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Article(id=1307013274315289481, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307013104081068276, articleNumber=null, orderNo=null, doi=10.11843/j.issn.0366-6964.2026.02.009, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1750262400000, receivedDateStr=2025-06-19, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1789545847729, onlineDateStr=2026-09-16, pubDate=1771776000000, pubDateStr=2026-02-23, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789545847729, onlineIssueDateStr=2026-09-16, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789545847729, creator=13701087609, updateTime=1789545847729, updator=13701087609, issue=Issue{id=1307013104081068276, tenantId=1146029695717560320, journalId=1306274286668972124, year='2026', volume='57', issue='2', pageStart='569', pageEnd='1176', issueExtLink='null', onlineDate='null', pubDate='1771776000000', pubDateStr='2026-02-23', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789545807142, creator='13701087609', updateTime=1789609910854, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1307281975350419753, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307013104081068276, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1307281975350419754, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307013104081068276, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=668, endPage=676, ext={EN=ArticleExt(id=1307013274483061642, articleId=1307013274315289481, tenantId=1146029695717560320, journalId=1306274286668972124, language=EN, title=Regulatory Role of ZNF697 on the Proliferation and Differentiation of Skeletal Muscle Satellite Cells of Pig, columnId=1307007413870158054, journalTitle=Acta Veterinaria et Zootechnica Sinica, columnName=ANIMAL GENETICS AND BREEDING, runingTitle=null, highlight=null, articleAbstract=

This experiment aimed to investigate the effects of ZNF697 on the proliferation and differentiation of porcine skeletal muscle satellite cells （MuSCs）. In this experiment， porcine MuSCs were used as the research object. Flow cytometry， Real-time quantitative PCR， Western blot and immunofluorescence staining were employed to detect the effects of ZNF697 on the proliferation and differentiation of porcine MuSCs. The following results were obtained： 1） The expression of ZNF697 mRNA dynamically changes with the proliferation of porcine MuSCs. Interfering with ZNF697 inhibited the mRNA expression of Cyclin E （P<0.01） and Cyclin D （P<0.05） and the protein expression of Cyclin B （P<0.05）， promoted the mRNA expression of p21 （P<0.01）. Overexpression of ZNF697 showed the opposite results. 2） During the myogenic differentiation of porcine MuSCs， the expression of ZNF697 gradually increased. Interfering with ZNF697 significantly reduced the myogenic differentiation index （P<0.05） and inhibited the mRNA expression of MyoG （P<0.01）， the mRNA and protein expression of MyHC （P<0.05） and MyoD （P<0.01）. Overexpression of ZNF697 showed the opposite results. The results show that ZNF697 is a promoter of proliferation and myogenic differentiation in porcine MuSCs， which will provide a breeding target for high-lean-meat pig breeds.

, authors=Yang LI, Chenglong WEN, Yuhe WANG, Xiao LI, authorsList=Yang LI, Chenglong WEN, Yuhe WANG, Xiao LI, authorCompany=null, correspAuthors=Xiao LI, authorNote=null, correspAuthorsNote=
LI Xiao， E-mail：
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旨在探究ZNF697对猪骨骼肌卫星细胞（muscle satellite cells， MuSCs）增殖及分化的影响。本试验以猪MuSCs为研究对象，利用流式细胞术、实时荧光定量聚合酶链式反应（qRT-PCR）、蛋白质免疫印迹（Western blot）和免疫荧光染色等方法检测ZNF697对猪MuSCs增殖及分化的影响。获得以下研究结果：1）ZNF697 mRNA的表达量随猪MuSCs增殖而动态变化，干扰ZNF697抑制Cyclin E（P<0.01）、Cyclin D（P<0.05）mRNA表达和Cyclin B（P<0.05）蛋白表达，促进p21（P<0.01）mRNA表达，过表达ZNF697取得相反的结果。2）ZNF697在猪MuSCs成肌分化过程中表达逐渐升高，干扰ZNF697显著降低成肌分化指数（P<0.05），抑制MyoG（P<0.01）mRNA表达、MyHC（P<0.05）、MyoD（P<0.01）mRNA和蛋白表达，过表达ZNF697取得相反的结果。本研究鉴定了猪MuSCs增殖与成肌分化的促进因子——ZNF697，为培育高瘦肉率的优良猪种提供了育种靶点。

, authors=李杨, 文成龙, 王育禾, 李晓, authorsList=李杨, 文成龙, 王育禾, 李晓, authorCompany=null, correspAuthors=李晓, authorNote=

李杨，硕士，主要从事动物遗传育种研究，E-mail：

, correspAuthorsNote=
李晓，主要从事猪肉品质形成的调控机理研究，E-mail：
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李杨，硕士，主要从事动物遗传育种研究，E-mail：

