Myostatin gene promoter: structure,conservation and importance as a target for muscle modulation
|
摘要
Myostatin(MSTN) is one of the key factors regulating myogenesis. Because of its role as a negative regulator of muscle mass deposition, much interest has been given to its protein and, in recent years, several studies have analysed MSTN gene regulation. This review discusses the MSTN gene promoter, focusing on its structure in several animal species, both vertebrate and invertebrate. We report the important binding sites considering their degree of phylogenetic conservation and roles they play in the promoter activity. Finally, we discuss recent studies focusing on MSTN gene regulation via promoter manipulation and the potential applications they have both in medicine and agriculture.
Myostatin(MSTN) is one of the key factors regulating myogenesis. Because of its role as a negative regulator of muscle mass deposition, much interest has been given to its protein and, in recent years, several studies have analysed MSTN gene regulation. This review discusses the MSTN gene promoter, focusing on its structure in several animal species, both vertebrate and invertebrate. We report the important binding sites considering their degree of phylogenetic conservation and roles they play in the promoter activity. Finally, we discuss recent studies focusing on MSTN gene regulation via promoter manipulation and the potential applications they have both in medicine and agriculture.
Myostatin(MSTN) is one of the key factors regulating myogenesis. Because of its role as a negative regulator of muscle mass deposition, much interest has been given to its protein and, in recent years, several studies have analysed MSTN gene regulation. This review discusses the MSTN gene promoter, focusing on its structure in several animal species, both vertebrate and invertebrate. We report the important binding sites considering their degree of phylogenetic conservation and roles they play in the promoter activity. Finally, we discuss recent studies focusing on MSTN gene regulation via promoter manipulation and the potential applications they have both in medicine and agriculture.
引文
1.McPherron AC,Lawler AM,Lee SJ.Regulation of skeletal muscle mass in mice by a new TGF-beta superfamily member.Nature.1997;387:83-90.
2.Joulia D,Bernardi H,Garandel V,Rabenoelina F,Vernus B,Cabello G.Mechanisms involved in the inhibition of myoblast proliferation and differentiation by myostatin.Exp Cell Res.2003;286:263-75.
3.McPherron AC,Lee SJ.Double muscling in cattle due to mutations in the myostatin gene.Proc Natl Acad Sci U S A.1997;94:12457-61.
4.Clop A,Marcq F,Takeda H,Pirottin D,Tordoir X,BibéB,et al.A mutation creating a potential illegitimate microRNA target site in the myostatin gene affects muscularity in sheep.Nat Genet.2006;38:813-8.
5.Mosher DS,Quignon P,Bustamante CD,Sutter NB,Mellersh CS,Parker HG,et al.A mutation in the myostatin gene increases muscle mass and enhances racing performance in heterozygote dogs.PLoS Genet.2007;3:e79.
6.Shelton GD,Engvall E.Gross muscle hypertrophy in whippet dogs is caused by a mutation in the myostatin gene.Neuromuscul Disord NMD.2007;17:721-2.
7.Schuelke M,Wagner KR,Stolz LE,Hübner C,Riebel T,K?men W,et al.Myostatin mutation associated with gross muscle hypertrophy in a child.NEngl J Med.2004;350:2682-8.
8.Jespersen J,Kjaer M,Schjerling P.The possible role of myostatin in skeletal muscle atrophy and cachexia.Scand J Med Sci Sports.2006;16:74-82.
9.Joulia-Ekaza D,Cabello G.The myostatin gene:physiology and pharmacological relevance.Curr Opin Pharmacol.2007;7:310-5.
10.Wójcik S,Engel WK,McFerrin J,Askanas V.Myostatin is increased and complexes with amyloid-beta within sporadic inclusion-body myositis muscle fibers.Acta Neuropathol(Berl).2005;110:173-7.
11.Bogdanovich S,Krag TOB,Barton ER,Morris LD,Whittemore L-A,Ahima RS,et al.Functional improvement of dystrophic muscle by myostatin blockade.Nature.2002;420:418-21.
12.Tang L,Yan Z,Wan Y,Han W,Zhang Y.Myostatin DNA vaccine increases skeletal muscle mass and endurance in mice.Muscle Nerve.2007;36:342-8.
13.Takata H,Terada K,Oka H,Sunada Y,Moriguchi T,Nohno T.Involvement of Wnt4 signaling during myogenic proliferation and differentiation of skeletal muscle.Dev Dyn Off Publ Am Assoc Anat.2007;236:2800-7.
14.Allen DL,Loh AS.Posttranscriptional mechanisms involving microRNA-27a and b contribute to fast-specific and glucocorticoid-mediated myostatin expression in skeletal muscle.Am J Physiol Cell Physiol.2011;300:C124-37.
15.Wilt F,Hake S.Principles of developmental biology.New York:W.W.Norton&Company,Inc;2004.
16.Heintzman ND,Ren B.The gateway to transcription:identifying,characterizing and understanding promoters in the eukaryotic genome.Cell Mol Life Sci.2007;64:386-400.
17.Ma K,Mallidis C,Artaza J,Taylor W,Gonzalez-Cadavid N,Bhasin S.Characterization of 5’-regulatory region of human myostatin gene:regulation by dexamethasone in vitro.Am J Physiol Endocrinol Metab.2001;281:E1128-36.
18.Zammit PS.Function of the myogenic regulatory factors Myf5,MyoD,Myogenin and MRF4 in skeletal muscle,satellite cells and regenerative myogenesis.Semin Cell Dev Biol.2017;72:19-32.
19.Spiller MP,Kambadur R,Jeanplong F,Thomas M,Martyn JK,Bass JJ,et al.The myostatin gene is a downstream target gene of basic helix-loop-helix transcription factor MyoD.Mol Cell Biol.2002;22:7066-82.
20.Fan H,Zhang R,Tesfaye D,Tholen E,Looft C,H?lker M,et al.Sulforaphane causes a major epigenetic repression of myostatin in porcine satellite cells.Epigenetics.2012;7:1379-90.
21.Taylor MV,Hughes SM.Mef2 and the skeletal muscle differentiation program.Semin Cell Dev Biol.2017;72:33-44.
22.Ríos R,Carneiro I,Arce VM,Devesa J.Myostatin regulates cell survival during C2C12 myogenesis.Biochem Biophys Res Commun.2001;280:561-6.
23.Artaza JN,Bhasin S,Mallidis C,Taylor W,Ma K,Gonzalez-Cadavid NF.Endogenous expression and localization of myostatin and its relation to myosin heavy chain distribution in C2C12 skeletal muscle cells.J Cell Physiol.2002;190:170-9.
24.Salerno MS,Thomas M,Forbes D,Watson T,Kambadur R,Sharma M.Molecular analysis of fiber type-specific expression of murine myostatin promoter.Am J Physiol Cell Physiol.2004;287:C1031-40.
25.Grade CVC,Salerno MS,Schubert FR,Dietrich S,Alvares LE.An evolutionarily conserved Myostatin proximal promoter/enhancer confers basal levels of transcription and spatial specificity in vivo.Dev Genes Evol.2009;219:497-508.
26.Valdés JA,Flores S,Fuentes EN,Osorio-Fuentealba C,Jaimovich E,Molina A.IGF-1 induces IP3-dependent calcium signal involved in the regulation of myostatin gene expression mediated by NFAT during myoblast differentiation.J Cell Physiol.2013;228:1452-63.
