羊胎儿期皮肤毛囊发育及调控机制的研究概述

doi:10.16431/j.cnki.1671-7236.2023.02.021

摘要/Abstract

摘要： 中国绵山羊养殖历史悠久,品种资源丰富,在国民经济中起重要作用。羊毛按纤维类型可分为同质毛和异质毛,同质毛的特点是被毛由同一类型的羊毛纤维组成,其特点是羊毛的纤维直径、长度、卷曲等外表特征基本相同,异质毛的特点是被毛中兼有绒毛、有髓毛、无卷曲和少卷毛等不同类型的羊毛纤维,其化学、物理性能均不相同。毛囊是控制哺乳动物被毛生长的重要器官,是唯一具有周期性发育并可终生再生的器官,控制着被毛发生发育与自然脱落。毛囊依据其形成时间和结构可分为初级毛囊(primary follicles)和次级毛囊(secondary follicles),初级毛囊生长髓质发达的粗毛,次级毛囊产生无髓毛的绒毛。有髓毛较粗长且弯曲少,多用于加工粗纺织品、毛毯、地毯、毡制品;无髓毛直径一般不超40 μm,弯曲多,是毛纺工业的优质原料。毛囊的发育受到不同信号通路和基因的影响,如Wnt信号通路、骨形态发生蛋白质(BMP)信号通路、成纤维细胞生长因子(FGF)信号通路、音猬因子(SHH)信号通路、Notch信号通路等以及角蛋白家族基因、BMPs家族基因、同源异型盒基因(Hox)等。作者对国内外对绵山羊胎儿期皮肤毛囊发育及调控机制进行综述,以期为提高以绵山羊被毛产量及改善被毛品质为主要育种目标的选育提供参考。

关键词: 绵山羊; 胎儿期; 毛囊发育; 调控机制

Abstract: China has a long history of goat breeding and rich variety resources,which play an important role in the national economy.Wool can be divided into homogeneous fleece and heterogeneous fleece according to fiber type.The characteristic of homogeneous fleece is that the coat is composed of the same type of wool fibers,and its characteristics are that the fiber diameter,length,curl,and other appearance characteristics of wool are basically the same.The characteristic of heterogeneous fleece is that there are different types of wool fibers in the coat,such as down fleeces,medullary fleeces,and non-curly and oligo-curly fleeces.The chemical,physical and mechanical properties of wool fibers of heterogeneous fleece are different from those of homogeneous fleece. Hair follicle is an important organ with periodic development and lifelong regeneration,which controls the development and natural shedding of the hair.The hair follicles can be divided into primary follicles and secondary follicles according to their formation time and structure.The primary follicles develop subsequently developed coarse hairs,while secondary hair follicles produce unmyelinated villi.Being relatively thicker,longer,and less curved,the medullated wool is mainly used to process coarse textiles,blankets,carpets,and felt products.In contrast,non-medullated wool is an excellent raw material for the wool spinning industry because it is generally not more than 40 μm in diameter and has much curvature.Hair follicle development is affected by different genes and signaling pathways,such as Wnt signaling pathway,bone morphogenetic protein (BMP) signaling pathway,fibroblast growth factor (FGF) signaling pathway,sonic hedgehog(SHH) signaling pathway,Notch signaling pathway,keratin family genes, BMPs family genes,homologous box genes (Hox),etc.The author reviewes the literature on the development and regulation of hair follicles in the fetal skin of sheep,intending to provide a reference for the selection and breeding of sheep whose main breeding objectives are to improve coat yield and coat quality.

Key words: sheep and goats; fetal stag; hair follicle development; regulatory mechanism

中图分类号:

S813.1

秦志云, 姜怀志. 羊胎儿期皮肤毛囊发育及调控机制的研究概述[J]. 中国畜牧兽医, 2023, 50(2): 638-646.

QIN Zhiyun, JIANG Huaizhi. An Overview of Research on Skin Hair Follicle Development and Regulatory Mechanism in Sheep During the Fetal Period[J]. China Animal Husbandry and Veterinary Medicine, 2023, 50(2): 638-646.