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李杨，硕士，主要从事动物遗传育种研究，E-mail：

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fundingSource=陕西省创新能力支撑计划(2023-CX-TD-57), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1307282800508429137, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, xref=null, ext=[AuthorCompanyExt(id=1307282800516817746, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, companyId=1307282800508429137, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=Shaanxi Provincial Key Laboratory of Animal Genetics，Breeding and Reproduction，Laboratory of Animal Fat Deposition and Muscle Development，College of Animal Science and Technology，Northwest A&F University，Yangling 712100，China), AuthorCompanyExt(id=1307282800525206355, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, companyId=1307282800508429137, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=西北农林科技大学动物科技学院，动物脂肪沉积与肌肉发育实验室，陕西省动物遗传育种与繁殖重点实验室，杨凌 712100)])], figs=[ArticleFig(id=1307282804287497072, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Fig.1, caption=Interference of ZNF697 inhibits the proliferation of pig MuSCs

A. The mRNA expression of ZNF697 in proliferating porcine skeletal muscle satellite cells for different time （0， 12， 24， 36， and 48 h）； B. The transfection efficiency of si-ZNF697； C. Flow cytometry analysis of the proportion of cells in different phases of the cell cycle； D. Quantitative analysis of the percentage of cells in different phases； E. Immunofluorescence staining with EdU （red） and Hoechst （blue）， scale bar： 200 μm； F. The ratio of EdU-positive cells was quantified； G. The relative mRNA expression of Cyclin E， Cyclin D， and p21； H. The proteins expression and quantitative results of Cyclin E and Cyclin D. n=3

, figureFileSmall=CpDdN8C4OLPvxbH2hHw1eQ==, figureFileBig=bVq59S8oN3SEfvfVlZ9dCg==, tableContent=null), ArticleFig(id=1307282804530766705, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=图1, caption=干扰ZNF697抑制猪MuSCs的增殖

A.猪MuSCs增殖不同时间（0、12、24、36、48 h）ZNF697 mRNA相对表达量；B. si-ZNF697的转染效率；C.流式细胞术分析细胞周期不同时期的细胞比例；D.不同时期细胞百分比统计分析；E. EdU（红色）和Hoechst（蓝色）免疫荧光染色，比例尺：200 μm；F.统计EdU阳性细胞比例；G. Cyclin E、Cyclin D、p21 mRNA相对表达量；H. Cyclin E和Cyclin D蛋白相对表达量及统计结果。n=3

, figureFileSmall=CpDdN8C4OLPvxbH2hHw1eQ==, figureFileBig=bVq59S8oN3SEfvfVlZ9dCg==, tableContent=null), ArticleFig(id=1307282804623041394, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Fig.2, caption=Overexpression of ZNF697 promotes the proliferation of pig MuSCs

A. The mRNA expression of ZNF697 were detected by qRT-PCR； B. Immunofluorescence staining with EdU（red）and Hoechst（blue），scale bar：200 μm； C. The ratio of EdU-positive cells was quantified； D. The proteins expression and quantitative results of Cyclin B and p21. n=3

, figureFileSmall=fALDViwrNKJ7kder9x4LTA==, figureFileBig=E6wlPROmTpOircCL/DGhPQ==, tableContent=null), ArticleFig(id=1307282804681761651, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=图2, caption=过表达ZNF697促进猪MuSCs的增殖

A. qRT-PCR检测ZNF697 mRNA相对表达量；B. EdU（红色）和Hoechst （蓝色）免疫荧光染色，比例尺：200 μm；C. 统计EdU阳性细胞比例；D. Cyclin B和p21蛋白相对表达量及统计结果。n=3

, figureFileSmall=fALDViwrNKJ7kder9x4LTA==, figureFileBig=E6wlPROmTpOircCL/DGhPQ==, tableContent=null), ArticleFig(id=1307282804753064820, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Fig.3, caption=Interference of ZNF697 inhibits the differentiation of pig MuSCs

A. The mRNA expression of ZNF697 in differentiating porcine skeletal muscle satellite cells for different time （0，2，4， and 6 days）； B. Differentiation index statistical results， differentiation index is calculated as the percentage of nuclei in MyHC-positive cells；C. Immunofluorescence staining with DAPI （blue） and MyHC （green），scale bar：100 μm； D. The mRNA expression of MyHC，MyoG，and MyoD were detected by qRT-PCR； E. The proteins expression and quantitative results of MyHC，MyoG，and MyoD. n=3

