仿刺参基质金属蛋白酶基因MMP-16的克隆及表达
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摘要
基质金属蛋白酶(MMPs)是一种能够降解细胞外基质的蛋白水解酶类。为研究MMPs在仿刺参免疫防御中的作用,本实验采用RACE技术克隆了仿刺参基质金属蛋白酶16基因(Aj-MMP-16)的cDNA全长序列,并对其序列特征和功能进行了初步分析;采用实时荧光定量PCR(qRT-PCR)方法,分别分析了Aj-MMP-16基因在仿刺参不同组织、不同"化皮"体壁组织以及病原菌刺激后体腔细胞中的表达情况。结果显示,Aj-MMP-16基因的cDNA全长为2 976 bp,包括一个342 bp的5′非编码区,一个963 bp的3′非编码区;开放阅读框(ORF)为1 671 bp,编码557个氨基酸,预测蛋白分子量为63.11 ku,等电点为4.79。Aj-MMP-16具有典型的MMPs家族蛋白结构:N-端前肽区、铰链区、催化区、C-端类血红素结合区和跨膜区。Aj-MMP-16与其他物种的MMPs具有一定的相似性,与紫色球海胆的MMP-16相似性最高。Aj-MMP-16基因mRNA在仿刺参各组织中均有表达,表达量由高到低为呼吸树、肠、体腔细胞、管足、肌肉、体壁;在"化皮病"不同阶段,AjMMP-16基因mRNA在"化皮"体壁组织中的表达量显著高于正常体壁组织;灿烂弧菌和蜡样芽孢杆菌刺激后,体腔细胞中Aj-MMP-16基因mRNA表达量显著升高。Aj-MMP-16基因可能在仿刺参内脏再生、炎症发生以及免疫应答中起着重要的作用。
Matrix metalloproteinases(MMPs) are proteolytic enzymes that degrade extracellular matrix. In order to study the function of MMPs in the immune defense of Apostichopus japonicus, the full-length cDNA sequence of the matrix metalloproteinase 16 gene, named Aj-MMP-16, was cloned using RACE method. The sequence characteristics and function of Aj-MMP-16 were preliminarily analyzed. The results showed that the full-length cDNA of this gene was 2 976 bp, including a 5′ non-coding region of 342 bp, a 3′ non-coding region of 963 bp,and an open reading frame(ORF) of 1 671 bp encoding 557 amino acids. The predicted molecular weight of AjMMP-16 protein was 63.11 ku and isoelectric point was 4.79. Functional domain analysis revealed the typical MMPs family protein structure of Aj-MMP-16 including N-terminal propeptide region, hinge region, catalytic region, hemopexin-like domain and transmembrane region. Multiple sequence alignment and phylogenetic analysis showed that Aj-MMP-16 shared a certain degree of conservatism with MMPs of other species and had the highest identity with MMP-16 of Strongylocentrotus purpuratus. Quantitative real time PCR showed that the Aj-MMP-16 mRNA was expressed in all tissues of A. japonicus, and the expression levels were from high to low in the order of respiratory tree, intestine, coelomocytes, tube feet, muscle and body wall. At different stages of skin ulceration syndrome progression, the expression of Aj-MMP-16 mRNA in ulcerate body wall was significantly higher than that of the normal body wall. After the pathogenic bacteria challenge, Aj-MMP-16 mRNA expression increased significantly in coelomocytes. The results suggested that Aj-MMP-16 may play important roles in visceral regeneration, inflammation and immune response in sea cucumber.