参考文献

[1] 高丽霞,刘志红,李金泉.角蛋白5基因在内蒙古绒山羊皮肤毛囊发育周期中的表达模式分析[J].中国畜牧杂志,2018,54(7):27-30. GAO L X,LIU Z H,LI J Q.Analysis of the expression pattern of keratin 5 gene in the development cycle of hair follicle skin of Inner Mongolia cashmere goat[J]. Chinese Journal of Animal Science,2018,54(7):27-30.(in Chinese)
[2] 李玉荣.内蒙古绒山羊不同时期皮肤中相关基因差异表达及原位杂交的研究[D].呼和浩特:内蒙古农业大学,2008. LI Y R.Study on difference expressing and in situ hybrition of skin correlational gene in Inner Mongolian cashmere goat at different stage[D].Hohhot:Inner Mongolia Agricultural University,2008.(in Chinese)
[3] 国家遗传资源委员会.中国畜禽遗传资源志-羊志[M].北京:中国农业出版社,2011. NATIONAL COMMISSION ON GENETIC RESOURCES.Animal Genetic Resources in China:Sheep and Goats[M].Beijing:China Agriculture Press,2011.(in Chinese)
[4] 陈耀春.中国羊业现状和存在的问题[J].农技服务,2004,2:4-6. CHEN Y C.Current situation and existing problems of China’s sheep industry[J]. Agricultural Technology Service,2004,2:4-6.(in Chinese)
[5] 赵印.2021年中国羊业生产形势及2022年趋势展望[J].畜牧产业,2022,3:4. ZHAO Y.Production situation of Chinese sheep industry in 2021 and trend outlook in 2022[J]. Animal Agriculture,2022,3:4.
[6] 辛国桢.澳洲美利奴羊[J].吉林畜牧兽医,1997,4:22. XIN G Z.Australian Merino sheep[J]. Jilin Animal Husbandry and Veterinary Medicine,1997,4:22.(in Chinese)
[7] 道良佐.澳洲美利奴羊的育种[J].国外畜牧学(草食家畜),1983,3:4-6. DAO Z L.Breeding of Australian Merino sheep[J]. Foreign Animal Husbandry (Herbivorous Livestock),1983,3:4-6.(in Chinese)
[8] 羊业分会新西兰羊业考察篇[J].饲料与畜牧,2019,3:54-60. Sheep industry branch New Zealand sheep industry investigation[J]. Feed and Animal Husbandry,2019,3:54-60.(in Chinese)
[9] 史志诚,杨黎旭,邵先华,等.美国的肉羊业[J].新疆畜牧业,1995,5:27-30. SHI Z C,YANG L X,SHAO X H,et al.Meat and sheep industry in the United States[J]. Xinjiang Animal Husbandry, 1995,5:27-30.(in Chinese)
[10] 郭旭.英国现代养羊业的特点及主要经验[J].农业技术与装备,2009,23:55-57. GUO X.Characteristics and main experience of modern sheep breeding industry in Britain[J]. Agricultural Technology and Equipment,2009,23:55-57.(in Chinese)
[11] 杨旭.外源褪黑激素对绒山羊绒毛和皮肤毛囊周期性变化的影响[D].通辽:内蒙古民族大学,2018. YANG X.Effects of exogenous melatonin on the periodic changes of villus and skin hair follicles in cashmere go[D].Tongliao:Inner Mongolia University for Nationalities,2018.(in Chinese)
[12] 杨涛.毛囊生长影响因素及毛囊干细胞分化的诱导研究[D].福州:福建医科大学,2011. YANG T.Study on the influence factor in hair follicle growth and the induced differentiation of hair follicle stem cell[D].Fuzhou:Fujian Medical University,2011.(in Chinese)
[13] DRISKELL R R,CLAVEL C,RENDL M,et al.Hair follicle dermal papilla cells at a glance[J]. Journal of Cell Science,2011,124(8):1179-1182.
[14] 张志丹.Wnt10b调控人毛囊生长的分子机制实验研究[D].广州:南方医科大学,2013. ZHANG Z D.The experimental study of the molecular regulation mechanism of Wnt10b on the human hair follicles[D].Guangzhou:Southern Medical University,2013.(in Chinese)
[15] 席海燕,郑琰,周欢敏.内蒙古绒山羊皮肤干细胞定位的初步研究[J].湖北农业科学,2009,48(2):4. XI H Y,ZHENG Y,ZHOU H M.Localization of skin stem cells of Inner Mongolia cashmere goats[J]. Hubei Agricultural Sciences,2009,48(2):4.(in Chinese)