, figureFileSmall=H1XXz0z+RotZPKqH7VKhWg==, figureFileBig=ly8qEtPhqhSiXjPdN7/3PQ==, tableContent=null), ArticleFig(id=1307282804954391413, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=图3, caption=干扰ZNF697抑制猪MuSCs的分化

A. 猪 MuSCs 诱导分化不同时间（0、2、4、6 d）ZNF697 mRNA 相对表达量；B. 分化指数统计结果，MyHC 阳性细胞细胞核比例表示分化指数；C. DAPI（蓝色）和 MyHC（绿色）免疫荧光染色，比例尺：100 μm；D. qRT-PCR 检测 MyHC、MyoG 和 MyoDmRNA 相对表达量；E. MyHC、MyoG 和 MyoD 蛋白相对表达量及统计结果。n=3

, figureFileSmall=H1XXz0z+RotZPKqH7VKhWg==, figureFileBig=ly8qEtPhqhSiXjPdN7/3PQ==, tableContent=null), ArticleFig(id=1307282805021500278, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Fig.4, caption=Overexpression of ZNF697 promotes the differentiation of pig MuSCs

A. Immunofluorescence staining with DAPI （blue） and MyHC （green）， scale bar： 100 μm； B. Differentiation index statistical results， differentiation index is calculated as the percentage of nuclei in MyHC-positive cells； C. The mRNA expression of MyHC，MyoG，and MyoD were detected by qRT-PCR； D. The proteins expression and quantitative results of MyHC and MyoG. n=3

, figureFileSmall=whRKVg9Qnx947SGmuR4t+Q==, figureFileBig=fp7mHn9jIf+NywE9rWRnEw==, tableContent=null), ArticleFig(id=1307282805382210424, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=图4, caption=过表达ZNF697促进猪MuSCs的分化

A. DAPI（蓝色）和MyHC（绿色）免疫荧光染色，比例尺：100 μm；B. 分化指数统计结果，MyHC阳性细胞细胞核比例表示分化指数；C. qRT-PCR检测MyHC、MyoG和MyoD mRNA相对表达量；D. MyHC和MyoG蛋白相对表达量及统计结果。n=3

, figureFileSmall=whRKVg9Qnx947SGmuR4t+Q==, figureFileBig=fp7mHn9jIf+NywE9rWRnEw==, tableContent=null), ArticleFig(id=1307282805461902201, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Table 1, caption=

The primer sequences used for real-time qPCR

, figureFileSmall=null, figureFileBig=null, tableContent=
基因 Gene正向序列（5′→3′） Forward primer反向序列（3′→5′） Reverse primer
ZNF697TGAGTGAGGAAGAAGGGGTTGTCCGGGAAATGCTGTCTGATT
p27TGTCTACGAGTTTGCAGCCATGGTCGGCCTTGCAAATTTCT
Cyclin EAGAAGGAAAGGGATGCGAAGGCCAAGGCTGATTGCCACACT
Cyclin DTACACCGACAACTCCATCCGGAGGGCGGGTTGGAAATGAA
p21ACGTCTCAGGAGGACCATGTAGAAGATCAGCCGGCGTTTG
MyHCTTGAAAAGACGAAGCAGCGACAGGAAGGAGGGCTGGAAGAG
MyoGATGAGACATCCCCCTACTTCTACCAGTCCCCAGCCCCTTATCTTCC
MyoDGGCTGCCCAAGGTGGAAATCTGCGTCTGAGTCACCGCTGTAG
β-actinAGATCAAGATCATCGCGCCTATGCAACTAACAGTCCGCCT
), ArticleFig(id=1307282805822612346, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=表1, caption=

实时定量 PCR 的引物序列

, figureFileSmall=null, figureFileBig=null, tableContent=
基因 Gene正向序列（5′→3′） Forward primer反向序列（3′→5′） Reverse primer
ZNF697TGAGTGAGGAAGAAGGGGTTGTCCGGGAAATGCTGTCTGATT
p27TGTCTACGAGTTTGCAGCCATGGTCGGCCTTGCAAATTTCT
Cyclin EAGAAGGAAAGGGATGCGAAGGCCAAGGCTGATTGCCACACT
Cyclin DTACACCGACAACTCCATCCGGAGGGCGGGTTGGAAATGAA
p21ACGTCTCAGGAGGACCATGTAGAAGATCAGCCGGCGTTTG
MyHCTTGAAAAGACGAAGCAGCGACAGGAAGGAGGGCTGGAAGAG
MyoGATGAGACATCCCCCTACTTCTACCAGTCCCCAGCCCCTTATCTTCC
MyoDGGCTGCCCAAGGTGGAAATCTGCGTCTGAGTCACCGCTGTAG
β-actinAGATCAAGATCATCGCGCCTATGCAACTAACAGTCCGCCT
), ArticleFig(id=1307282807470973819, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=EN, label=Table 2, caption=