27.Zuloaga R,Fuentes EN,Molina A,Valdés JA.The cAMP response element binding protein(CREB)is activated by insulin-like growth factor-1(IGF-1)and regulates myostatin gene expression in skeletal myoblast.Biochem Biophys Res Commun.2013;440:258-64.
28.Grade CVC,Mantovani CS,Fontoura MA,Yusuf F,Brand-Saberi B,Alvares LE.CREB,NF-Y and MEIS1 conserved binding sites are essential to balance Myostatin promoter/enhancer activity during early myogenesis.Mol Biol Rep.2017;44:419-27.
29.Rossi G,Antonini S,Bonfanti C,Monteverde S,Vezzali C,Tajbakhsh S,et al.Nfix regulates temporal progression of muscle regeneration through modulation of myostatin expression.Cell Rep.2016;14:2238-49.
30.Pearen MA,Ryall JG,Maxwell MA,Ohkura N,Lynch GS,Muscat GEO.The orphan nuclear receptor,NOR-1,is a target of beta-adrenergic signaling in skeletal muscle.Endocrinology.2006;147:5217-27.
31.Allen DL,Cleary AS,Hanson AM,Lindsay SF,Reed JM.CCAAT/enhancer binding protein-delta expression is increased in fast skeletal muscle by food deprivation and regulates myostatin transcription in vitro.Am J Physiol Regul Integr Comp Physiol.2010;299:R1592-601.
32.Deng B,Wen J,Ding Y,Gao Q,Huang H,Ran Z,et al.Functional analysis of pig myostatin gene promoter with some adipogenesis-and myogenesisrelated factors.Mol Cell Biochem.2012;363:291-9.
33.Qin J,Du R,Yang Y-Q,Zhang H-Q,Li Q,Liu L,et al.Dexamethasoneinduced skeletal muscle atrophy was associated with upregulation of myostatin promoter activity.Res Vet Sci.2013;94:84-9.
34.Allen DL,Unterman TG.Regulation of myostatin expression and myoblast differentiation by FoxO and SMAD transcription factors.Am J Physiol Cell Physiol.2007;292:C188-99.
35.Carneiro I,Castro-Piedras I,Mu?oz A,Labandeira-García JL,Devesa J,Arce VM.Hypothyroidism is associated with increased myostatin expression in rats.J Endocrinol Invest.2008;31:773-8.
36.Qiu J,Thapaliya S,Runkana A,Yang Y,Tsien C,Mohan ML,et al.Hyperammonemia in cirrhosis induces transcriptional regulation of myostatin by an NF-κB-mediated mechanism.Proc Natl Acad Sci U S A.2013;110:18162-7.
37.Sundaresan NR,Saxena VK,Singh R,Jain P,Singh KP,Anish D,et al.Expression profile of myostatin mRNA during the embryonic organogenesis of domestic chicken(Gallus Gallus domesticus).Res Vet Sci.2008;85:86-91.
38.Allen DL,Cleary AS,Speaker KJ,Lindsay SF,Uyenishi J,Reed JM,et al.Myostatin,activin receptor IIb,and follistatin-like-3 gene expression are altered in adipose tissue and skeletal muscle of obese mice.Am J Physiol Endocrinol Metab.2008;294:E918-27.
39.Wang B-W,Chang H,Kuan P,Shyu K-G.Angiotensin II activates myostatin expression in cultured rat neonatal cardiomyocytes via p38 MAP kinase and myocyte enhance factor 2 pathway.J Endocrinol.2008;197:85-93.
40.CrisàA,Marchitelli C,Savarese MC,Valentini A.Sequence analysis of myostatin promoter in cattle.Cytogenet Genome Res.2003;102:48-52.
41.Forbes D,Jackman M,Bishop A,Thomas M,Kambadur R,Sharma M.Myostatin auto-regulates its expression by feedback loop through Smad7dependent mechanism.J Cell Physiol.2006;206:264-72.
42.Han S-H,Cho I-C,Ko M-S,Kim E-Y,Park S-P,Lee S-S,et al.A promoter polymorphism of MSTN g.-371T>A and its associations with carcass traits in Korean cattle.Mol Biol Rep.2012;39:3767-72.
43.He YL,Wu YH,Quan FS,Liu YG,Zhang Y.Comparative analysis of myostatin gene and promoter sequences of Qinchuan and Red Angus cattle.Genet Mol Res.2013;12:3398-406.
44.Sarti FM,Lasagna E,Ceccobelli S,Di Lorenzo P,Filippini F,Sbarra F,et al.Influence of single nucleotide polymorphisms in the myostatin and myogenic factor 5 muscle growth-related genes on the performance traits of Marchigiana beef cattle.J Anim Sci.2014;92:3804-10.
45.Jank M,Zwierzchowski L,Siadkowska E,Budasz-?widerska M,Sadkowski T,Oprz?dek J,et al.Polymorphism in the 5’flanking region of the myostatin gene affects myostatin and TGF-β1expression in bovine skeletal muscle.JAnim Feed Sci.2006;15:381-91.
46.Stinckens A,Luyten T,Bijttebier J,Van den Maagdenberg K,Dieltiens D,Janssens S,et al.Characterization of the complete porcine MSTN gene and expression levels in pig breeds differing in muscularity.Anim Genet.2008;39:586-96.
47.Yu Z,Li Y,Meng Q,Yuan J,Zhao Z,Li W,et al.Comparative analysis of the pig BAC sequence involved in the regulation of myostatin gene.Sci China C Life Sci.2005;48:168-80.
48.Li J,Deng J,Yu S,Zhang J,Cheng D,Wang H.The virtual element in proximal promoter of porcine myostatin is regulated by myocyte enhancer factor 2C.Biochem Biophys Res Commun.2012;419:175-81.
49.Yu L,Tang H,Wang J,Wu Y,Zou L,Jiang Y,et al.Polymorphisms in the 5’regulatory region of myostatin gene are associated with early growth traits in Yorkshire pigs.Sci China C Life Sci.2007;50:642-7.
50.Liu X,Wang J,Li R,Yang X,Sun Q,Albrecht E,et al.Maternal dietary protein affects transcriptional regulation of myostatin gene distinctively at weaning and finishing stages in skeletal muscle of Meishan pigs.Epigenetics.2011;6:899-907.
51.Jia Y,Gao G,Song H,Cai D,Yang X,Zhao R.Low-protein diet fed to crossbred sows during pregnancy and lactation enhances myostatin gene expression through epigenetic regulation in skeletal muscle of weaning piglets.Eur J Nutr.2016;55:1307-14.
52.Tu P-A,Lo L-L,Chen Y-C,Hsu C-C,Shiau J-W,Lin E-C,et al.Polymorphisms in the promoter region of myostatin gene are associated with carcass traits in pigs.J Anim Breed Genet.2014;131:116-22.
53.Guimaraes SEF,Stahl CH,Lonergan SM,Geiger B,Rothschild MF.Myostatin promoter analysis and expression pattern in pigs.Livest Sci.2007;112:143-50.
54.Liu D,Xu Q,Zang L,Liang S,Wu Y,Wei S,et al.Identification and genetic effect of haplotypes in the promoter region of porcine myostatin gene.Anim Genet.2011;42:6-14.