Matrix metalloproteinases(MMPs) are proteolytic enzymes that degrade extracellular matrix. In order to study the function of MMPs in the immune defense of Apostichopus japonicus, the full-length cDNA sequence of the matrix metalloproteinase 16 gene, named Aj-MMP-16, was cloned using RACE method. The sequence characteristics and function of Aj-MMP-16 were preliminarily analyzed. The results showed that the full-length cDNA of this gene was 2 976 bp, including a 5′ non-coding region of 342 bp, a 3′ non-coding region of 963 bp,and an open reading frame(ORF) of 1 671 bp encoding 557 amino acids. The predicted molecular weight of AjMMP-16 protein was 63.11 ku and isoelectric point was 4.79. Functional domain analysis revealed the typical MMPs family protein structure of Aj-MMP-16 including N-terminal propeptide region, hinge region, catalytic region, hemopexin-like domain and transmembrane region. Multiple sequence alignment and phylogenetic analysis showed that Aj-MMP-16 shared a certain degree of conservatism with MMPs of other species and had the highest identity with MMP-16 of Strongylocentrotus purpuratus. Quantitative real time PCR showed that the Aj-MMP-16 mRNA was expressed in all tissues of A. japonicus, and the expression levels were from high to low in the order of respiratory tree, intestine, coelomocytes, tube feet, muscle and body wall. At different stages of skin ulceration syndrome progression, the expression of Aj-MMP-16 mRNA in ulcerate body wall was significantly higher than that of the normal body wall. After the pathogenic bacteria challenge, Aj-MMP-16 mRNA expression increased significantly in coelomocytes. The results suggested that Aj-MMP-16 may play important roles in visceral regeneration, inflammation and immune response in sea cucumber.
引文
[1]Michel G,Tonon T,Scornet D,et al.The cell wall polysaccharide metabolism of the brown alga ectocarpus siliculosus.Insights into the evolution of extracellular matrix polysaccharides in eukaryotes[J].New Phytologist,2010,188(1):82-97.
[2]Paye A,Truong A,Yip C,et al.EGFR activation and signaling in cancer cells are enhanced by the membranebound metalloprotease MT4-MMP[J].Cancer Research,2014,74(23):6758-6770.
[3]Olczyk P,Mencner?,Komosinska-Vassev K.The role of the extracellular matrix components in cutaneous wound healing[J].BioMed Research International,2014,2014:747584.
[4]Abedin M,King N.Diverse evolutionary paths to cell adhesion[J].Trends in Cell Biology,2010,20(12):734-742.
[5]Visse R,Nagase H.Matrix metalloproteinases and tissue inhibitors of metalloproteinases:structure,function,and biochemistry[J].Circulation Research,2003,92(8):827-839.
[6]VanS aun M N,Matrisian L M.Matrix metalloproteinases and cellular motility in development and disease[J].Birth Defects Research-Part C:Embryo Today:Reviews,2006,78(1):69-79.
[7]Massova I,Kotra L P,Fridman R,et al.Matrix metalloproteinases:structures,evolution,and diversification[J].The FASEB Journal,1998,12(12):1075-1095.
[8]Rojkind M.Role of metalloproteinases in liver fibrosis[J].Alcoholism:Clinical&Experimental Research,1999,23(5):934-939.
[9]Belotti D,Paganoni P,Manenti L,et al.Matrix metalloproteinases(MMP9 and MMP2)induce the release of vascular endothelial growth factor(VEGF)by ovarian carcinoma cells[J].Cancer Research,2003,63(17):5224-5229.
[10]Pei D,Majmudar G,Weiss S J.Hydrolytic inactivation of a breast carcinoma cell-derived serpin by human stromelysin-3[J].Journal of Biological Chemistry,1994,269(41):25849-25855.
[11]Freitas V S,de Araújo C R F,Alves P M,et al.Immunohistochemical expression of matrilysins(MMP-7 and MMP-26)in ameloblastomas and adenomatoid odontogenic tumors[J].Oral Surgery,Oral Medicine,Oral Pathology,Oral Radiology,and Endodontology,2009,108(3):417-424.
[12]Puttabyatappa M,Jacot T A,Al-Alem L F,et al.Ovarian membrane-type matrix metalloproteinases:induction of MMP14 and MMP16 during the periovulatory period in the rat,macaque,and human[J].Biology of Reproduction,2014,91(2):34.