Antibodies used in Western blot

, figureFileSmall=null, figureFileBig=null, tableContent=
抗体 Antibody宿主 Host稀释比例 Dilution ratio
MyHCRabbit1∶500
MyoDMouse1∶500
MyoGMouse1∶1 000
Cyclin EMouse1∶1 000
Cyclin BMouse1∶1 000
p21Mouse1∶1 000
ZNF697Rabbit1∶500
β-actinRabbit1∶1 500
Anti-rabbit-HRP/1∶5 000
Anti-mouse-HRP/1∶5 000
), ArticleFig(id=1307282807529694077, tenantId=1146029695717560320, journalId=1306274286668972124, articleId=1307013274315289481, language=CN, label=表2, caption=

蛋白质印迹中使用的抗体

, figureFileSmall=null, figureFileBig=null, tableContent=
抗体 Antibody宿主 Host稀释比例 Dilution ratio
MyHCRabbit1∶500
MyoDMouse1∶500
MyoGMouse1∶1 000
Cyclin EMouse1∶1 000
Cyclin BMouse1∶1 000
p21Mouse1∶1 000
ZNF697Rabbit1∶500
β-actinRabbit1∶1 500
Anti-rabbit-HRP/1∶5 000
Anti-mouse-HRP/1∶5 000
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ZNF697对猪骨骼肌卫星细胞增殖及分化的调控作用
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李杨 , 文成龙 , 王育禾 , 李晓
畜牧兽医学报 | 遗传育种 2026,57(2): 668-676
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畜牧兽医学报 |遗传育种 2026 , 57 (2) : 668 -676
ZNF697对猪骨骼肌卫星细胞增殖及分化的调控作用
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李杨 , 文成龙, 王育禾, 李晓
作者信息
  • 西北农林科技大学动物科技学院，动物脂肪沉积与肌肉发育实验室，陕西省动物遗传育种与繁殖重点实验室，杨凌 712100
通讯作者:
李晓，主要从事猪肉品质形成的调控机理研究，E-mail：
作者简介:

李杨，硕士，主要从事动物遗传育种研究，E-mail：

Regulatory Role of ZNF697 on the Proliferation and Differentiation of Skeletal Muscle Satellite Cells of Pig
Yang LI , Chenglong WEN, Yuhe WANG, Xiao LI
Affiliations
  • Shaanxi Provincial Key Laboratory of Animal Genetics，Breeding and Reproduction，Laboratory of Animal Fat Deposition and Muscle Development，College of Animal Science and Technology，Northwest A&F University，Yangling 712100，China
    • Corresponding Author:
    LI Xiao， E-mail：
出版时间: 2026-02-23 doi: 10.11843/j.issn.0366-6964.2026.02.009
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旨在探究ZNF697对猪骨骼肌卫星细胞（muscle satellite cells， MuSCs）增殖及分化的影响。本试验以猪MuSCs为研究对象，利用流式细胞术、实时荧光定量聚合酶链式反应（qRT-PCR）、蛋白质免疫印迹（Western blot）和免疫荧光染色等方法检测ZNF697对猪MuSCs增殖及分化的影响。获得以下研究结果：1）ZNF697 mRNA的表达量随猪MuSCs增殖而动态变化，干扰ZNF697抑制Cyclin E（P<0.01）、Cyclin D（P<0.05）mRNA表达和Cyclin B（P<0.05）蛋白表达，促进p21（P<0.01）mRNA表达，过表达ZNF697取得相反的结果。2）ZNF697在猪MuSCs成肌分化过程中表达逐渐升高，干扰ZNF697显著降低成肌分化指数（P<0.05），抑制MyoG（P<0.01）mRNA表达、MyHC（P<0.05）、MyoD（P<0.01）mRNA和蛋白表达，过表达ZNF697取得相反的结果。本研究鉴定了猪MuSCs增殖与成肌分化的促进因子——ZNF697，为培育高瘦肉率的优良猪种提供了育种靶点。

