以鹿茸为模型探索软骨组织发生
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- 英文论文题名：Exploration of Cartilage Development Using Deer Antler Model
- 论文作者：路晓 ; 孙红梅 ; 张伟 ; 李春义
- 英文论文作者：Lu Xiao ; Sun Hongmei ; Zhang Wei ; Li Chunyi ; State Key Laboratory of Special Economic Animal Molecular Biology ; Institute of Special Animal and Plant Sciences of CAAS
- 年：2016
- 作者机构：中国农业科学院特产研究所特种经济动物分子生物学国家重点实验室;
- 论文关键词：鹿茸 ; 软骨发生 ; 调控机制
- 英文论文关键词：antler ; cartilage generation ; regulation mechanism
- 会议召开时间：2016-06-01
- 会议录名称：“十二五” 鹿科学研究进展
- 语种：中文
- 分类号：Q136
- 学会代码：ZXMY
- 学会名称：中国畜牧业协会
- 页数：10
- 文件大小：782k
- 原文格式：O
摘要
软骨组织的研究一直依赖于哺乳动物胚胎模型,这种方法存在诸如取材困难、难以明确界定分层边界等弊端。而鹿茸非常独特,可以作为一种新的生物医学模型对软骨发育进行研究。本文将传统的软骨研究模型和鹿茸这种新型的模型进行了对比,分析了鹿茸作为软骨研究模型的优势,同时对鹿茸软骨组织发生与生长的组织基础及分子调控机制的研究进展进行了综述,并对调控鹿茸软骨组织中血管生成的调控因子进行了探讨,以期为寻找更有效的软骨发育研究模型提供理论参考。
Studies on cartilage tissues has been relying on embryo model of mammals.This method has difficulties in obtaining material and clearly defining layered boundaries.While,the antler is unique that can be used as a new biomedicine model to study cartilage development.Through comparing the traditional cartilage research model with the new antler model,this paper analyzed the advantages of antlers being a new cartilage research model,reviewed the turning up of antler cartilage and tissue foundation of its growth,and molecular regulation mechanism,discussed the regulatory factors in angiogenesis of antler cartilage,so as to provide theoretical basis for finding a more effective model in studying cartilage development.
Studies on cartilage tissues has been relying on embryo model of mammals.This method has difficulties in obtaining material and clearly defining layered boundaries.While,the antler is unique that can be used as a new biomedicine model to study cartilage development.Through comparing the traditional cartilage research model with the new antler model,this paper analyzed the advantages of antlers being a new cartilage research model,reviewed the turning up of antler cartilage and tissue foundation of its growth,and molecular regulation mechanism,discussed the regulatory factors in angiogenesis of antler cartilage,so as to provide theoretical basis for finding a more effective model in studying cartilage development.
引文
[1]Shi Jianbo,Jiang Xun,Di Jingfang,et al..Effect of basic fibroblast growth factor and insulin on the proliferation and differentiation of mouse chondrocyte[J].CJTER,2005,9(10):234-236.
[2]Newman AP.Articular cartilage repair[J].Am J Sports Med,1998,26(2):309-324.
[3]Khan I M,Gilbert S J,Singhrao S K,et al..Cartilage integration;evaluation of the reasons for failure of integration during cartilage repair.A review[J].Eur Cell Mater,2008,16:26-39.
[4]Shapiro F,Koide S,Glimcher M J.Cell origin and differentiation in the repair of full-thickness defects of articular cartilage[J].J Bone Joint Surg Am,1993,75(4):532-553.
[5]Frenkel S R,Di Cesare P E.Scaffolds for articular cartilage repair[J].Ann Biomed Eng,2004,32(1):26-34.
[6]Ahmed T A,Hincke M T.Strategies for articular cartilage lesion repair and functional restoration[J].Tissue Eng Part B Rev.,2010,16(3):305-329.
[7]Goss R J.Future directions in antler research[J].Anat Rec,1995,241(3):291-302.
[8]Li C,Suttie J M,Clark D E.Histological examination of antler regeneration in red deer(Cervuselaphus)[J].Anat Rec,2005,282(2):163-174.
[9]Banks W J.Histological and ultrastructural aspects of cervine antler development[J].Anat Rec,1973,175(2):175-487.
[10]Frasier M B,Banks W J,Newbrey J W.Characterization of developing antler cartilage matrix.I.Selected histochemical and enzymatic assessment[J].Calcif Tissue Res,1975,17(4):273-88.