[13]Bugel S M,Wehmas L C,La Du J K,et al.Phenotype anchoring in zebrafish reveals a potential role for matrix metalloproteinases(MMPs)in tamoxifen's effects on skin epithelium[J].Toxicology and Applied Pharmacology,2016,296:31-41.
[14]Wang C,Zhan C L,Cai Q F,et al.Expression and characterization of common carp(Cyprinus carpio)matrix metalloproteinase-2 and its activity against type Icollagen[J].Journal of Biotechnology,2014,177:45-52.
[15]Ogiwara K,Takano N,Shinohara M,et al.Gelatinase Aand membrane-type matrix metalloproteinases 1 and 2are responsible for follicle rupture during ovulation in the medaka[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102(24):8442-8447.
[16]Tsukamoto H,Yokoyama Y,Suzuki T,et al.Molecular cloning and expression of gelatinases(MMP-2 and MMP-9)in the pufferfish Takifugu rubripes[J].Comparative Biochemistry and Physiology-Part B:Biochemistry and Molecular Biology,2007,148(3):295-302.
[17]罗少杰,闫芳,郑哲,等.马氏珠母贝基质金属蛋白酶基因MMP-17的克隆及表达分析[J].水产学报,2015,39(7):978-988.Luo S J,Yan F,Zheng Z,et al.Molecular cloning and expression analysis of matrix metalloproteinase 17 gene from Pinctada martensii[J].Journal of Fisheries of China,2015,39(7):978-988(in Chinese).
[18]Wang K J,Ren H L,Xu D D,et al.Identification of the up-regulated expression genes in hemocytes of variously colored abalone(Haliotis diversicolor Reeve,1846)challenged with bacteria[J].Developmental&Comparative Immunology,2008,32(11):1326-1347.
[19]Chovar-Vera O,Valenzuela-Mu?oz V,GallardoEscárate C.Molecular characterization of collagen IVevidences early transcription expression related to the immune response against bacterial infection in the red abalone(Haliotis rufescens)[J].Fish&Shellfish Immunology,2015,42(2):241-248.
[20]Lepage T,Gache C.Purification and characterization of the sea urchin embryo hatching enzyme[J].Journal of Biological Chemistry,1989,264(9):4787-4793.
[21]Ingersoll E P,Pendharkar N C.Characterization and expression of two matrix metalloproteinase genes during sea urchin development[J].Gene Expression Patterns,2005,5(6):727-732.
[22]Qui?ones J L,Rosa R,Ruiz D L,et al.Extracellular matrix remodeling and metalloproteinase involvement during intestine regeneration in the sea cucumber Holothuria glaberrima[J].Developmental Biology,2002,250(1):181-197.
[23]Wu H L,Hu Y Q,Shen J D,et al.Identification of a novel gelatinolytic metalloproteinase(GMP)in the body wall of sea cucumber(Stichopus japonicus)and its involvement in collagen degradation[J].Process Biochemistry,2013,48(5-6):871-877.
[24]Yang A F,Zhou Z C,Pan Y J,et al.RNA sequencing analysis to capture the transcriptome landscape during skin ulceration syndrome progression in sea cucumber Apostichopus japonicus[J].BMC Genomics,2016,17:459.
[25]汪笑宇,周遵春,关晓燕,等.仿刺参及养殖环境中溶藻弧菌和灿烂弧菌的PCR快速检测[J].中国农业科技导报,2010,12(3):125-130.Wang X Y,Zhou Z C,Guan X Y,et al.Rapid PCRdetection for Vibrio alginolyticus and Vibrio splendidus in sea cucumber Apostichopus japonicus and its culturing environment[J].Journal of Agricultural Science and Technology,2010,12(3):125-130(in Chinese).