猪  /  骨骼肌卫星细胞  /  增殖  /  ZNF697

This experiment aimed to investigate the effects of ZNF697 on the proliferation and differentiation of porcine skeletal muscle satellite cells （MuSCs）. In this experiment， porcine MuSCs were used as the research object. Flow cytometry， Real-time quantitative PCR， Western blot and immunofluorescence staining were employed to detect the effects of ZNF697 on the proliferation and differentiation of porcine MuSCs. The following results were obtained： 1） The expression of ZNF697 mRNA dynamically changes with the proliferation of porcine MuSCs. Interfering with ZNF697 inhibited the mRNA expression of Cyclin E （P<0.01） and Cyclin D （P<0.05） and the protein expression of Cyclin B （P<0.05）， promoted the mRNA expression of p21 （P<0.01）. Overexpression of ZNF697 showed the opposite results. 2） During the myogenic differentiation of porcine MuSCs， the expression of ZNF697 gradually increased. Interfering with ZNF697 significantly reduced the myogenic differentiation index （P<0.05） and inhibited the mRNA expression of MyoG （P<0.01）， the mRNA and protein expression of MyHC （P<0.05） and MyoD （P<0.01）. Overexpression of ZNF697 showed the opposite results. The results show that ZNF697 is a promoter of proliferation and myogenic differentiation in porcine MuSCs， which will provide a breeding target for high-lean-meat pig breeds.