[11]Price J S,Oyajobi B O,Nalin AM,et al..Chondrogenesis in the regenerating antler tip in red deer;expression of collagen types I,IIA,IIB,and X demonstrated by in situ nucleic acid hybridization and immunocytochemistry[J].Dev Dyn,1996,205(3):332-347.
[12]孙红梅,杨福合,邢秀梅,等.鹿茸再生干细胞成骨诱导(微粒体)培养体系的建立[J].吉林农业大学学报,2010,32(6):680-683.Sun H M,Yang F H,Xing X M,et al..Establishment of Micromass Culture System for Antler Stem Cell Induction Toward Osteoblast Differentiation in Vitros[J].Journal of Jilin Agricultural University,2010,32(6):680-683.
[13]Li C,Littlejohn R P,Carson ID,et al..Effects of testosterone on pedicle formation and its transformation to antler in castrated male,freemartin and normal female red deer(Cervuselaphus)[J].Gen Comp Endocrinol,2003,131(1):21-31.
[14]Price J,Faucheux C,Allen S.Deer antlers as a model of mammalian regeneration[J].Curr Top Dev Biol,2005,67:1-48.
[15]Nieto-Diaz M,Pita-Thomas D W,Munoz-Galdeano T,et al..Deer antler innervation and regeneration[J].Frontiers in bioscience:a journal and virtual library,2012,17;1 389-1 401.
[16]Li C.Clark D E,Lord E A,ea al.Sampling technique to discriminate the different tissue layers of growing antler tips for gene discovery[J].Anat Rec,2002,268(2):125-130.
[17]Kierdorf U,Li C,Price J S.Improbable appendages:Deer antler renewal as a unique case of mammalian regeneration[J].Semin Cell Dev Biol,2009,20(5),535-542.
[18]Li C,Yang F,Sheppard A.Adult stem cells and mammalian epimorphic regeneration-insights from studying annual renewal of deer antlers[J].Curr Stem Cell Res Ther,2009,4(3),237-251.
[19]Li C.Histogenetic aspects of deer antler development[J].Front Biosci(Elite Ed),2013,5:479-489.
[20]Li C,Suttie J M,Clark D E.Morphological observation of antler regeneration in red deer(Cervuselaphus)[J].JMorphol,2004,262(3):731-740.
[21]Banks W J,Newbrey J W.Antler development as aunique modification of mammalian endochondral ossification[A].Antler development in Cervidae,Caesar Kleberg Wildlife Research Institute Kingsville,Texas,1983,1:279-306.
[22]Yang Z H,Gu L J,Zhang D L,et al..Red deer antler extract accelerates hair growth by stimulating expression of insulin-like growth factor I in full-thickness wound healing rat model[J].Asian-Australasian journal of animal sciences,2012,25(5):708-716.
[23]李沐.MicroRNA介导的IGF-1基因沉默与端粒酶对鹿茸细胞增殖抑制的影响[D].长春:吉林农业大学,硕士论文,2013.Li M.Study on IGF-1 Gene Silence Mediated by microRNA and the Effect of Telomerase Against Proliferation of Antler Cells[D].Changchun;Jilin Agricultural University,Master Dissertation,2013.
[24]王守堂.PTHrP及其受体在梅花鹿茸角中的表达与调节[D].长春:吉林大学,硕士论文.2013.Wang S T.Expression and Regulation of PTHrP and Its Receptor in Sika Deer Antler[D].Changchun;Jilin University,Master Dissertation,2013.
[25]Barling P M,Lai A K,Nicholson L F.Distribution of EGF and its receptor in growing red deer antler[J].CellBiolInt,2005,29(3):229-236.
[26]Wegner M.From head to toes;the multiple facets of Sox proteins[J].Nucleic Acids Res,1999,27(6):1 409-1 420.
[27]江玲霞.猪Sox9基因cDNA的全长克隆及功能预测[D].南京:南京农业大学,硕士学位论文.2008.Jiang L X.Cloning and Function Prediction of the Full-Length cDNA of Porcine Sox9 Gene[D].Nanjing;Nanjing Agricultural UniVersity,Master Dissertation,2008.
[28]赵辰,黄伟,梁熙,等.Smad7在Sox9增强BMP2成软骨效应中的作用[J].第三军医大学学报,2015,37(2):95-100.Zhao C,Huang W,Liang X,et al..Role of Smad7 in Sox9-potentiated and BMP2-induced differentiation of mouse mesenchymal stem cells into chondrocytes[J].J Third Mil Med Univ,2015,37(2):95-100.