[26]艾海新,于晶晶,郑方亮,等.刺参“腐皮综合症”致病菌LNUB415的分离及防治的初步研究[J].微生物学杂志,2012,32(2):68-72.Ai H X,Yu J J,Zheng F L,et al.Isolation and control of pathogen bacteria LNUB415 that caused“Skin Ulcer Syndrome”on spinous sea cucumber(Apostichopus japonicus)[J].Journal of Microbiology,2012,32(2):68-72(in Chinese).
[27]杨爱馥,周遵春,董颖,等.仿刺参cytb和β-actin基因表达稳定性比较[J].中国农业科技导报,2010,12(1):79-84.Yang A F,Zhou Z C,Dong Y,et al.Stability comparison of cytb andβ-actin genes expression in sea cucumber(Apostichopus japonicus)[J].Journal of Agricultural Science and Technology,2010,12(1):79-84(in Chinese).
[28]Livak K J,Schmittgen T D.Analysis of relative gene expression data using real-time quantitative PCR and the2-ΔΔCT method[J].Methods,2001,25(4):402-408.
[29]Van Wart H E,Birkedal-Hansen H.The cysteine switch:a principle of regulation of metalloproteinase activity with potential applicability to the entire matrix metalloproteinase gene family[J].Proceedings of the National Academy of Sciences of the United States of America,1990,87(14):5578-5582.
[30]Jiang W P,Bond J S.Families of metalloendopeptidases and their relationships[J].FEBS Letters,1992,312(2-3):110-114.
[31]Nagase H,Visse R,Murphy G.Structure and function of matrix metalloproteinases and TIMPs[J].Cardiovascular Research,2006,69(3):562-573.
[32]Overall C M.Molecular determinants of metalloproteinase substrate specificity:matrix metalloproteinase substrate binding domains,modules,and exosites[J].Molecular Biotechnology,2002,22(1):51-86.
[33]Roy R,Yang J,Moses M A.Matrix metalloproteinases as novel biomarker s and potential therapeutic targets in human cancer[J].Journal of Clinical Oncology,2009,27(31):5287-5297.
[34]Hatfield K J,Reikvam H,Bruserud?.The crosstalk between the matrix metalloprotease system and the chemokine network in acute myeloid leukemia[J].Current Medicinal Chemistry,2010,17(36):4448-4461.
[35]Miao T,Wan Z X,Sun L N,et al.Extracellular matrix remodeling and matrix metalloproteinases(ajMMP-2like and ajMMP-16 like)characterization during intestine regeneration of sea cucumber Apostichopus japonicus[J].Comparative Biochemistry and Physiology-Part B:Biochemistry and Molecular Biology,2017,212:12-23.
[36]杨红生,周毅,张涛.刺参生物学--理论与实践[M].北京:科学出版社,2014.Yang H S,Zhou Y,Zhang T.The Biology of Sea Cucumber Apostichopus japonicus:theory and Practice[M].Beijing:Science Press,2014(in Chinese).
[37]Mott J D,Werb Z.Regulation of matrix biology by matrix metalloproteinases[J].Current Opinion in Cell Biology,2004,16(5):558-564.
[38]Majkowska I,Shitomi Y,Ito N,et al.Discoidin domain receptor 2 mediates collagen-induced activation of membrane-type 1 matrix metalloproteinase in human fibroblasts[J].Journal of Biological Chemistry,2017,292(16):6633-6643.
[39]Serra R,Grande R,Buffone G,et al.Extracellular matrix assessment of infected chronic venous leg ulcers:role of metalloproteinases and inflammatory cytokines[J].International Wound Journal,2016,13(1):53-58.
[40]Amato B,Coretti G,Compagna R,et al.Role of matrix metalloproteinases in non‐healing venous ulcers[J].International Wound Journal,2015,12(6):641-645.
[41]Latifa K,Sondess S,Hajer G,et al.Evaluation of physiological risk factors,oxidant-antioxidant imbalance,proteolytic and genetic variations of matrix metalloproteinase-9 in patients with pressure ulcer[J].Scientific Reports,2016,6:29371.