pig  /  skeletal muscle satellite cells  /  proliferation  /  ZNF697
李杨, 文成龙, 王育禾, 李晓. ZNF697对猪骨骼肌卫星细胞增殖及分化的调控作用. 畜牧兽医学报, 2026 , 57 (2) : 668 -676 . DOI: 10.11843/j.issn.0366-6964.2026.02.009
Yang LI, Chenglong WEN, Yuhe WANG, Xiao LI. Regulatory Role of ZNF697 on the Proliferation and Differentiation of Skeletal Muscle Satellite Cells of Pig[J]. Acta Veterinaria et Zootechnica Sinica, 2026 , 57 (2) : 668 -676 . DOI: 10.11843/j.issn.0366-6964.2026.02.009
正文
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我国猪肉产量和消费量居世界第一。猪产肉量与骨骼肌生长发育密切相关。骨骼肌卫星细胞（skeletal muscle satellite cells，MuSCs）是一类存在于肌肉基底膜与肌纤维之间的成体干细胞，呈扁平状，广泛地参与骨骼肌生长、发育和修复等生理过程，Pax7是MuSCs的标志基因［1］。MuSCs通常处于静息状态，在生长、急性损伤和衰老等刺激下被激活，进而增殖和分化，促进骨骼肌生成与肌肉再生［2］。MuSCs增殖及成肌分化受到多种微环境因子的复杂调控［3⁃5］。相关研究报道，Cyclin D、Cyclin E调控G1期到S期的转换，Cyclin B是G2/M期周期蛋白标志物，与细胞周期素依赖性激酶（CDK）抑制因子p21等共同参与调控细胞增殖，因此常作为检测细胞增殖的标志物［6⁃10］。MyoD、MyoG等作为成肌分化标志物表达并促进MuSCs向肌原细胞分化，促进骨骼肌的发育与成熟［11⁃13］。探究MuSCs增殖与分化的分子调控机制对于促进猪肉生产、改善猪肉品质具有重要的理论和实践价值。
锌指蛋白697（zinc finger protein 697，ZNF697）又名Zfp697，位于猪的4号染色体上，属于C2H2型锌指蛋白家族。全表型组关联分析（phenome-wide association analysis，PheWAS）显示，ZNF697与猪的肉质与胴体性状高度关联［14］。本课题组前期对猪原代MuSCs增殖期及分化3 d肌管的RNA-seq分析（相关内容未公开发表）显示，增殖期的MuSCs中ZNF697的表达量显著高于肌管，暗示ZNF697对MuSCs的促增殖作用。通过NCBI序列对比，猪和小鼠的ZNF697氨基酸序列分别含有553和569个氨基酸，序列同源性为84.56%（https：//blast.ncbi.nlm.nih.gov/），表明ZNF697在猪和小鼠上可能具有相似的功能。Correia等［15］运用Mef2c-Cre构建骨骼肌特异性ZNF697敲除小鼠，发现ZNF697缺失导致CTX诱导的肌肉再生受阻，RNA-seq分析显示肌肉再生相关基因表达受到抑制。MuSCs在肌肉再生过程中发挥关键作用，提示该基因可能调控MuSCs增殖和/或成肌分化。
因此，本试验以猪MuSCs为研究对象，通过干扰和过表达探究ZNF697对猪MuSCs增殖及分化的作用，为提高猪肉产量和改善猪肉品质提供新的育种靶点。
1　材料与方法
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1.1　试验动物
3日龄大白猪来自西北农林科技大学三站试验猪场。将大白猪用戊巴比妥钠（Sigma）麻醉，耳源性静脉注射10% KCl（0.5 mL·kg-1）处死。37 ℃温水清洗仔猪体表3遍，之后浸泡于75%酒精中消毒30 min。在超净工作台进行解剖，分离猪背最长肌和比目鱼肌。解剖器械超声清洗30 min，之后在超净工作台照射紫外60 min。取样后迅速放于液氮保存，后续用于检测ZNF697的组织表达情况。
1.2　主要试剂
I型胶原酶（SCR103）购自美国Sigma-Aldrich公司；胰蛋白酶（C0202）和Lipofectamine 2000转染试剂（11668-019）购自美国Thermo Fisher Scientific公司；DMEM/F12高糖培养基（SH30023.01）和胎牛血清（SV30208.02）购自美国Cytiva HyClone公司；青霉素/链霉素（B540734-0010）和马血清（E510006-0100）购自中国生工生物工程（上海）股份有限公司；Trizol试剂（AG21102）购自艾科瑞生物科技有限公司；蛋白裂解液和蛋白酶抑制剂购自美国AbMole BioScience公司；荧光定量（EG20117M）和反转录试剂盒（EG15139M-1-2）购自江苏愚公生物科技有限公司；荧光定量引物由北京擎科生物科技有限公司合成；BeyoClick™ EdU-594细胞增殖检测试剂盒（C0078S）购买自中国上海碧云天生物技术有限公司；本研究中siRNA由广州锐博生物技术有限公司合成；pcDNA3.1-ZNF697载体由通用生物公司合成。
1.3　猪MuSCs的分离及培养
3日龄杜长大仔公猪3头，选用背最长肌，均匀剪碎成1 mm3大小。猪MuSCs的分离方法已有报道［16］，采用双酶法进行肌肉消化，在37°C的水浴中用0.1%的I型胶原酶消化90 min，之后换用0.25%的胰蛋白酶消化30 min。用含10%胎牛血清的DMEM/F12高糖培养基终止消化，之后将细胞悬液依次通过70目和200目筛网过滤。细胞重悬于生长培养基中，生长培养基含DMEM/F12、20%胎牛血清和1%青霉素/链霉素。差速贴壁法纯化骨骼肌卫星细胞：分离得到的细胞悬液先在细胞培养皿中自由沉降1 d，然后上清液转入用基质胶孵育后的培养皿中贴壁2 d，得到F1代骨骼肌卫星细胞。细胞培养于37℃，5% CO2培养箱中，每2 d更换新的培养基。密度达到90%时更换为分化培养基诱导分化5 d，分化培养基含DMEM/F12、5%马血清和1%青霉素/链霉素。