[16] 郭雪峰,包鹏甲,常永芳,等.哺乳动物毛囊发育及调控研究进展[J].中国畜牧兽医,2019,46(2):387-394. GUO X F,BAO P J,CHANG Y F,et al.Research progress on the development and regulation of mammalian hair follicles[J]. China Animal Husbandry & Veterinary Medicine,2019,46(2):387-394.(in Chinese)
[17] 吴素芳,马涛,李健宇,等.调控毛囊发育的Wnt信号通路研究进展[J].动物医学进展,2019,40(4):92-95. WU S F,MA T,LI J Y,et al.Progress on Wnt signaling regulating hair follicle development[J]. Progress in Veterinary Medicine,2019,40(4):92-95.(in Chinese)
[18] 陈维云.山羊毛囊发育和毛色相关基因的克隆与表达分析[D].泰安:山东农业大学,2012. CHEN W Y.Molecular cloning and expression analysis of genes involved in the hair follicle development and coat color in Jining Gray goat[D].Taian:Shandong Agricultural University,2012.(in Chinese)
[19] 范一星.不同毛被类型内蒙古绒山羊皮肤毛囊周期性生长及相关分子机理研究[D].内蒙古农业大学,2019. FAN Y X.Study on different hair types of Inner Mongolia cashmere goat in periodic changes of skin and hair follicles and related molecular mechanisms[D].Hohhot:Inner Mongolia Agricultural University,2019.(in Chinese)
[20] 姜维.太行黑山羊毛囊生长—退行期皮肤组织差异表达基因的筛选与鉴定[D].杨凌:西北农林科技大学,2012. JIANG W.Screening and identifying regulatory genes associated with Taihang Black goat hair follicle anagen-catagen transition[D].Yangling:Northwest A & F University,2012.(in Chinese)
[21] 贺延玉,罗玉柱,程李香,等.河西绒山羊毛囊超微结构的周期性变化[J].中国农业科学,2012,45(13):2779-2786. HE Y Y,LUO Y Z,CHENG L X,et al.Identification of the secondary follicle cycling of Hexi cashmere goat[J]. Scientia Agricultura Sinica,2012,45(13):2779-2786.(in Chinese)
[22] 吕雪峰,叶小芳,郑文新,等.新疆山羊皮肤毛囊结构及周期性发育规律的研究[J].安徽农业科学,2012,40(24):12094-12096. LÜ X F,YE X F,ZHENG W X,et al.Study on the structure and cyclical development of skin and hair follicles in Xinjiang goats[J]. Journal of Anhui Agricultural Sciences,2012,40(24):12094-12096.(in Chinese)
[23] 金凤,李金泉,刘东军,等.内蒙古罕山白绒山羊毛囊生长规律的研究[J].中国草食动物,2008,26(1):17-18. JIN F,LI J,LIU D J,et al.Study on the growth law of White Velvet goat wool sac in Hanshan Inner Mongolia[J]. China Herbivores Science, 2008,26(1):17-18.(in Chinese)
[24] 张春兰,李金泉,尹俊,等.内蒙古阿尔巴斯绒山羊毛囊生长周期性变化的研究[J].中国畜牧杂志,2005,41(9):10-13. ZHANG C L,LI J Q,YIN J,et al.Study on hair follicles periodical variety in Inner Mongolia Arbas White cashmere goats[J]. Chinese Journal of Animal Science,2005,41(9):10-13.(in Chinese)
[25] 赵艳丽,姜怀志,张世伟,等.辽宁绒山羊皮肤毛囊细胞凋亡特点的研究[J].中国畜牧兽医,2011,38(8):35-38. ZHAO Y L,JIANG H Z,ZHANG S W,et al.Study on hair follicle apoptosis of Liaoning cashmere goats[J]. China Animal Husbandry & Veterinary Medicine,2011,38(8):35-38.(in Chinese)
[26] LYNE A G,HOLLIS D E.The structure and development of the epidermis in sheep fetuses[J]. Journal of Ultrastructure Research,1972,38(5-6):444-458.
[27] 杨涵羽璐,杨永林,杨华,等.中国美利奴羊(新疆军垦型)胎儿期皮肤毛囊结构及形态学观察[J].西南农业学报,2021,34(1):202-207. YANG H Y L,YANG Y L,YANG H,et al.Fetal skin hair follicle development and morphology of Chinese Merino(Xinjiang Junken type)[J]. Southwest China Journal of Agricultural Sciences,2021,34(1):202-207.(in Chinese)