[42]Ke F,Wang Y,Hong J,et al.Characterization of MMP-9 gene from a normalized cDNA library of kidney tissue of yellow catfish(Pelteobagrus fulvidraco)[J].Fish&Shellfish Immunology,2015,45(2):260-267.
[43]Oh S Y,Lee S J,Jung Y H,et al.Arachidonic acid promotes skin wound healing through induction of human MSC migration by MT3-MMP-mediated fibronectin degradation[J].Cell Death&Disease,2015,6(5):e1750.
[44]Tracy L E,Minasian R A,Caterson E J.Extracellular matrix and dermal fibroblast function in the healing wound[J].Advances in Wound Care,2016,5(3):119-136.
[2]Paye A,Truong A,Yip C,et al.EGFR activation and signaling in cancer cells are enhanced by the membranebound metalloprotease MT4-MMP[J].Cancer Research,2014,74(23):6758-6770.
[3]Olczyk P,Mencner?,Komosinska-Vassev K.The role of the extracellular matrix components in cutaneous wound healing[J].BioMed Research International,2014,2014:747584.
[4]Abedin M,King N.Diverse evolutionary paths to cell adhesion[J].Trends in Cell Biology,2010,20(12):734-742.
[5]Visse R,Nagase H.Matrix metalloproteinases and tissue inhibitors of metalloproteinases:structure,function,and biochemistry[J].Circulation Research,2003,92(8):827-839.
[6]VanS aun M N,Matrisian L M.Matrix metalloproteinases and cellular motility in development and disease[J].Birth Defects Research-Part C:Embryo Today:Reviews,2006,78(1):69-79.
[7]Massova I,Kotra L P,Fridman R,et al.Matrix metalloproteinases:structures,evolution,and diversification[J].The FASEB Journal,1998,12(12):1075-1095.
[8]Rojkind M.Role of metalloproteinases in liver fibrosis[J].Alcoholism:Clinical&Experimental Research,1999,23(5):934-939.
[9]Belotti D,Paganoni P,Manenti L,et al.Matrix metalloproteinases(MMP9 and MMP2)induce the release of vascular endothelial growth factor(VEGF)by ovarian carcinoma cells[J].Cancer Research,2003,63(17):5224-5229.
[10]Pei D,Majmudar G,Weiss S J.Hydrolytic inactivation of a breast carcinoma cell-derived serpin by human stromelysin-3[J].Journal of Biological Chemistry,1994,269(41):25849-25855.
[11]Freitas V S,de Araújo C R F,Alves P M,et al.Immunohistochemical expression of matrilysins(MMP-7 and MMP-26)in ameloblastomas and adenomatoid odontogenic tumors[J].Oral Surgery,Oral Medicine,Oral Pathology,Oral Radiology,and Endodontology,2009,108(3):417-424.
[12]Puttabyatappa M,Jacot T A,Al-Alem L F,et al.Ovarian membrane-type matrix metalloproteinases:induction of MMP14 and MMP16 during the periovulatory period in the rat,macaque,and human[J].Biology of Reproduction,2014,91(2):34.
[13]Bugel S M,Wehmas L C,La Du J K,et al.Phenotype anchoring in zebrafish reveals a potential role for matrix metalloproteinases(MMPs)in tamoxifen's effects on skin epithelium[J].Toxicology and Applied Pharmacology,2016,296:31-41.
[14]Wang C,Zhan C L,Cai Q F,et al.Expression and characterization of common carp(Cyprinus carpio)matrix metalloproteinase-2 and its activity against type Icollagen[J].Journal of Biotechnology,2014,177:45-52.
[15]Ogiwara K,Takano N,Shinohara M,et al.Gelatinase Aand membrane-type matrix metalloproteinases 1 and 2are responsible for follicle rupture during ovulation in the medaka[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102(24):8442-8447.