55.Tu P-A,Shiau J-W,Ding S-T,Lin E-C,Wu M-C,Wang P-H.The association of genetic variations in the promoter region of myostatin gene with growth traits in Duroc pigs.Anim Biotechnol.2012;23:291-8.
56.Bongiorni S,Tilesi F,Bicorgna S,Iacoponi F,Willems D,Gargani M,et al.Promoter polymorphisms in genes involved in porcine myogenesis influence their transcriptional activity.BMC Genet.2014;15:119.
57.Du R,Chen Y-F,An X-R,Yang X-Y,Ma Y,Zhang L,et al.Cloning and sequence analysis of myostatin promoter in sheep.DNA Seq J DNA Seq Mapp.2005;16:412-7.
58.Song XC,Xu C,Yue ZG,Wang L,Wang GW,Yang FH.Bioinformatic analysis based on the complete coding region of the MSTN gene within and among different species.Genet Mol Res.2016;15.https://doi.org/10.4238/gmr.15025031.
59.Du R,An X,Chen Y,Qin J.Functional analysis of the Myostatin gene promoter in sheep.Sci China C Life Sci.2007;50:648-54.
60.Du R,An X-R,Chen Y-F,Qin J.Some motifs were important for myostatin transcriptional regulation in sheep(Ovis aries).J Biochem Mol Biol.2007;40:547-53.
61.Wang J,Zhou H,Hu J,Li S,Luo Y,Hickford JGH.Two single nucleotide polymorphisms in the promoter of the ovine myostatin gene(MSTN)and their effect on growth and carcass muscle traits in New Zealand Romney sheep.J Anim Breed Genet.2016;133:219-26.
62.Singh SP,Kumar R,Kumari P,Kumar S,Mitra A.Characterization of 5’upstream region and investigation of TTTTA deletion in 5’UTR of myostatin(MSTN)gene in Indian goat breeds.Anim Biotechnol.2014;25:55-68.
63.Li X,Liu Z,Zhou R,Zheng G,Gong Y,Li L.Deletion of TTTTA in 5′UTRof goat MSTN gene and its distribution in different population groups and genetic effect on bodyweight at different ages.Front Agric China.2008;2:103-9.
64.Dall’Olio S,Fontanesi L,Nanni Costa L,Tassinari M,Minieri L,Falaschini A.Analysis of horse myostatin gene and identification of single nucleotide polymorphisms in breeds of different morphological types.J Biomed Biotechnol.2010;2010.https://doi.org/10.1155/2010/542945.
65.Stefaniuk M,Kaczor U,Augustyn R,Gurgul A,Kulisa M,Podstawski Z.Identification of a new haplotype within the promoter region of the MSTNgene in horses from five of the most common breeds in Poland.Folia Biol(Praha).2014;62:219-22.
66.Hill EW,McGivney BA,Gu J,Whiston R,Machugh DE.A genome-wide SNP-association study confirms a sequence variant(g.66493737C>T)in the equine myostatin(MSTN)gene as the most powerful predictor of optimum racing distance for Thoroughbred racehorses.BMC Genomics.2010;11:552.
67.Petersen JL,Mickelson JR,Rendahl AK,Valberg SJ,Andersson LS,Axelsson J,et al.Genome-wide analysis reveals selection for important traits in domestic horse breeds.PLoS Genet.2013;9:e1003211.
68.van den Hoven R,Gür E,Schlamanig M,Hofer M,Onmaz AC,Steinborn R.Putative regulation mechanism for the MSTN gene by a CpG island generated by the SINE marker Ins227bp.BMC Vet Res.2015;11:138.
69.Dall’Olio S,Scotti E,Fontanesi L,Tassinari M.Analysis of the 227 bp short interspersed nuclear element(SINE)insertion of the promoter of the myostatin(MSTN)gene in different horse breeds.Vet Ital.2014;50:193-7.
70.Rooney MF,Hill EW,Kelly VP,Porter RK.The“speed gene”effect of myostatin arises in Thoroughbred horses due to a promoter proximal SINEinsertion.PloS One.2018;13:e0205664.
71.Santagostino M,Khoriauli L,Gamba R,Bonuglia M,Klipstein O,Piras FM,et al.Genome-wide evolutionary and functional analysis of the Equine Repetitive Element 1:an insertion in the myostatin promoter affects gene expression.BMC Genet.2015;16:126.
72.Petersen JL,Valberg SJ,Mickelson JR,McCue ME.Haplotype diversity in the equine myostatin gene with focus on variants associated with race distance propensity and muscle fiber type proportions.Anim Genet.2014;45:827-35.
73.Gu Z,Zhang Y,Shi P,Zhang Y-P,Zhu D,Li H.Comparison of avian myostatin genes.Anim Genet.2004;35:470-2.
74.Hu W,Chen S,Zhang R,Lin Y.Single nucleotide polymorphisms in the upstream regulatory region alter the expression of myostatin.In Vitro Cell Dev Biol Anim.2013;49:417-23.
75.Paswan C,Bhattacharya TK,Nagaraj CS,Chaterjee RN,Jayashankar MR.SNPs in minimal promoter of myostatin(GDF-8)gene and its association with body weight in broiler chicken.J Appl Anim Res.2014;42:304-9.
76.Kumar STB,Dilbaghi N,Ahlawat SPS,Mishra B,Tantia MS,Vijh RK.Genetic relationship among chicken populations of india based on SNP markers of Myostatin Gene(GDF 8).Int J Poult Sci.2007;6:684-8.
77.Genxi Z,Guojun D,Jinyu W,Yue W,Fuxiang D,Zhang L,et al.Polymorphisms in 5’-upstream region of the myostatin gene in four chicken breeds and its relationship with growth traits in the Bian chicken.Afr JBiotechnol.2012;11:9677-9682.
78.Lu J,Hou S,Huang W,Yu J,Wang W.Polymorphisms in the myostatin gene and their association with growth and carcass traits in duck.Afr JBiotechnol.2011;10:11309-12.
79.Meyer A,Schartl M.Gene and genome duplications in vertebrates:the oneto-four(-to-eight in fish)rule and the evolution of novel gene functions.Curr Opin Cell Biol.1999;11:699-704.
80.Ostbye T-KK,Wetten OF,Tooming-Klunderud A,Jakobsen KS,Yafe A,Etzioni S,et al.Myostatin(MSTN)gene duplications in Atlantic salmon(Salmo salar):evidence for different selective pressure on teleost MSTN-1 and-2.Gene.2007;403:159-69.
81.Xu C,Wu G,Zohar Y,Du S-J.Analysis of myostatin gene structure,expression and function in zebrafish.J Exp Biol.2003;206:4067-79.
82.Kerr T,Roalson EH,Rodgers BD.Phylogenetic analysis of the myostatin gene sub-family and the differential expression of a novel member in zebrafish.Evol Dev.2005;7:390-400.
83.Funkenstein B,Balas V,Rebhan Y,Pliatner A.Characterization and functional analysis of the 5’flanking region of Sparus aurata myostatin-1 gene.Comp Biochem Physiol A Mol Integr Physiol.2009;153:55-62.
84.Nadjar-Boger E,Funkenstein B.Myostatin-2 gene structure and polymorphism of the promoter and first intron in the marine fish Sparus aurata:evidence for DNA duplications and/or translocations.BMC Genet.2011;12:22.