[28] 程瑞禾,路步高,谢成侠,等.湖羊胎儿期皮肤和毛囊发生的研究[J].南京农业大学学报,1981,4(4):96-108. CHENG R H,LU B G,XIE C X,et al.Development of skin and hair follicle at the fetal stages of the sheep[J]. Journal of Nanjing Agricultural University,1981,4(4):96-108.(in Chinese)
[29] 董彬,崔志峰,尹逊河,等.济宁青山羊皮肤毛囊的组织学特性研究[J].山东农业大学学报(自然科学版),2010,41(2):258-262. DONG B,CUI Z F,YIN S H,et al.Studies on histological characteristics of Jining Gray goat skins and hair follicle[J]. Journal of Shandong Agricultural University(Natural Science Edition),2010,41(2):258-262.(in Chinese)
[30] 胡艳霞,董彬,崔志峰,等.济宁青山羊不同发育阶段皮肤MTR1A的差异表达量及其与毛囊组织学结构的相关性研究[A].全国动物遗传育种学术讨论会[C].2009. HU Y X,DONG B,CUI Z F,et al.Differential expression of skin MTR1A in Jining Green goats at different developmental stages and their correlation with hair follicle histological structure[A].National Symposium on Animal Genetics and Breeding[C].2009.(in Chinese)
[31] 尹俊.内蒙古绒山羊毛囊发育、生长周期及相关基因的研究[D].呼和浩特:内蒙古大学,2004. YIN J.Study on the development、growth cycle and related genes of wool sacs in Inner Mongolia[D].Hohhot:Inner Mongolia University,2004.(in Chinese)
[32] 张燕军,尹俊,李长青,等.内蒙古阿尔巴斯绒山羊胎儿期皮肤毛囊发生发育规律研究[J].畜牧兽医学报,2006,8:761-768. ZHANG Y J,YIN J,LI C C,et al.Study on developmengt of skin and hair follicle from fetal Inner Mongolian Arbas cashmere goats[J]. Acta Veterinaria et Zootechnica Sinca,2006,8:761-768.(in Chinese)
[33] 陈洋,陈辉,常青,等.辽宁绒山羊胎儿期皮肤毛囊发生发育的研究[J].中国畜牧杂志,2013,49(11):18-20. CHEN Y,CHEN H,CHANG Q,et al.Study on the occurrence and development of skin hair follicles in the fetal stage of Liaoning cashmere goat[J]. Chinese Journal of Animal Science,2013,49(11):18-20.(in Chinese)
[34] 尹定子,宋海云.Wnt信号通路:调控机理和生物学意义[J].中国细胞生物学学报,2011,33(2):103-111. YIN D Z,SONG H Y.Regulation of Wnt signaling:Mechanism and biological significance[J]. Chinese Journal of Cell Biology,2011,33(2):103-111.(in Chinese)
[35] 吴海青,马跃军,勿都巴拉,等.阿尔巴斯绒山羊产绒量性状相关基因挖掘的研究[J].畜牧与饲料科学,2020,41(1):34-41. WU H Q,MA Y J,WU D B L,et al.Mining of cashmere yield associated genes in Alpas cashmere goats[J]. Animal Husbandry and Feed Science,2020,41(1):34-41.(in Chinese)
[36] 王正森.BMP和FGF信号在小鼠毛囊发育中的作用及机制[D].福州:福建师范大学,2021. WANG Z S.The role and mechanism of BMP and FGF signaling in the development of hair follicle in mice[D].Fuzhou:Fujian Normal University,2021.(in Chinese)
[37] 吴萍.Notch信号通路与绒山羊绒毛生长的研究[D].呼和浩特:内蒙古农业大学,2010. WU P.The research on Notch signal pathway and the cashmere growth in cashmere goats[D].Hohhot:Inner Mongolia University,2010.(in Chinese)
[38] 刘公言,白莉雅,李福昌,等.毛囊发育与周期性生长的调控信号通路研究进展[J].畜牧与兽医,2021,53(1):125-129. LIU G Y,BAI Y L,LI F C,et al.Progress in research on the regulatory signaling pathway of hair follicle development and periodic growth[J]. Animal Husbandry & Veterinary Medicine, 2021,53(1):125-129.(in Chinese)
[39] 王乐乐,张燕军,杨坤,等.Wnt通路中部分基因在绒山羊胚胎皮肤毛囊中的表达[J].中国生物化学与分子生物学报,2016,32(1):99-105. WANG L L,ZHANG Y J,YANG K,et al.Wnt pathway related gene expression in skin follicles of cashmere goat embryo[J]. Chinese Journal of Biochemistry and Molecular Biology,2016,32(1):99-105.(in Chinese)