[16]Tsukamoto H,Yokoyama Y,Suzuki T,et al.Molecular cloning and expression of gelatinases(MMP-2 and MMP-9)in the pufferfish Takifugu rubripes[J].Comparative Biochemistry and Physiology-Part B:Biochemistry and Molecular Biology,2007,148(3):295-302.
[17]罗少杰,闫芳,郑哲,等.马氏珠母贝基质金属蛋白酶基因MMP-17的克隆及表达分析[J].水产学报,2015,39(7):978-988.Luo S J,Yan F,Zheng Z,et al.Molecular cloning and expression analysis of matrix metalloproteinase 17 gene from Pinctada martensii[J].Journal of Fisheries of China,2015,39(7):978-988(in Chinese).
[18]Wang K J,Ren H L,Xu D D,et al.Identification of the up-regulated expression genes in hemocytes of variously colored abalone(Haliotis diversicolor Reeve,1846)challenged with bacteria[J].Developmental&Comparative Immunology,2008,32(11):1326-1347.
[19]Chovar-Vera O,Valenzuela-Mu?oz V,GallardoEscárate C.Molecular characterization of collagen IVevidences early transcription expression related to the immune response against bacterial infection in the red abalone(Haliotis rufescens)[J].Fish&Shellfish Immunology,2015,42(2):241-248.
[20]Lepage T,Gache C.Purification and characterization of the sea urchin embryo hatching enzyme[J].Journal of Biological Chemistry,1989,264(9):4787-4793.
[21]Ingersoll E P,Pendharkar N C.Characterization and expression of two matrix metalloproteinase genes during sea urchin development[J].Gene Expression Patterns,2005,5(6):727-732.
[22]Qui?ones J L,Rosa R,Ruiz D L,et al.Extracellular matrix remodeling and metalloproteinase involvement during intestine regeneration in the sea cucumber Holothuria glaberrima[J].Developmental Biology,2002,250(1):181-197.
[23]Wu H L,Hu Y Q,Shen J D,et al.Identification of a novel gelatinolytic metalloproteinase(GMP)in the body wall of sea cucumber(Stichopus japonicus)and its involvement in collagen degradation[J].Process Biochemistry,2013,48(5-6):871-877.
[24]Yang A F,Zhou Z C,Pan Y J,et al.RNA sequencing analysis to capture the transcriptome landscape during skin ulceration syndrome progression in sea cucumber Apostichopus japonicus[J].BMC Genomics,2016,17:459.
[25]汪笑宇,周遵春,关晓燕,等.仿刺参及养殖环境中溶藻弧菌和灿烂弧菌的PCR快速检测[J].中国农业科技导报,2010,12(3):125-130.Wang X Y,Zhou Z C,Guan X Y,et al.Rapid PCRdetection for Vibrio alginolyticus and Vibrio splendidus in sea cucumber Apostichopus japonicus and its culturing environment[J].Journal of Agricultural Science and Technology,2010,12(3):125-130(in Chinese).
[26]艾海新,于晶晶,郑方亮,等.刺参“腐皮综合症”致病菌LNUB415的分离及防治的初步研究[J].微生物学杂志,2012,32(2):68-72.Ai H X,Yu J J,Zheng F L,et al.Isolation and control of pathogen bacteria LNUB415 that caused“Skin Ulcer Syndrome”on spinous sea cucumber(Apostichopus japonicus)[J].Journal of Microbiology,2012,32(2):68-72(in Chinese).
[27]杨爱馥,周遵春,董颖,等.仿刺参cytb和β-actin基因表达稳定性比较[J].中国农业科技导报,2010,12(1):79-84.Yang A F,Zhou Z C,Dong Y,et al.Stability comparison of cytb andβ-actin genes expression in sea cucumber(Apostichopus japonicus)[J].Journal of Agricultural Science and Technology,2010,12(1):79-84(in Chinese).