85.Nadjar-Boger E,Hinits Y,Funkenstein B.Structural and functional analysis of myostatin-2 promoter alleles from the marine fish Sparus aurata:evidence for strong muscle-specific promoter activity and post-transcriptional regulation.Mol Cell Endocrinol.2012;361:51-68.
86.Xue L,Dong X,Zhang X,Diallo A.Organization and functional analysis of the 5’flanking regions of myostatin-1 and 2 genes from Larimichthys crocea.DNA Cell Biol.2012;31:845-55.
87.Chen Y-M,Wei C-Y,Chien C-H,Chang H-W,Huang S-I,Yang H-L,et al.Myostatin gene organization and nodavirus-influenced expression in orange-spotted grouper(Epinephelus coioides).Comp Biochem Physiol Part D Genomics Proteomics.2007;2:215-27.
88.Ye H-Q,Chen S-L,Xu J-Y.Molecular cloning and characterization of the Myf5 gene in sea perch(Lateolabrax japonicus).Comp Biochem Physiol BBiochem Mol Biol.2007;147:452-9.
89.De Santis C,Evans BS,Smith-Keune C,Jerry DR.Molecular characterization,tissue expression and sequence variability of the barramundi(Lates calcarifer)myostatin gene.BMC Genomics.2008;9:82.
90.Li H,Fan J,Liu S,Yang Q,Mu G,He C.Characterization of a myostatin gene(MSTN1)from spotted halibut(Verasper variegatus)and association between its promoter polymorphism and individual growth performance.Comp Biochem Physiol B Biochem Mol Biol.2012;161:315-22.
91.Nadjar-Boger E,Maccatrozzo L,Radaelli G,Funkenstein B.Genomic cloning and promoter functional analysis of myostatin-2 in shi drum,Umbrina cirrosa:conservation of muscle-specific promoter activity.Comp Biochem Physiol B Biochem Mol Biol.2013;164:99-110.
92.Roberts SB,Goetz FW.Myostatin protein and RNA transcript levels in adult and developing brook trout.Mol Cell Endocrinol.2003;210:9-20.
93.Garikipati DK,Gahr SA,Rodgers BD.Identification,characterization,and quantitative expression analysis of rainbow trout myostatin-1a and myostatin-1b genes.J Endocrinol.2006;190:879-88.
94.Garikipati DK,Gahr SA,Roalson EH,Rodgers BD.Characterization of rainbow trout myostatin-2 genes(rtMSTN-2a and-2b):genomic organization,differential expression,and pseudogenization.Endocrinology.2007;148:2106-15.
95.Hu X,Guo H,He Y,Wang S,Zhang L,Wang S,et al.Molecular characterization of myostatin gene from Zhikong scallop Chlamys farreri(Jones et Preston 1904).Genes Genet Syst.2010;85:207-18.
96.Zhuo RQ,Zhou TT,Yang SP,Chan SF.Characterization of a molt-related myostatin gene(FmMstn)from the banana shrimp Fenneropenaeus merguiensis.Gen Comp Endocrinol.2017;248:55-68.
97.Li S,Zhou Z,Dong Y,Sun H,Gao S,Chen Z,et al.Molecular characterization,expression analysis of the myostatin gene and its association with growth traits in sea cucumber(Apostichopus japonicus).Comp Biochem Physiol B Biochem Mol Biol.2016;201:12-20.
98.Allen DL,Du M.Comparative functional analysis of the cow and mouse myostatin genes reveals novel regulatory elements in their upstream promoter regions.Comp Biochem Physiol B Biochem Mol Biol.2008;150:432-9.
99.He Y,Wu Y,Lan Z,Liu Y,Zhang Y.Molecular analysis of the first intron in the bovine myostatin gene.Mol Biol Rep.2011;38:4643-9.
100.Han D-S,Huang H-P,Wang T-G,Hung M-Y,Ke J-Y,Chang K-T,et al.Transcription activation of myostatin by trichostatin A in differentiated C2C12 myocytes via ASK1-MKK3/4/6-JNK and p38 mitogen-activated protein kinase pathways.J Cell Biochem.2010;111:564-73.
101.Abbas EM,Takayanagi A,Shimizu N,Kato M.Methylation status and chromatin structure of the myostatin gene promoter region in the sea perch Lateolabrax japonicus(Perciformes).Genet Mol Res.2011;10:3306-15.
102.Nogalska A,Wojcik S,Engel WK,McFerrin J,Askanas V.Endoplasmic reticulum stress induces myostatin precursor protein and NF-kappaB in cultured human muscle fibers:relevance to inclusion body myositis.Exp Neurol.2007;204:610-8.
103.Samant SA,Kanwal A,Pillai VB,Bao R,Gupta MP.The histone deacetylase SIRT6 blocks myostatin expression and development of muscle atrophy.Sci Rep.2017;7:11877.
104.Roberts TC,Andaloussi SE,Morris KV,McClorey G,Wood MJ.Small RNA-mediated epigenetic myostatin silencing.Mol Ther Nucleic Acids.2012;1:e23.
105.Yu R,Chen J-A,Xu J,Cao J,Wang Y,Thomas SS,et al.Suppression of muscle wasting by the plant-derived compound ursolic acid in a model of chronic kidney disease.J Cachexia Sarcopenia Muscle.2017;8:327-41.
106.Salehian B,Mahabadi V,Bilas J,Taylor WE,Ma K.The effect of glutamine on prevention of glucocorticoid-induced skeletal muscle atrophy is associated with myostatin suppression.Metabolism.2006;55:1239-47.
107.Zhao LH,Zhao YH,Liang H,Yun T,Han XJ,Zhang ML,et al.A promoter trap vector for knocking out bovine myostatin gene with high targeting efficiency.Genet Mol Res.2015;14:2750-61.
108.Lee SJ,McPherron AC.Regulation of myostatin activity and muscle growth.Proc Natl Acad Sci U S A.2001;98:9306-11.
109.McFarlane C,Langley B,Thomas M,Hennebry A,Plummer E,Nicholas G,et al.Proteolytic processing of myostatin is auto-regulated during myogenesis.Dev Biol.2005;283:58-69.
110.Ovcharenko I,Loots GG,Giardine BM,Hou M,Ma J,Hardison RC,et al.Mulan:multiple-sequence local alignment and visualization for studying function and evolution.Genome Res.2005;15:184-94.
111.Katoh K,Standley DM.MAFFT multiple sequence alignment software version 7:improvements in performance and usability.Mol Biol Evol.2013;30:772-80.
112.Erb I,González-Vallinas JR,Bussotti G,Blanco E,Eyras E,Notredame C.Use of ChIP-Seq data for the design of a multiple promoter-alignment method.Nucleic Acids Res.2012;40:e52.
113.Hal TA.Bio Edit:a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT.Nucleic Acids Symp Ser.1999;41:95-8.
114.Jiang Y-L,Li N,Du L-X,Wu C-X.Relationship of T-->A mutation in the promoter region of myostatin gene with growth traits in swine.Yi Chuan Xue Bao.2002;29:413-6.
115.Sternstein I,Reissmann M,Maj D,Bieniek J,Brockmann GA.A new single nucleotide polymorphism in the rabbit(Oryctolagus cuniculus)myostatin(MSTN)gene is associated with carcass composition traits.Anim Genet.2014;45:596-9.