1.4　细胞转染
细胞培养于6孔或12孔细胞培养板，铺满皿底70%~80%时进行转染。根据Lipofectamine 2000转染试剂说明书，将靶向ZNF697的si-ZNF697和pcDNA3.1-ZNF697载体分别转染进细胞，6~12 h后更换培养基。
1.5　RNA提取、反转录及qRT-PCR
Trizol法提取细胞和组织的总RNA。分光光度计测定RNA浓度，取A260/A280值为1.8~2.0的样品使用AGEvo M-MLV反转录试剂盒进行反转录，之后按照SYBR Green Kit试剂盒和基因特异引物进行qRT-PCR反应，反应体系为：cDNA 2 μL，上、下游引物各0.2 μL，Taq SYBR Green qPCR PreMix 5 μL，ddH2O 2.6 μL，总体积为10 μL。引物序列见表1，内参为β-actin。数据采用2-ΔΔCt进行分析。
1.6　蛋白质免疫印迹（Western blot）检测
加入适量的裂解液（RIPA∶蛋白酶抑制剂为100∶1），冰上裂解30 min，4 ℃、12 000 r·min-1离心10 min，保留上清液。按照上清液∶缓冲液为4∶1的比例添加上样缓冲液，煮沸10 min。BCA蛋白检测试剂盒检测蛋白浓度。制备10%SDS-PAGE凝胶，每个上样孔加入20 μg蛋白样品，80 V电泳至样品进入分离后，调电压至120 V，待样品中溴酚蓝迁移至凝胶最下端时，电泳完毕。然后转膜2.5 h到PVDF膜上。封闭液封闭膜2 h。一抗4 ℃摇床过夜孵育，摇床上孵育二抗2 h。曝光显影。本研究中所用抗体见表2。
1.7　流式细胞术检测
将密度为15%~20%的细胞接种于6 cm培养皿，细胞密度达到40%~50%时转染si-ZNF697或pcDNA3.1-ZNF697，待细胞汇合度为80%左右时用胰酶消化。收集细胞沉淀，冷乙醇4 ℃过夜固定，PBS清洗后加入500 µL染色工作液（含50 µg·mL-1碘化丙啶的PBS，100 µg·mL-1 RNase A，0.2% Triton X-100），4 ℃避光孵育30 min。流式细胞仪检测细胞数量，ModFit量化计数结果。
1.8　EdU染色
将对数生长期的细胞接种于6孔或12孔细胞培养板，培养至正常生长阶段。细胞密度达到30%~40%时转染si-ZNF697或pcDNA3.1-ZNF697，并按照BeyoClick™ EdU-594细胞增殖检测试剂盒进行EdU染色，染色结束后在荧光显微镜下观察并计数。
1.9　免疫荧光染色
转染后的细胞诱导分化5 d，弃去培养基，PBS洗涤2次。将细胞用4%多聚甲醛固定10 min，0.5% Triton X-100通透10 min，用5% BSA的封闭液封闭1 h，一抗4 ℃孵育过夜，二抗室温孵育2 h，细胞经PBS洗涤后进行DAPI染色，置于荧光显微镜下观察、拍照。
1.10　数据分析
使用GraphPad Prism 9.5.0进行数据整理和绘图，结果以“平均值±标准误”表示，使用Student’s-t检验或单因素方差分析（one-way analysis of variance，ANOVA）。*P<0.05表示显著，**P<0.01、***P<0.001表示极显著。
2　结果
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2.1　干扰ZNF697对猪MuSCs增殖的影响
在MuSCs增殖过程中，ZNF697表达量在24 h最高（图1A）。转染siRNA，24 h之后检测转染效率，ZNF697 mRNA水平极显著降低（P<0.01，图1B）。对细胞周期的检测发现，干扰ZNF697导致G1期细胞比例显著高于对照组，S期细胞比例则显著低于对照组（P<0.05，图1C、1D），提示干扰ZNF697抑制猪MuSCs增殖。EdU染色结果显示干扰ZNF697显著降低了EdU阳性细胞数目（P<0.05，图1E、1F）。qRT-PCR检测发现干扰ZNF697后，Cyclin E mRNA表达水平极显著降低（P<0.01），Cyclin D mRNA表达水平显著降低（P<0.05），p21 mRNA水平极显著升高（P<0.01，图1G）。Westem blot结果表明，干扰ZNF697极显著降低Cyclin E蛋白水平（P<0.01），显著降低Cyclin B的蛋白水平（P<0.05，图1H），与mRNA结果一致。以上结果表明，干扰ZNF697抑制猪MuSCs增殖。
2.2　过表达ZNF697对猪MuSCs增殖的影响
转染pcDNA3.1-ZNF697显著提高ZNF697 mRNA水平，约15倍以上（P<0.05，图2A）。EdU染色结果显示，与对照组相比，过表达ZNF697 EdU阳性细胞数量显著提高（P<0.05，图2B、2C）。Western blot结果显示，在增殖过程中，ZNF697过表达显著促进Cyclin B表达（P<0.05），极显著抑制p21表达（P<0.01，图2D）。结果表明，过表达ZNF697促进猪MuSCs增殖。
2.3　干扰ZNF697对猪成肌分化的影响
在MuSCs分化过程中，ZNF697表达水平逐渐上升（图3A）。转染si-ZNF697显著降低MyHC阳性细胞比例，分化指数显著降低（P<0.05，图3B、3C）。RT-qPCR结果显示，干扰ZNF697显著降低分化标志基因MyHC mRNA的表达水平（P<0.05），极显著降低MyoG和MyoD mRNA表达水平（P<0.01，图3D）。Western blot结果同样显示，干扰ZNF697 后可显著降低MyHC蛋白水平（P<0.05），极显著降低MyoD蛋白水平（P<0.01，图3E）。以上结果表明，干扰ZNF697 抑制猪MuSCs的分化。