[28]Livak K J,Schmittgen T D.Analysis of relative gene expression data using real-time quantitative PCR and the2-ΔΔCT method[J].Methods,2001,25(4):402-408.
[29]Van Wart H E,Birkedal-Hansen H.The cysteine switch:a principle of regulation of metalloproteinase activity with potential applicability to the entire matrix metalloproteinase gene family[J].Proceedings of the National Academy of Sciences of the United States of America,1990,87(14):5578-5582.
[30]Jiang W P,Bond J S.Families of metalloendopeptidases and their relationships[J].FEBS Letters,1992,312(2-3):110-114.
[31]Nagase H,Visse R,Murphy G.Structure and function of matrix metalloproteinases and TIMPs[J].Cardiovascular Research,2006,69(3):562-573.
[32]Overall C M.Molecular determinants of metalloproteinase substrate specificity:matrix metalloproteinase substrate binding domains,modules,and exosites[J].Molecular Biotechnology,2002,22(1):51-86.
[33]Roy R,Yang J,Moses M A.Matrix metalloproteinases as novel biomarker s and potential therapeutic targets in human cancer[J].Journal of Clinical Oncology,2009,27(31):5287-5297.
[34]Hatfield K J,Reikvam H,Bruserud?.The crosstalk between the matrix metalloprotease system and the chemokine network in acute myeloid leukemia[J].Current Medicinal Chemistry,2010,17(36):4448-4461.
[35]Miao T,Wan Z X,Sun L N,et al.Extracellular matrix remodeling and matrix metalloproteinases(ajMMP-2like and ajMMP-16 like)characterization during intestine regeneration of sea cucumber Apostichopus japonicus[J].Comparative Biochemistry and Physiology-Part B:Biochemistry and Molecular Biology,2017,212:12-23.
[36]杨红生,周毅,张涛.刺参生物学--理论与实践[M].北京:科学出版社,2014.Yang H S,Zhou Y,Zhang T.The Biology of Sea Cucumber Apostichopus japonicus:theory and Practice[M].Beijing:Science Press,2014(in Chinese).
[37]Mott J D,Werb Z.Regulation of matrix biology by matrix metalloproteinases[J].Current Opinion in Cell Biology,2004,16(5):558-564.
[38]Majkowska I,Shitomi Y,Ito N,et al.Discoidin domain receptor 2 mediates collagen-induced activation of membrane-type 1 matrix metalloproteinase in human fibroblasts[J].Journal of Biological Chemistry,2017,292(16):6633-6643.
[39]Serra R,Grande R,Buffone G,et al.Extracellular matrix assessment of infected chronic venous leg ulcers:role of metalloproteinases and inflammatory cytokines[J].International Wound Journal,2016,13(1):53-58.
[40]Amato B,Coretti G,Compagna R,et al.Role of matrix metalloproteinases in non‐healing venous ulcers[J].International Wound Journal,2015,12(6):641-645.
[41]Latifa K,Sondess S,Hajer G,et al.Evaluation of physiological risk factors,oxidant-antioxidant imbalance,proteolytic and genetic variations of matrix metalloproteinase-9 in patients with pressure ulcer[J].Scientific Reports,2016,6:29371.
[42]Ke F,Wang Y,Hong J,et al.Characterization of MMP-9 gene from a normalized cDNA library of kidney tissue of yellow catfish(Pelteobagrus fulvidraco)[J].Fish&Shellfish Immunology,2015,45(2):260-267.
[43]Oh S Y,Lee S J,Jung Y H,et al.Arachidonic acid promotes skin wound healing through induction of human MSC migration by MT3-MMP-mediated fibronectin degradation[J].Cell Death&Disease,2015,6(5):e1750.
[44]Tracy L E,Minasian R A,Caterson E J.Extracellular matrix and dermal fibroblast function in the healing wound[J].Advances in Wound Care,2016,5(3):119-136.