116.Qiao XB,Xu KY,Li B,Luan X,Xia T,Fan XZ.Rabbit MSTN gene polymorphisms and genetic effect analysis.Genet Mol Res.2014;13:2590-7.
117.Pe?aloza C,Hamilton A,Guy DR,Bishop SC,Houston RD.A SNP in the 5′flanking region of the myostatin-1b gene is associated with harvest traits in Atlantic salmon(Salmo salar).BMC Genet.2013;14:112.
118.Fan S,Xu Y,Liu B,He W,Zhang B,Su J,et al.Molecular characterization and expression analysis of the myostatin gene and its association with growth traits in Noble scallop(Chlamys nobilis).Comp Biochem Physiol B Biochem Mol Biol.2017;212:24-31.
119.Ferrell RE,Conte V,Lawrence EC,Roth SM,Hagberg JM,Hurley BF.Frequent sequence variation in the human myostatin(GDF8)gene as a marker for analysis of muscle-related phenotypes.Genomics.1999;62:203-7.
2.Joulia D,Bernardi H,Garandel V,Rabenoelina F,Vernus B,Cabello G.Mechanisms involved in the inhibition of myoblast proliferation and differentiation by myostatin.Exp Cell Res.2003;286:263-75.
3.McPherron AC,Lee SJ.Double muscling in cattle due to mutations in the myostatin gene.Proc Natl Acad Sci U S A.1997;94:12457-61.
4.Clop A,Marcq F,Takeda H,Pirottin D,Tordoir X,BibéB,et al.A mutation creating a potential illegitimate microRNA target site in the myostatin gene affects muscularity in sheep.Nat Genet.2006;38:813-8.
5.Mosher DS,Quignon P,Bustamante CD,Sutter NB,Mellersh CS,Parker HG,et al.A mutation in the myostatin gene increases muscle mass and enhances racing performance in heterozygote dogs.PLoS Genet.2007;3:e79.
6.Shelton GD,Engvall E.Gross muscle hypertrophy in whippet dogs is caused by a mutation in the myostatin gene.Neuromuscul Disord NMD.2007;17:721-2.
7.Schuelke M,Wagner KR,Stolz LE,Hübner C,Riebel T,K?men W,et al.Myostatin mutation associated with gross muscle hypertrophy in a child.NEngl J Med.2004;350:2682-8.
8.Jespersen J,Kjaer M,Schjerling P.The possible role of myostatin in skeletal muscle atrophy and cachexia.Scand J Med Sci Sports.2006;16:74-82.
9.Joulia-Ekaza D,Cabello G.The myostatin gene:physiology and pharmacological relevance.Curr Opin Pharmacol.2007;7:310-5.
10.Wójcik S,Engel WK,McFerrin J,Askanas V.Myostatin is increased and complexes with amyloid-beta within sporadic inclusion-body myositis muscle fibers.Acta Neuropathol(Berl).2005;110:173-7.
11.Bogdanovich S,Krag TOB,Barton ER,Morris LD,Whittemore L-A,Ahima RS,et al.Functional improvement of dystrophic muscle by myostatin blockade.Nature.2002;420:418-21.
12.Tang L,Yan Z,Wan Y,Han W,Zhang Y.Myostatin DNA vaccine increases skeletal muscle mass and endurance in mice.Muscle Nerve.2007;36:342-8.
13.Takata H,Terada K,Oka H,Sunada Y,Moriguchi T,Nohno T.Involvement of Wnt4 signaling during myogenic proliferation and differentiation of skeletal muscle.Dev Dyn Off Publ Am Assoc Anat.2007;236:2800-7.
14.Allen DL,Loh AS.Posttranscriptional mechanisms involving microRNA-27a and b contribute to fast-specific and glucocorticoid-mediated myostatin expression in skeletal muscle.Am J Physiol Cell Physiol.2011;300:C124-37.
15.Wilt F,Hake S.Principles of developmental biology.New York:W.W.Norton&Company,Inc;2004.
16.Heintzman ND,Ren B.The gateway to transcription:identifying,characterizing and understanding promoters in the eukaryotic genome.Cell Mol Life Sci.2007;64:386-400.
17.Ma K,Mallidis C,Artaza J,Taylor W,Gonzalez-Cadavid N,Bhasin S.Characterization of 5’-regulatory region of human myostatin gene:regulation by dexamethasone in vitro.Am J Physiol Endocrinol Metab.2001;281:E1128-36.
18.Zammit PS.Function of the myogenic regulatory factors Myf5,MyoD,Myogenin and MRF4 in skeletal muscle,satellite cells and regenerative myogenesis.Semin Cell Dev Biol.2017;72:19-32.
19.Spiller MP,Kambadur R,Jeanplong F,Thomas M,Martyn JK,Bass JJ,et al.The myostatin gene is a downstream target gene of basic helix-loop-helix transcription factor MyoD.Mol Cell Biol.2002;22:7066-82.
20.Fan H,Zhang R,Tesfaye D,Tholen E,Looft C,H?lker M,et al.Sulforaphane causes a major epigenetic repression of myostatin in porcine satellite cells.Epigenetics.2012;7:1379-90.
21.Taylor MV,Hughes SM.Mef2 and the skeletal muscle differentiation program.Semin Cell Dev Biol.2017;72:33-44.
22.Ríos R,Carneiro I,Arce VM,Devesa J.Myostatin regulates cell survival during C2C12 myogenesis.Biochem Biophys Res Commun.2001;280:561-6.
23.Artaza JN,Bhasin S,Mallidis C,Taylor W,Ma K,Gonzalez-Cadavid NF.Endogenous expression and localization of myostatin and its relation to myosin heavy chain distribution in C2C12 skeletal muscle cells.J Cell Physiol.2002;190:170-9.
24.Salerno MS,Thomas M,Forbes D,Watson T,Kambadur R,Sharma M.Molecular analysis of fiber type-specific expression of murine myostatin promoter.Am J Physiol Cell Physiol.2004;287:C1031-40.
25.Grade CVC,Salerno MS,Schubert FR,Dietrich S,Alvares LE.An evolutionarily conserved Myostatin proximal promoter/enhancer confers basal levels of transcription and spatial specificity in vivo.Dev Genes Evol.2009;219:497-508.
26.Valdés JA,Flores S,Fuentes EN,Osorio-Fuentealba C,Jaimovich E,Molina A.IGF-1 induces IP3-dependent calcium signal involved in the regulation of myostatin gene expression mediated by NFAT during myoblast differentiation.J Cell Physiol.2013;228:1452-63.
27.Zuloaga R,Fuentes EN,Molina A,Valdés JA.The cAMP response element binding protein(CREB)is activated by insulin-like growth factor-1(IGF-1)and regulates myostatin gene expression in skeletal myoblast.Biochem Biophys Res Commun.2013;440:258-64.
28.Grade CVC,Mantovani CS,Fontoura MA,Yusuf F,Brand-Saberi B,Alvares LE.CREB,NF-Y and MEIS1 conserved binding sites are essential to balance Myostatin promoter/enhancer activity during early myogenesis.Mol Biol Rep.2017;44:419-27.
29.Rossi G,Antonini S,Bonfanti C,Monteverde S,Vezzali C,Tajbakhsh S,et al.Nfix regulates temporal progression of muscle regeneration through modulation of myostatin expression.Cell Rep.2016;14:2238-49.