[29]Mount J G,Muzylak M,Allen S,et al..Evidence that the canonical Wntsignalling pathway regulates deer antler regeneration[J].Dev Dyn,2006,235(5):1 390-1 399.
[30]Zhang Y,Marsboom G,Toth P T,et al..Mitochondrial respiration regulates adipogenic differentiation of human mesenchymal stem cells[J].PLoS One,2013,8(10):1-12.
[31]Huang P I,Chen Y C,Chen L H,et al..PGC-1 a mediates differentiation of mesenchymal stem cells to brown adipose cells[J].J AtherosclerThromb,2011,18(11):966-980.
[32]Gersbach C A,Byers B A,Pavlath G K,et al..Runx2/Cbfal stimulates transdifferentiation of primary skeletal myoblasts into a mineralizing osteoblastic phenotype[J].Exp Cell Res,2004,300(2):406-417.
[33]Sun H,Yang F,Chu W,et al..Lentiviral-mediated rnai knockdown of cbfa1 gene inhibits endochondral ossification of antler stem cells in micromassculture[J].PloS one,2012,7(10):e47367.
[34]李婷.李沐.孟星宇.等.microRNA介导的IGF1基因沉默对鹿茸软骨细胞增殖的影响[J].西北农林科技大学学报,2013,41(11):7-12.Li T,Li M,Meng X Y,et al..Effect of microRNA-mediated IGF1 gene silencing on the proliferation of deer antler cartilage cells[J].Journal of Northwest A&F University,2013,41(11):7-12.
[35]Huarte M,Sanz-Ezquerro J J,Roncal F,et al..PA subunit from influenza virus polymerase complex interacts with a cellular protein with homology to a family of transcriptional activators[J].J Virol,2001,75(18):8 597-8 604.
[36]Guo J,Wang W,Liao P,et al..Identification of serum biomarkers forpancreaticadenocarcinoma by proteomic analysis[J].Cancer Sci,2009,100(12):2292-2301.
[37]张登禄,韩金祥,崔亚洲,等.胰腺癌转移相关基因C140rf166的真核表达及其蛋白相互作用的蛋白质组学筛选[J].中国医药生物技术,2010,5(3):189-192.Zhang D L,Han J X,Cui Y Z,et al..The eukaryotic expression of pancreatic cancer relted gene C14orfl66 and screening of its interacting proteins by Proteomics methods[J].Chin Med Biotechnol,2010,5(3):189-192.
[38]张登禄,韩金祥,崔亚洲,等.C140rf166异位表达对Hela细胞增殖与迁移能力的影响[J].中国肿瘤防治杂志,2010,17(10):740-743.Zhang D L,Han J X,Cui Y Z,et al..Ectopic expression of C14orf166 and its effect on Hela cell proliferation and immigration[J].CHIN J CANCER PREV TREAT,2010,17(10):740-743.
[39]郭静会.胰腺癌血清蛋白质指纹图谱及CCR7与胰腺癌淋巴结转移相关性研究[D].上海:复旦大学.2010.Guo J H.The study of Protein Profiling of Pancreatic cancer and correlation of CCR7 with lymph node metastasis of Pancreatic caneer[D].Shanghai:Fudan University,Doctoral Dissertation,2010.
[40]Howng S L,Hsu H C,Cheng T S,et al..A novel ninein-interaction protein,CGI-99,blocks ninein phosphorylation by GSK3βand is highly expressed in brain tumors[J].FEBS letters,2004,566(1):162-168.
[41]王静.脑发育不同阶段蛋白质组学及血清脑红蛋白含量的研究[D].北京:中国人民解放军军医进修学院,2006.Wang J.Brain Porteomie Study in Dieffrent Developmental Stages And The Study On Expression Level of Neuorglobin in Serum[D].Beijing;Chinese PLA Postgraduate Medical School,Master Dissertation,2006.
[42]Perez-Gonzalez A,Pazo A,Navajas R,et al..hCLE/C14orf'166 associates with DDXl-HSP C117-FAM98B in a novel transcription-dependent shuttling RNA-transporting complex[J].PloS one,2014,9(3):e90957.
