein NOV homolog	Role in proliferation, adhesion, migration, differentiation, survival	Classical		↑					²⁶^,³⁶
Protein RCC2	mitosis and cytokinesis								³⁵^,³⁶
Protein VAC14 homolog	phosphatidylinositol biosynthetic process and signal transduction						↑		³¹
Protocadherin-1, gamma C3, 18, 19, beta 14, Fat 4	cell to cell adhesion and interaction processes	Classical						19, Fat 4↓	²⁷^,³⁵^,³⁶
Ras suppressor protein 1	regulation of cell substrate adhesion	Non-classical			↑				34, 35, 36
Ras-related protein Rab 5A, 14, 35	Protein transport, transport	Non-classical							²⁹^,³³^,³⁵^,³⁶
Reticulocalbin 1, 2, 3	Calcium, metal binding	Classical						↓	⁶^,²⁶^,²⁸^,34, 35, 36^,³⁹
Reticulon 2, 4	Neurogenesis				4↑				33, 34, 35, 36
Rho GDP dissociation inhibitor (GDI) α	Controls rho proteins homeostasis	Non-classical		↓	↑				²⁸^,³⁵
Semaphorin 3(A,B, C, D, E), 4(B, C), 5A, 6(A, B), 7A	Role in differenciation, inflammation and neurogenesis	Classical		3A, 3D, 3E, 6A (isoform 1)↑ 7A↑ and↓				3 (A, B), 4B, 4C, 6A↓	26, 27, 28, 29^,35, 36, 37
Septin 2, 7, 9, 11	Cell cycle, cell division, differentiation, mitosis				11↑				²⁶^,²⁸^,34, 35, 36^,³⁸
Serotransferrin	Iron transport		↑						¹¹
SH3 domain-binding glutamic acid-rich protein									⁶^,³⁰^,³¹^,³⁵^,³⁶
Slit homolog 2 protein	Differentiation, neurogenesis	Classical						↓	²⁷^,³⁶
Spondin 2	cell adhesion, immunity	Classical						↓	⁶^,²⁶^,²⁷^,³¹^,³²^,³⁵
Stanniocalcin 1, 2	Stimulates renal phosphate reabsorption	Classical							⁶^,²⁸^,³¹^,³²^,³⁵^,³⁶^,³⁸
Stathmin 1, 2	Regulation of the microtubule filament							1 ↑	⁶^,30, 31, 32^,³⁵^,³⁶^,³⁹
Sterile α motif domain-containing protein 3				↓					³²
Sushi repeat-containing protein SRPX2	angiogenesis, cell adhesion							↓	⁶^,26, 27, 28^,³⁶
Syntenin-1	Tumorigenesis, exosome biosynthesis								⁶^,²⁷^,²⁸^,³²^,³⁵^,³⁶
Talin 1, 2	connections in the cytoskelettal structure								²⁸^,³³^,³⁵^,³⁶^,³⁸
Telethonin	Muscle assembly regulating factor								¹¹
Testican 1, 2	Neurogenesis	Classical				2↑			⁶^,25, 26, 27, 28^,³⁶
Tetraspanin 4, 6, 7, 9, 14	may be involved in cell proliferation and motility	Non-classical							³³^,³⁵^,³⁶
Thrombospondin 1, 2	mediates cell to cell and cell to matrix interactions	Classical		↑				1↑	⁶^,26, 27, 28^,³¹^,³²^,35, 36, 37
Transcobalamin 2	Vitamin B12 binding, transport protein	Classical						↓	²⁶^,²⁷^,³⁵^,³⁶^,³⁹
Transcription elongation factor B polypeptide 2 (Elongin-B)	Transcription factor					↑			25, 26, 27, 28^,³⁵^,³⁶
Translationally-controlled tumor protein	involved in microtubule stabilization and calcium binding	Non-classical		↑	↑				⁶^,²⁷^,²⁸^,³¹^,³²^,34, 35, 36, 37, 38
Ubiquilin 1, 2, 3	Regulation of protein degradation								²⁸^,³⁵^,³⁶
Vacuolar protein sorting-associated protein 26A, 29, 35, 37C VTA1 homolog	protein transporter			↑	26A, 35↑				⁶^,26, 27, 28^,33, 34, 35, 36
Vascular cell adhesion protein 1	Cell–cell recognition	Classical		Isoform 1↑					²⁷^,²⁸^,³⁶
Vasorin		Classical							⁶^,²⁷^,²⁸^,³⁶
Vigilin	Lipid metabolism	Non-classical						↑	²⁸^,³⁵^,³⁶
Vitamin K-dependent protein S	Blood coagulation, fibrinolysis, hemostasis	Classical						↓	⁶^,²⁶^,²⁷^,³⁶
WNT1-inducible-signaling pathway protein 1	Associated with cell survival	Classical							²⁷^,³⁵^,³⁶
α-crystallin A chain (HSP4), B chain	chaperone, eye-lens protection				B↑				⁶^,¹¹^,²⁶^,³⁰^,33, 34, 35, 36^,³⁹
β-2-microglobulin	component of the class I MHC	Classical	↑					↓	²⁶^,³⁸^,³⁹