30.Pearen MA,Ryall JG,Maxwell MA,Ohkura N,Lynch GS,Muscat GEO.The orphan nuclear receptor,NOR-1,is a target of beta-adrenergic signaling in skeletal muscle.Endocrinology.2006;147:5217-27.
31.Allen DL,Cleary AS,Hanson AM,Lindsay SF,Reed JM.CCAAT/enhancer binding protein-delta expression is increased in fast skeletal muscle by food deprivation and regulates myostatin transcription in vitro.Am J Physiol Regul Integr Comp Physiol.2010;299:R1592-601.
32.Deng B,Wen J,Ding Y,Gao Q,Huang H,Ran Z,et al.Functional analysis of pig myostatin gene promoter with some adipogenesis-and myogenesisrelated factors.Mol Cell Biochem.2012;363:291-9.
33.Qin J,Du R,Yang Y-Q,Zhang H-Q,Li Q,Liu L,et al.Dexamethasoneinduced skeletal muscle atrophy was associated with upregulation of myostatin promoter activity.Res Vet Sci.2013;94:84-9.
34.Allen DL,Unterman TG.Regulation of myostatin expression and myoblast differentiation by FoxO and SMAD transcription factors.Am J Physiol Cell Physiol.2007;292:C188-99.
35.Carneiro I,Castro-Piedras I,Mu?oz A,Labandeira-García JL,Devesa J,Arce VM.Hypothyroidism is associated with increased myostatin expression in rats.J Endocrinol Invest.2008;31:773-8.
36.Qiu J,Thapaliya S,Runkana A,Yang Y,Tsien C,Mohan ML,et al.Hyperammonemia in cirrhosis induces transcriptional regulation of myostatin by an NF-κB-mediated mechanism.Proc Natl Acad Sci U S A.2013;110:18162-7.
37.Sundaresan NR,Saxena VK,Singh R,Jain P,Singh KP,Anish D,et al.Expression profile of myostatin mRNA during the embryonic organogenesis of domestic chicken(Gallus Gallus domesticus).Res Vet Sci.2008;85:86-91.
38.Allen DL,Cleary AS,Speaker KJ,Lindsay SF,Uyenishi J,Reed JM,et al.Myostatin,activin receptor IIb,and follistatin-like-3 gene expression are altered in adipose tissue and skeletal muscle of obese mice.Am J Physiol Endocrinol Metab.2008;294:E918-27.
39.Wang B-W,Chang H,Kuan P,Shyu K-G.Angiotensin II activates myostatin expression in cultured rat neonatal cardiomyocytes via p38 MAP kinase and myocyte enhance factor 2 pathway.J Endocrinol.2008;197:85-93.
40.CrisàA,Marchitelli C,Savarese MC,Valentini A.Sequence analysis of myostatin promoter in cattle.Cytogenet Genome Res.2003;102:48-52.
41.Forbes D,Jackman M,Bishop A,Thomas M,Kambadur R,Sharma M.Myostatin auto-regulates its expression by feedback loop through Smad7dependent mechanism.J Cell Physiol.2006;206:264-72.
42.Han S-H,Cho I-C,Ko M-S,Kim E-Y,Park S-P,Lee S-S,et al.A promoter polymorphism of MSTN g.-371T>A and its associations with carcass traits in Korean cattle.Mol Biol Rep.2012;39:3767-72.
43.He YL,Wu YH,Quan FS,Liu YG,Zhang Y.Comparative analysis of myostatin gene and promoter sequences of Qinchuan and Red Angus cattle.Genet Mol Res.2013;12:3398-406.
44.Sarti FM,Lasagna E,Ceccobelli S,Di Lorenzo P,Filippini F,Sbarra F,et al.Influence of single nucleotide polymorphisms in the myostatin and myogenic factor 5 muscle growth-related genes on the performance traits of Marchigiana beef cattle.J Anim Sci.2014;92:3804-10.
45.Jank M,Zwierzchowski L,Siadkowska E,Budasz-?widerska M,Sadkowski T,Oprz?dek J,et al.Polymorphism in the 5’flanking region of the myostatin gene affects myostatin and TGF-β1expression in bovine skeletal muscle.JAnim Feed Sci.2006;15:381-91.
46.Stinckens A,Luyten T,Bijttebier J,Van den Maagdenberg K,Dieltiens D,Janssens S,et al.Characterization of the complete porcine MSTN gene and expression levels in pig breeds differing in muscularity.Anim Genet.2008;39:586-96.
47.Yu Z,Li Y,Meng Q,Yuan J,Zhao Z,Li W,et al.Comparative analysis of the pig BAC sequence involved in the regulation of myostatin gene.Sci China C Life Sci.2005;48:168-80.
48.Li J,Deng J,Yu S,Zhang J,Cheng D,Wang H.The virtual element in proximal promoter of porcine myostatin is regulated by myocyte enhancer factor 2C.Biochem Biophys Res Commun.2012;419:175-81.
49.Yu L,Tang H,Wang J,Wu Y,Zou L,Jiang Y,et al.Polymorphisms in the 5’regulatory region of myostatin gene are associated with early growth traits in Yorkshire pigs.Sci China C Life Sci.2007;50:642-7.
50.Liu X,Wang J,Li R,Yang X,Sun Q,Albrecht E,et al.Maternal dietary protein affects transcriptional regulation of myostatin gene distinctively at weaning and finishing stages in skeletal muscle of Meishan pigs.Epigenetics.2011;6:899-907.
51.Jia Y,Gao G,Song H,Cai D,Yang X,Zhao R.Low-protein diet fed to crossbred sows during pregnancy and lactation enhances myostatin gene expression through epigenetic regulation in skeletal muscle of weaning piglets.Eur J Nutr.2016;55:1307-14.
52.Tu P-A,Lo L-L,Chen Y-C,Hsu C-C,Shiau J-W,Lin E-C,et al.Polymorphisms in the promoter region of myostatin gene are associated with carcass traits in pigs.J Anim Breed Genet.2014;131:116-22.
53.Guimaraes SEF,Stahl CH,Lonergan SM,Geiger B,Rothschild MF.Myostatin promoter analysis and expression pattern in pigs.Livest Sci.2007;112:143-50.
54.Liu D,Xu Q,Zang L,Liang S,Wu Y,Wei S,et al.Identification and genetic effect of haplotypes in the promoter region of porcine myostatin gene.Anim Genet.2011;42:6-14.
55.Tu P-A,Shiau J-W,Ding S-T,Lin E-C,Wu M-C,Wang P-H.The association of genetic variations in the promoter region of myostatin gene with growth traits in Duroc pigs.Anim Biotechnol.2012;23:291-8.
56.Bongiorni S,Tilesi F,Bicorgna S,Iacoponi F,Willems D,Gargani M,et al.Promoter polymorphisms in genes involved in porcine myogenesis influence their transcriptional activity.BMC Genet.2014;15:119.
57.Du R,Chen Y-F,An X-R,Yang X-Y,Ma Y,Zhang L,et al.Cloning and sequence analysis of myostatin promoter in sheep.DNA Seq J DNA Seq Mapp.2005;16:412-7.
58.Song XC,Xu C,Yue ZG,Wang L,Wang GW,Yang FH.Bioinformatic analysis based on the complete coding region of the MSTN gene within and among different species.Genet Mol Res.2016;15.https://doi.org/10.4238/gmr.15025031.