[43]Ariel Rodriguez,Alicia Pe'rez-Gonza'lez,Amelia Nieto.Cellular Human CLE/C14orfl66 Protein Interacts with Influenza Virus Polymerase and Is Required for Viral Replication[J].Journal of virology,2011,85(22):12 062-12 066.
[44]宋家升.H5N1亚型禽流感病毒对家鸭致病力分子机制的研究[D].北京:中国农业科学院.2010.Song J S.Detecting virulence determinants of H5N1 avain influenza virus for ducks[D].Beijing;Chinese Academy of Agricultural Sciences,Doctoral Dissertation,2010.
[45]Li C,Waldrup K A,Corson ID,et al..Histogenesis of antlerogenie tissues cultivated in diffusion chambers in vivo in red deer(Cervuselaphus)[J].J Exp Zool,1995,272(5):345-355.
[46]Li C,Harris A J,Suttie J M.Tissue interactions and antlerogenesis;new findings revealed by a xenograft approach[J].J Exp Zool,2001,290(1):18-30.
[47]Li C,Suttie J M.Deer antlerogenie periosteum;a piece of postnatally retained embryonic tissue[J].AnatEmbryol,2001,204(5):375-388.
[48]Clark D E,Lord E A,Suttie J M.Expression of VEGF and pleiotrophin in deer antler[J].Anat Rec A Discov Mol Cell Evol Biol,2006,288(12):1 281-1 293.
[49]Li C,Harper A,Puddick J,et al..Proteomes and signalling pathways of antler stem cells[J].PLoS One,2012,7(1):e30026.
[2]Newman AP.Articular cartilage repair[J].Am J Sports Med,1998,26(2):309-324.
[3]Khan I M,Gilbert S J,Singhrao S K,et al..Cartilage integration;evaluation of the reasons for failure of integration during cartilage repair.A review[J].Eur Cell Mater,2008,16:26-39.
[4]Shapiro F,Koide S,Glimcher M J.Cell origin and differentiation in the repair of full-thickness defects of articular cartilage[J].J Bone Joint Surg Am,1993,75(4):532-553.
[5]Frenkel S R,Di Cesare P E.Scaffolds for articular cartilage repair[J].Ann Biomed Eng,2004,32(1):26-34.
[6]Ahmed T A,Hincke M T.Strategies for articular cartilage lesion repair and functional restoration[J].Tissue Eng Part B Rev.,2010,16(3):305-329.
[7]Goss R J.Future directions in antler research[J].Anat Rec,1995,241(3):291-302.
[8]Li C,Suttie J M,Clark D E.Histological examination of antler regeneration in red deer(Cervuselaphus)[J].Anat Rec,2005,282(2):163-174.
[9]Banks W J.Histological and ultrastructural aspects of cervine antler development[J].Anat Rec,1973,175(2):175-487.
[10]Frasier M B,Banks W J,Newbrey J W.Characterization of developing antler cartilage matrix.I.Selected histochemical and enzymatic assessment[J].Calcif Tissue Res,1975,17(4):273-88.
[11]Price J S,Oyajobi B O,Nalin AM,et al..Chondrogenesis in the regenerating antler tip in red deer;expression of collagen types I,IIA,IIB,and X demonstrated by in situ nucleic acid hybridization and immunocytochemistry[J].Dev Dyn,1996,205(3):332-347.
[12]孙红梅,杨福合,邢秀梅,等.鹿茸再生干细胞成骨诱导(微粒体)培养体系的建立[J].吉林农业大学学报,2010,32(6):680-683.Sun H M,Yang F H,Xing X M,et al..Establishment of Micromass Culture System for Antler Stem Cell Induction Toward Osteoblast Differentiation in Vitros[J].Journal of Jilin Agricultural University,2010,32(6):680-683.
[13]Li C,Littlejohn R P,Carson ID,et al..Effects of testosterone on pedicle formation and its transformation to antler in castrated male,freemartin and normal female red deer(Cervuselaphus)[J].Gen Comp Endocrinol,2003,131(1):21-31.
[14]Price J,Faucheux C,Allen S.Deer antlers as a model of mammalian regeneration[J].Curr Top Dev Biol,2005,67:1-48.
[15]Nieto-Diaz M,Pita-Thomas D W,Munoz-Galdeano T,et al..Deer antler innervation and regeneration[J].Frontiers in bioscience:a journal and virtual library,2012,17;1 389-1 401.