2.4　过表达ZNF697对成肌分化的影响
相比于对照组，过表达ZNF697显著增加了MyHC阳性细胞的比例，分化指数显著升高（P<0.05，图4A、4B）。同时，过表达ZNF697极显著升高成肌标志基因MyHC（P<0.01）、MyoD mRNA表达水平（P<0.001，图4C）。Western blot 结果表明，MyHC（P<0.01）和MyoG蛋白水平极显著升高（P<0.001，图4D），与mRNA结果一致。以上结果表明，过表达ZNF697 促进MuSCs分化。
3　讨论
收起
MuSCs具有干细胞活性，可以增殖、分化为肌细胞，促进肌肉生长和损伤后再生［17］。ZNF697是锌指蛋白家族（ZNFs）的一员，该家族在调节转录、DNA修复、信号转导、蛋白质降解等诸多功能中发挥作用［18］。相关研究发现，ZNFs 如Klf2、Klf3和Klf4等参与调节骨骼肌发育［19］。Hayashi等［20］研究表明，MuSCs缺失Krüppel样因子5（Klf5）抑制肌管形成。Hirata等［21］研究发现，小鼠骨骼肌中缺乏KLF15可以抵御肌肉萎缩。本研究通过检索PigBiobank数据库发现ZNF697与猪肉质性状（腰大肌深度、背膘厚度）显著相关，检索Ensembl数据库发现ZNF697上游SNP位点rs80909317与猪腰大肌深度显著相关（https：//www.ensembl.org/），此外，检索GeneAtlas数据库发现ZNF697上游位点rs49886和rs404937与人的生长性状高度相关（http：//geneatlas.roslin.ed.ac.uk/），然而ZNF697对于猪骨骼肌发育的调控作用仍不清楚。试验结果表明ZNF697促进MuSCs成肌分化［14］。
目前对ZNF697报道较少且集中于炎症反应。Guo等［22］发现ZNF697在促炎肺泡巨噬细胞中上调，并根据该基因表达量构建预测肺腺癌风险预后模型。Martínez-Aguilar等［23］运用GWAS分析发现ZNF697 rs10494227位点A等位基因与干扰素β（IFNβ）治疗反应相关。Correia等［15］发现ZNF697促进趋化因子配体1（Ccl1）、干扰素调节因子1（Irf1）等炎症相关基因的表达，促进骨骼肌的组织修复，过表达ZNF697会导致细胞凋亡以及NF-κB/TNF-α、IL2/STAT5等相关通路的富集。研究发现NF-κB/TNF-α、IL2/STAT5参与炎症反应［24⁃25］。MuSCs在受到炎症因子的刺激下激活并进入细胞周期，实现自我更新，从而维持干细胞池的稳态［26］。ZNF697通过促进炎症反应进而促进MuSCs的激活是未来研究的方向。本研究发现，在MuSCs中干扰ZNF697显著抑制Cyclin B，促进p21 mRNA表达水平，进而促进MuSCs的增殖。此外，IFNγ通路可通过ZNF697上的STAT位点发挥作用，注射IFNγ导致小鼠肌管ZNF697 mRNA水平显著升高［15］。最近的研究发现IFNγ长期刺激阻断MuSCs激活，抑制MuSCs的增殖［27］。而本研究发现过表达ZNF697促进MuSCs增殖，这表明长期的IFNγ刺激可能导致ZNF697水平的进一步变化，进而导致ZNF697的作用失效。对于IFNγ通路与ZNF697之间的具体作用仍需要深入探究。
锌指蛋白家族成员大多数为转录因子，通常与DNA结合发挥特定功能，调节基因的转录等。然而，研究发现ZNF697主要作为RNA结合蛋白发挥作用［15］。相关研究表明C2H2型锌指蛋白也可以结合RNA，参与转录后调控［28］。Bye-A-Jee等［29］发现在成年小鼠Pax7阳性细胞中RNA结合蛋白Zfp36l1与Zfp36l2缺失会导致骨骼肌重量显著降低。ZNF697属于C2H2型锌指蛋白，Correia等［15］发现ZNF697作为RNA结合蛋白靶向miR-206-3p。该miRNA已被证明可以促进骨骼肌再生［30］，表明ZNF697可能通过调节miRNA等调控MuSCs的生理功能。有关ZNF697作为转录因子或RNA结合蛋白的功能仍有待进一步研究。
4　结论
收起
本研究发现，干扰ZNF697显著抑制猪MuSCs增殖及成肌分化，过表达促进该过程。本研究结果为进一步探索ZNF697对MuSCs的具体作用机制提供研究依据，为提高猪肉产量提供新的育种靶点。
基金
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  • 农业生物育种国家科技重大专项(2023ZD0404702)
  • 国家生猪产业技术体系(CARS-35-PIG)
  • 陕西省创新能力支撑计划(2023-CX-TD-57)
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2026年第57卷第2期
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doi: 10.11843/j.issn.0366-6964.2026.02.009
  • 接收时间：2025-06-19
  • 首发时间：2026-09-16
  • 出版时间：2026-02-23
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  • 收稿日期：2025-06-19
基金
农业生物育种国家科技重大专项(2023ZD0404702)
国家生猪产业技术体系(CARS-35-PIG)
陕西省创新能力支撑计划(2023-CX-TD-57)
作者信息
    西北农林科技大学动物科技学院，动物脂肪沉积与肌肉发育实验室，陕西省动物遗传育种与繁殖重点实验室，杨凌 712100

通讯作者:

李晓，主要从事猪肉品质形成的调控机理研究，E-mail：
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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