[40] WU Z,HAI E,DI Z,et al.Using WGCNA (weighted gene co-expression network analysis) to identify the Hub genes of skin hair follicle development in fetus stage of Inner Mongolia cashmere goat[J]. PLoS One,2020,15(12):e0243507.
[41] NIE Y,LI S,ZHENG X,et al.Transcriptome reveals long non-coding RNAs and mRNAs involved in primary wool follicle induction in Carpet sheep fetal skin[J]. Frontiers Physiology, 2018,9:446.
[42] 杨坤.EDA、EDAR在绒山羊胎儿皮肤毛囊中的表达及对相关基因表达的影响[D].呼和浩特:内蒙古农业大学,2017. YANG K.EDA,EDAR Expression in the fetal hair follicles of cashmere goat and the effects on expression of related genes[D].Hohhot:Inner Mongolia Agricultural University,2017.(in Chinese)
[43] 赵帅. Wnt10 b,β-catenin, FGF18基因在甘肃高山细毛羊胎儿皮肤毛囊中的表达规律研究[J].甘肃农业大学学报,2015,50(5):6-14. ZHAO S.Expression patterns of Wnt10 b,β-catenin and FGF18 genes in fetal hair follicles of Gansu alpine fine-wool sheep[J]. Journal of Gansu Agricultural University,2015,50(5):6-14.(in Chinese)
[44] 高晔.陕北白绒山羊皮肤毛囊发育早期的转录组学分析及 VDR基因的功能研究[D].杨凌:西北农林科技大学,2017. GAO Y.Comparative transcriptome analysis of fetal skin reveals key genes related to hair follicle morphogenesis in Shanbei White cashmere goats and functional research of VDR[D].Yangling:Northwest A&F University,2017.(in Chinese)
[45] 尚方正,韩文静,吴志红,等.非编码RNA及其在畜禽中的应用研究进展[J].中国畜牧兽医,2021,48(1):235-247. SHANG F Z,HAN W J,WU Z H,et al.Research Advance on Application of non-coding RNA in Livestock and Poultry[J]. China Animal Husbandry & Veterinary Medicine,2021,48(1):235-247.(in Chinese)
[46] 李志娟,高士争,赵素梅.microRNA的生物学功能及其在动物肌肉和脂肪组织中表达的研究进展[J].中国畜牧兽医,2012,39(7):102-107. LI Z J,GAO S Z,ZHAO S M.Research progress of biological function of microRNA and its expression in animal muscle and adipose tissue[J]. China Animal Husbandry & Veterinary Medicine,2012,39(7):102-107.(in Chinese)
[47] YANG H Y L,YANG H,SHI G Q,et al.Expression profile analysis of microRNAs during hair follicle development in the sheep foetus[J]. Bioscience, Biotechnology, and Biochemistry,2019,83(6):1045-1061.
[48] 阿布来提·苏来曼.苏博美利奴羊毛囊发育相关lncRNA的筛选与功能分析[D].乌鲁木齐:新疆农业大学,2018. SULAYMAN A.Identification and functional analysis of lncRNA related to development of Subo Merino’s hair follicle[D].Urumqi:Xinjiang Agricultural University,2018.(in Chinese)
[49] 张小伟.苏博美利奴羊毛囊发育相关ncRNA分子调控网络的构建及关键基因的验证[D].乌鲁木齐:新疆农业大学,2021. ZHANG X W.Construction of ncRNA network related to hair follicle development in Subo Merino sheep[D].Urumqi:Xinjiang Agricultural University,2021.(in Chinese)
[50] HAI E,HAN W,WU Z,et al.Chi-miR-370-3p regulates hair follicle morphogenesis of Inner Mongolian cashmere goats[J]. G3-Genes Genomes Genetics,2021,11(5):jkab091.
[51] 尚方正,马荣,戎友俊,等.绒山羊胎儿期毛囊发生发育关键circRNA的鉴定及功能预测分析[J].中国农业大学学报,2022,27(4):207-218. SHANG F Z,MA R,RONG Y J,et al.Identification and functional prediction analysis of key circRNA for follicular development in the fetal stage of cashmere goat[J]. Journal of China Agricultural University 2022,27(4):207-218.(in Chinese)