[16]Li C.Clark D E,Lord E A,ea al.Sampling technique to discriminate the different tissue layers of growing antler tips for gene discovery[J].Anat Rec,2002,268(2):125-130.
[17]Kierdorf U,Li C,Price J S.Improbable appendages:Deer antler renewal as a unique case of mammalian regeneration[J].Semin Cell Dev Biol,2009,20(5),535-542.
[18]Li C,Yang F,Sheppard A.Adult stem cells and mammalian epimorphic regeneration-insights from studying annual renewal of deer antlers[J].Curr Stem Cell Res Ther,2009,4(3),237-251.
[19]Li C.Histogenetic aspects of deer antler development[J].Front Biosci(Elite Ed),2013,5:479-489.
[20]Li C,Suttie J M,Clark D E.Morphological observation of antler regeneration in red deer(Cervuselaphus)[J].JMorphol,2004,262(3):731-740.
[21]Banks W J,Newbrey J W.Antler development as aunique modification of mammalian endochondral ossification[A].Antler development in Cervidae,Caesar Kleberg Wildlife Research Institute Kingsville,Texas,1983,1:279-306.
[22]Yang Z H,Gu L J,Zhang D L,et al..Red deer antler extract accelerates hair growth by stimulating expression of insulin-like growth factor I in full-thickness wound healing rat model[J].Asian-Australasian journal of animal sciences,2012,25(5):708-716.
[23]李沐.MicroRNA介导的IGF-1基因沉默与端粒酶对鹿茸细胞增殖抑制的影响[D].长春:吉林农业大学,硕士论文,2013.Li M.Study on IGF-1 Gene Silence Mediated by microRNA and the Effect of Telomerase Against Proliferation of Antler Cells[D].Changchun;Jilin Agricultural University,Master Dissertation,2013.
[24]王守堂.PTHrP及其受体在梅花鹿茸角中的表达与调节[D].长春:吉林大学,硕士论文.2013.Wang S T.Expression and Regulation of PTHrP and Its Receptor in Sika Deer Antler[D].Changchun;Jilin University,Master Dissertation,2013.
[25]Barling P M,Lai A K,Nicholson L F.Distribution of EGF and its receptor in growing red deer antler[J].CellBiolInt,2005,29(3):229-236.
[26]Wegner M.From head to toes;the multiple facets of Sox proteins[J].Nucleic Acids Res,1999,27(6):1 409-1 420.
[27]江玲霞.猪Sox9基因cDNA的全长克隆及功能预测[D].南京:南京农业大学,硕士学位论文.2008.Jiang L X.Cloning and Function Prediction of the Full-Length cDNA of Porcine Sox9 Gene[D].Nanjing;Nanjing Agricultural UniVersity,Master Dissertation,2008.
[28]赵辰,黄伟,梁熙,等.Smad7在Sox9增强BMP2成软骨效应中的作用[J].第三军医大学学报,2015,37(2):95-100.Zhao C,Huang W,Liang X,et al..Role of Smad7 in Sox9-potentiated and BMP2-induced differentiation of mouse mesenchymal stem cells into chondrocytes[J].J Third Mil Med Univ,2015,37(2):95-100.
[29]Mount J G,Muzylak M,Allen S,et al..Evidence that the canonical Wntsignalling pathway regulates deer antler regeneration[J].Dev Dyn,2006,235(5):1 390-1 399.
[30]Zhang Y,Marsboom G,Toth P T,et al..Mitochondrial respiration regulates adipogenic differentiation of human mesenchymal stem cells[J].PLoS One,2013,8(10):1-12.
[31]Huang P I,Chen Y C,Chen L H,et al..PGC-1 a mediates differentiation of mesenchymal stem cells to brown adipose cells[J].J AtherosclerThromb,2011,18(11):966-980.
[32]Gersbach C A,Byers B A,Pavlath G K,et al..Runx2/Cbfal stimulates transdifferentiation of primary skeletal myoblasts into a mineralizing osteoblastic phenotype[J].Exp Cell Res,2004,300(2):406-417.
[33]Sun H,Yang F,Chu W,et al..Lentiviral-mediated rnai knockdown of cbfa1 gene inhibits endochondral ossification of antler stem cells in micromassculture[J].PloS one,2012,7(10):e47367.