59.Du R,An X,Chen Y,Qin J.Functional analysis of the Myostatin gene promoter in sheep.Sci China C Life Sci.2007;50:648-54.
60.Du R,An X-R,Chen Y-F,Qin J.Some motifs were important for myostatin transcriptional regulation in sheep(Ovis aries).J Biochem Mol Biol.2007;40:547-53.
61.Wang J,Zhou H,Hu J,Li S,Luo Y,Hickford JGH.Two single nucleotide polymorphisms in the promoter of the ovine myostatin gene(MSTN)and their effect on growth and carcass muscle traits in New Zealand Romney sheep.J Anim Breed Genet.2016;133:219-26.
62.Singh SP,Kumar R,Kumari P,Kumar S,Mitra A.Characterization of 5’upstream region and investigation of TTTTA deletion in 5’UTR of myostatin(MSTN)gene in Indian goat breeds.Anim Biotechnol.2014;25:55-68.
63.Li X,Liu Z,Zhou R,Zheng G,Gong Y,Li L.Deletion of TTTTA in 5′UTRof goat MSTN gene and its distribution in different population groups and genetic effect on bodyweight at different ages.Front Agric China.2008;2:103-9.
64.Dall’Olio S,Fontanesi L,Nanni Costa L,Tassinari M,Minieri L,Falaschini A.Analysis of horse myostatin gene and identification of single nucleotide polymorphisms in breeds of different morphological types.J Biomed Biotechnol.2010;2010.https://doi.org/10.1155/2010/542945.
65.Stefaniuk M,Kaczor U,Augustyn R,Gurgul A,Kulisa M,Podstawski Z.Identification of a new haplotype within the promoter region of the MSTNgene in horses from five of the most common breeds in Poland.Folia Biol(Praha).2014;62:219-22.
66.Hill EW,McGivney BA,Gu J,Whiston R,Machugh DE.A genome-wide SNP-association study confirms a sequence variant(g.66493737C>T)in the equine myostatin(MSTN)gene as the most powerful predictor of optimum racing distance for Thoroughbred racehorses.BMC Genomics.2010;11:552.
67.Petersen JL,Mickelson JR,Rendahl AK,Valberg SJ,Andersson LS,Axelsson J,et al.Genome-wide analysis reveals selection for important traits in domestic horse breeds.PLoS Genet.2013;9:e1003211.
68.van den Hoven R,Gür E,Schlamanig M,Hofer M,Onmaz AC,Steinborn R.Putative regulation mechanism for the MSTN gene by a CpG island generated by the SINE marker Ins227bp.BMC Vet Res.2015;11:138.
69.Dall’Olio S,Scotti E,Fontanesi L,Tassinari M.Analysis of the 227 bp short interspersed nuclear element(SINE)insertion of the promoter of the myostatin(MSTN)gene in different horse breeds.Vet Ital.2014;50:193-7.
70.Rooney MF,Hill EW,Kelly VP,Porter RK.The“speed gene”effect of myostatin arises in Thoroughbred horses due to a promoter proximal SINEinsertion.PloS One.2018;13:e0205664.
71.Santagostino M,Khoriauli L,Gamba R,Bonuglia M,Klipstein O,Piras FM,et al.Genome-wide evolutionary and functional analysis of the Equine Repetitive Element 1:an insertion in the myostatin promoter affects gene expression.BMC Genet.2015;16:126.
72.Petersen JL,Valberg SJ,Mickelson JR,McCue ME.Haplotype diversity in the equine myostatin gene with focus on variants associated with race distance propensity and muscle fiber type proportions.Anim Genet.2014;45:827-35.
73.Gu Z,Zhang Y,Shi P,Zhang Y-P,Zhu D,Li H.Comparison of avian myostatin genes.Anim Genet.2004;35:470-2.
74.Hu W,Chen S,Zhang R,Lin Y.Single nucleotide polymorphisms in the upstream regulatory region alter the expression of myostatin.In Vitro Cell Dev Biol Anim.2013;49:417-23.
75.Paswan C,Bhattacharya TK,Nagaraj CS,Chaterjee RN,Jayashankar MR.SNPs in minimal promoter of myostatin(GDF-8)gene and its association with body weight in broiler chicken.J Appl Anim Res.2014;42:304-9.
76.Kumar STB,Dilbaghi N,Ahlawat SPS,Mishra B,Tantia MS,Vijh RK.Genetic relationship among chicken populations of india based on SNP markers of Myostatin Gene(GDF 8).Int J Poult Sci.2007;6:684-8.
77.Genxi Z,Guojun D,Jinyu W,Yue W,Fuxiang D,Zhang L,et al.Polymorphisms in 5’-upstream region of the myostatin gene in four chicken breeds and its relationship with growth traits in the Bian chicken.Afr JBiotechnol.2012;11:9677-9682.
78.Lu J,Hou S,Huang W,Yu J,Wang W.Polymorphisms in the myostatin gene and their association with growth and carcass traits in duck.Afr JBiotechnol.2011;10:11309-12.
79.Meyer A,Schartl M.Gene and genome duplications in vertebrates:the oneto-four(-to-eight in fish)rule and the evolution of novel gene functions.Curr Opin Cell Biol.1999;11:699-704.
80.Ostbye T-KK,Wetten OF,Tooming-Klunderud A,Jakobsen KS,Yafe A,Etzioni S,et al.Myostatin(MSTN)gene duplications in Atlantic salmon(Salmo salar):evidence for different selective pressure on teleost MSTN-1 and-2.Gene.2007;403:159-69.
81.Xu C,Wu G,Zohar Y,Du S-J.Analysis of myostatin gene structure,expression and function in zebrafish.J Exp Biol.2003;206:4067-79.
82.Kerr T,Roalson EH,Rodgers BD.Phylogenetic analysis of the myostatin gene sub-family and the differential expression of a novel member in zebrafish.Evol Dev.2005;7:390-400.
83.Funkenstein B,Balas V,Rebhan Y,Pliatner A.Characterization and functional analysis of the 5’flanking region of Sparus aurata myostatin-1 gene.Comp Biochem Physiol A Mol Integr Physiol.2009;153:55-62.
84.Nadjar-Boger E,Funkenstein B.Myostatin-2 gene structure and polymorphism of the promoter and first intron in the marine fish Sparus aurata:evidence for DNA duplications and/or translocations.BMC Genet.2011;12:22.
85.Nadjar-Boger E,Hinits Y,Funkenstein B.Structural and functional analysis of myostatin-2 promoter alleles from the marine fish Sparus aurata:evidence for strong muscle-specific promoter activity and post-transcriptional regulation.Mol Cell Endocrinol.2012;361:51-68.
86.Xue L,Dong X,Zhang X,Diallo A.Organization and functional analysis of the 5’flanking regions of myostatin-1 and 2 genes from Larimichthys crocea.DNA Cell Biol.2012;31:845-55.
87.Chen Y-M,Wei C-Y,Chien C-H,Chang H-W,Huang S-I,Yang H-L,et al.Myostatin gene organization and nodavirus-influenced expression in orange-spotted grouper(Epinephelus coioides).Comp Biochem Physiol Part D Genomics Proteomics.2007;2:215-27.
88.Ye H-Q,Chen S-L,Xu J-Y.Molecular cloning and characterization of the Myf5 gene in sea perch(Lateolabrax japonicus).Comp Biochem Physiol BBiochem Mol Biol.2007;147:452-9.
89.De Santis C,Evans BS,Smith-Keune C,Jerry DR.Molecular characterization,tissue expression and sequence variability of the barramundi(Lates calcarifer)myostatin gene.BMC Genomics.2008;9:82.
90.Li H,Fan J,Liu S,Yang Q,Mu G,He C.Characterization of a myostatin gene(MSTN1)from spotted halibut(Verasper variegatus)and association between its promoter polymorphism and individual growth performance.Comp Biochem Physiol B Biochem Mol Biol.2012;161:315-22.
91.Nadjar-Boger E,Maccatrozzo L,Radaelli G,Funkenstein B.Genomic cloning and promoter functional analysis of myostatin-2 in shi drum,Umbrina cirrosa:conservation of muscle-specific promoter activity.Comp Biochem Physiol B Biochem Mol Biol.2013;164:99-110.
92.Roberts SB,Goetz FW.Myostatin protein and RNA transcript levels in adult and developing brook trout.Mol Cell Endocrinol.2003;210:9-20.
93.Garikipati DK,Gahr SA,Rodgers BD.Identification,characterization,and quantitative expression analysis of rainbow trout myostatin-1a and myostatin-1b genes.J Endocrinol.2006;190:879-88.
94.Garikipati DK,Gahr SA,Roalson EH,Rodgers BD.Characterization of rainbow trout myostatin-2 genes(rtMSTN-2a and-2b):genomic organization,differential expression,and pseudogenization.Endocrinology.2007;148:2106-15.
95.Hu X,Guo H,He Y,Wang S,Zhang L,Wang S,et al.Molecular characterization of myostatin gene from Zhikong scallop Chlamys farreri(Jones et Preston 1904).Genes Genet Syst.2010;85:207-18.
96.Zhuo RQ,Zhou TT,Yang SP,Chan SF.Characterization of a molt-related myostatin gene(FmMstn)from the banana shrimp Fenneropenaeus merguiensis.Gen Comp Endocrinol.2017;248:55-68.
97.Li S,Zhou Z,Dong Y,Sun H,Gao S,Chen Z,et al.Molecular characterization,expression analysis of the myostatin gene and its association with growth traits in sea cucumber(Apostichopus japonicus).Comp Biochem Physiol B Biochem Mol Biol.2016;201:12-20.
98.Allen DL,Du M.Comparative functional analysis of the cow and mouse myostatin genes reveals novel regulatory elements in their upstream promoter regions.Comp Biochem Physiol B Biochem Mol Biol.2008;150:432-9.
99.He Y,Wu Y,Lan Z,Liu Y,Zhang Y.Molecular analysis of the first intron in the bovine myostatin gene.Mol Biol Rep.2011;38:4643-9.
100.Han D-S,Huang H-P,Wang T-G,Hung M-Y,Ke J-Y,Chang K-T,et al.Transcription activation of myostatin by trichostatin A in differentiated C2C12 myocytes via ASK1-MKK3/4/6-JNK and p38 mitogen-activated protein kinase pathways.J Cell Biochem.2010;111:564-73.
101.Abbas EM,Takayanagi A,Shimizu N,Kato M.Methylation status and chromatin structure of the myostatin gene promoter region in the sea perch Lateolabrax japonicus(Perciformes).Genet Mol Res.2011;10:3306-15.
102.Nogalska A,Wojcik S,Engel WK,McFerrin J,Askanas V.Endoplasmic reticulum stress induces myostatin precursor protein and NF-kappaB in cultured human muscle fibers:relevance to inclusion body myositis.Exp Neurol.2007;204:610-8.
103.Samant SA,Kanwal A,Pillai VB,Bao R,Gupta MP.The histone deacetylase SIRT6 blocks myostatin expression and development of muscle atrophy.Sci Rep.2017;7:11877.
104.Roberts TC,Andaloussi SE,Morris KV,McClorey G,Wood MJ.Small RNA-mediated epigenetic myostatin silencing.Mol Ther Nucleic Acids.2012;1:e23.
105.Yu R,Chen J-A,Xu J,Cao J,Wang Y,Thomas SS,et al.Suppression of muscle wasting by the plant-derived compound ursolic acid in a model of chronic kidney disease.J Cachexia Sarcopenia Muscle.2017;8:327-41.
106.Salehian B,Mahabadi V,Bilas J,Taylor WE,Ma K.The effect of glutamine on prevention of glucocorticoid-induced skeletal muscle atrophy is associated with myostatin suppression.Metabolism.2006;55:1239-47.
107.Zhao LH,Zhao YH,Liang H,Yun T,Han XJ,Zhang ML,et al.A promoter trap vector for knocking out bovine myostatin gene with high targeting efficiency.Genet Mol Res.2015;14:2750-61.
108.Lee SJ,McPherron AC.Regulation of myostatin activity and muscle growth.Proc Natl Acad Sci U S A.2001;98:9306-11.
109.McFarlane C,Langley B,Thomas M,Hennebry A,Plummer E,Nicholas G,et al.Proteolytic processing of myostatin is auto-regulated during myogenesis.Dev Biol.2005;283:58-69.
110.Ovcharenko I,Loots GG,Giardine BM,Hou M,Ma J,Hardison RC,et al.Mulan:multiple-sequence local alignment and visualization for studying function and evolution.Genome Res.2005;15:184-94.
111.Katoh K,Standley DM.MAFFT multiple sequence alignment software version 7:improvements in performance and usability.Mol Biol Evol.2013;30:772-80.
112.Erb I,González-Vallinas JR,Bussotti G,Blanco E,Eyras E,Notredame C.Use of ChIP-Seq data for the design of a multiple promoter-alignment method.Nucleic Acids Res.2012;40:e52.
113.Hal TA.Bio Edit:a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT.Nucleic Acids Symp Ser.1999;41:95-8.
114.Jiang Y-L,Li N,Du L-X,Wu C-X.Relationship of T-->A mutation in the promoter region of myostatin gene with growth traits in swine.Yi Chuan Xue Bao.2002;29:413-6.
115.Sternstein I,Reissmann M,Maj D,Bieniek J,Brockmann GA.A new single nucleotide polymorphism in the rabbit(Oryctolagus cuniculus)myostatin(MSTN)gene is associated with carcass composition traits.Anim Genet.2014;45:596-9.
116.Qiao XB,Xu KY,Li B,Luan X,Xia T,Fan XZ.Rabbit MSTN gene polymorphisms and genetic effect analysis.Genet Mol Res.2014;13:2590-7.
117.Pe?aloza C,Hamilton A,Guy DR,Bishop SC,Houston RD.A SNP in the 5′flanking region of the myostatin-1b gene is associated with harvest traits in Atlantic salmon(Salmo salar).BMC Genet.2013;14:112.
118.Fan S,Xu Y,Liu B,He W,Zhang B,Su J,et al.Molecular characterization and expression analysis of the myostatin gene and its association with growth traits in Noble scallop(Chlamys nobilis).Comp Biochem Physiol B Biochem Mol Biol.2017;212:24-31.
119.Ferrell RE,Conte V,Lawrence EC,Roth SM,Hagberg JM,Hurley BF.Frequent sequence variation in the human myostatin(GDF8)gene as a marker for analysis of muscle-related phenotypes.Genomics.1999;62:203-7.