[34]李婷.李沐.孟星宇.等.microRNA介导的IGF1基因沉默对鹿茸软骨细胞增殖的影响[J].西北农林科技大学学报,2013,41(11):7-12.Li T,Li M,Meng X Y,et al..Effect of microRNA-mediated IGF1 gene silencing on the proliferation of deer antler cartilage cells[J].Journal of Northwest A&F University,2013,41(11):7-12.
[35]Huarte M,Sanz-Ezquerro J J,Roncal F,et al..PA subunit from influenza virus polymerase complex interacts with a cellular protein with homology to a family of transcriptional activators[J].J Virol,2001,75(18):8 597-8 604.
[36]Guo J,Wang W,Liao P,et al..Identification of serum biomarkers forpancreaticadenocarcinoma by proteomic analysis[J].Cancer Sci,2009,100(12):2292-2301.
[37]张登禄,韩金祥,崔亚洲,等.胰腺癌转移相关基因C140rf166的真核表达及其蛋白相互作用的蛋白质组学筛选[J].中国医药生物技术,2010,5(3):189-192.Zhang D L,Han J X,Cui Y Z,et al..The eukaryotic expression of pancreatic cancer relted gene C14orfl66 and screening of its interacting proteins by Proteomics methods[J].Chin Med Biotechnol,2010,5(3):189-192.
[38]张登禄,韩金祥,崔亚洲,等.C140rf166异位表达对Hela细胞增殖与迁移能力的影响[J].中国肿瘤防治杂志,2010,17(10):740-743.Zhang D L,Han J X,Cui Y Z,et al..Ectopic expression of C14orf166 and its effect on Hela cell proliferation and immigration[J].CHIN J CANCER PREV TREAT,2010,17(10):740-743.
[39]郭静会.胰腺癌血清蛋白质指纹图谱及CCR7与胰腺癌淋巴结转移相关性研究[D].上海:复旦大学.2010.Guo J H.The study of Protein Profiling of Pancreatic cancer and correlation of CCR7 with lymph node metastasis of Pancreatic caneer[D].Shanghai:Fudan University,Doctoral Dissertation,2010.
[40]Howng S L,Hsu H C,Cheng T S,et al..A novel ninein-interaction protein,CGI-99,blocks ninein phosphorylation by GSK3βand is highly expressed in brain tumors[J].FEBS letters,2004,566(1):162-168.
[41]王静.脑发育不同阶段蛋白质组学及血清脑红蛋白含量的研究[D].北京:中国人民解放军军医进修学院,2006.Wang J.Brain Porteomie Study in Dieffrent Developmental Stages And The Study On Expression Level of Neuorglobin in Serum[D].Beijing;Chinese PLA Postgraduate Medical School,Master Dissertation,2006.
[42]Perez-Gonzalez A,Pazo A,Navajas R,et al..hCLE/C14orf'166 associates with DDXl-HSP C117-FAM98B in a novel transcription-dependent shuttling RNA-transporting complex[J].PloS one,2014,9(3):e90957.
[43]Ariel Rodriguez,Alicia Pe'rez-Gonza'lez,Amelia Nieto.Cellular Human CLE/C14orfl66 Protein Interacts with Influenza Virus Polymerase and Is Required for Viral Replication[J].Journal of virology,2011,85(22):12 062-12 066.
[44]宋家升.H5N1亚型禽流感病毒对家鸭致病力分子机制的研究[D].北京:中国农业科学院.2010.Song J S.Detecting virulence determinants of H5N1 avain influenza virus for ducks[D].Beijing;Chinese Academy of Agricultural Sciences,Doctoral Dissertation,2010.
[45]Li C,Waldrup K A,Corson ID,et al..Histogenesis of antlerogenie tissues cultivated in diffusion chambers in vivo in red deer(Cervuselaphus)[J].J Exp Zool,1995,272(5):345-355.
[46]Li C,Harris A J,Suttie J M.Tissue interactions and antlerogenesis;new findings revealed by a xenograft approach[J].J Exp Zool,2001,290(1):18-30.
[47]Li C,Suttie J M.Deer antlerogenie periosteum;a piece of postnatally retained embryonic tissue[J].AnatEmbryol,2001,204(5):375-388.
[48]Clark D E,Lord E A,Suttie J M.Expression of VEGF and pleiotrophin in deer antler[J].Anat Rec A Discov Mol Cell Evol Biol,2006,288(12):1 281-1 293.
[49]Li C,Harper A,Puddick J,et al..Proteomes and signalling pathways of antler stem cells[J].PLoS One,2012,7(1):e30026.
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