家畜卵巢生理解析与结构功能探究

doi:10.16431/j.cnki.1671-7236.2025.10.005

摘要/Abstract

摘要： 卵巢作为雌性家畜生殖系统的核心组成部分，其结构、功能及其在生殖过程中的作用至关重要。文章系统综述了马、驴、牛、羊、猪等不同家畜卵巢的基本结构与功能特性，涵盖卵巢的位置分布、形态特征、皮质髓质结构以及独特的排卵窝特性等方面；深入剖析了卵巢特性对家畜发情周期、排卵特性以及繁殖性能的影响机制；详细介绍了CRISPR/Cas9基因编辑技术、孤雌激活技术以及高通量测序与生物信息学等新技术在家畜卵巢研究领域中的最新应用进展；同时，还展望了未来家畜卵巢研究的发展方向，重点聚焦于利用基因编辑技术优化家畜繁殖性能，以及借助大数据和人工智能技术提升繁殖效率等方面的潜力，旨在为深入理解家畜繁殖生物学机制、优化繁殖策略以及提高繁殖效率提供一定的科学依据，并为家畜繁殖技术的持续改进与创新提供有力支持。

关键词: 家畜; 卵巢; 繁殖; 形态; 发情周期; 排卵

Abstract: The ovary serves as a core component of the female reproductive system in livestock,with its structure,function,and role in the reproductive process being of paramount importance.This paper systematically reviews the basic structural and functional characteristics of ovaries in various livestock,including horses,donkeys,cattle/cows,sheep/goats,and pigs,specifically covering aspects such as the location distribution,morphological features,cortex-medulla structure,and unique ovulation fossa characteristics of ovaries.This paper delves into the mechanisms by which the ovarian characteristics influence the estrous cycle,ovulation characteristics,and reproductive performance of livestock.Furthermore,this paper comprehensively introduces the latest application progress of novel technologies in the research field of livestock ovaries,such as CRISPR/Cas9 gene editing,parthenogenesis activation,high-throughput sequencing,and bioinformatics.Additionally,this paper outlines the future research directions in the field of livestock ovaries,with a focus on optimizing reproductive performance through gene editing technology and enhancing reproductive efficiency with the aid of big data and artificial intelligence technologies.This paper aims to provide a solid scientific basis for a deeper understanding of the biological mechanisms of livestock reproduction,the optimization of reproduction strategies,and the improvement of reproductive efficiency,while also offering strong support for the continuous improvement and innovation of livestock reproduction technologies.

Key words: livestock; ovary; reproduction; morphology; estrous cycle; ovulation

中图分类号:

S821.3

何千倩, 刘琦, 刘源壹, 王娜, 温鑫, 韩宝祥, 格日乐其木格. 家畜卵巢生理解析与结构功能探究[J]. 中国畜牧兽医, 2025, 52(10): 4575-4591.

HE Qianiqan, LIU Qi, LIU Yuanyi, WANG Na, WEN Xin, HAN Baoxiang, Gerelchimeg. Research on Physiological Analysis and Structural Function Exploration of Livestock Ovarian[J]. China Animal Husbandry & Veterinary Medicine, 2025, 52(10): 4575-4591.

参考文献

[1] PAMULA M C,LEHMANN R.How germ granules promote germ cell fate[J].Nature Reviews Genetics,2024,25(11):803-821.
[2] HONG T K,SONG J H,LEE S B,et al.Germ cell derivation from pluripotent stem cells for understanding in vitro gametogenesis[J].Cells,2021,10(8):1889.
[3] BHARTI D,TIKKA M,LEE S Y,et al.Female germ cell development,functioning and associated adversities under unfavorable circumstances[J].International Journal of Molecular Sciences,2021,22(4):1979.
[4] TSUTSUI K,UBUKA T,UKENA K.Advancing reproductive neuroendocrinology through research on the regulation of GnIH and on its diverse actions on reproductive physiology and behavior[J].Frontiers in Neuroendocrinology,2022,64:100955.
[5] GRAY P B,STRAFTIS A A,BIRD B M,et al.Human reproductive behavior,life history,and the challenge hypothesis:A 30-year review,retrospective and future directions[J]. Hormones and Behavior,2020,123:104530.
[6] KOHLBRENNER T,BERGER S,LARANJEIRA A C,et al.Actomyosin-mediated apical constriction promotes physiological germ cell death in C.elegans[J].PLoS Biology,2024,22(8):e3002775.
[7] SCHUMACHER A,GEHLEN H.Health of Polo horses[J].Animals (Basel),2024,14(12):1735.
[8] 陶力.蒙古马卵巢发育规律及调控机理研究[D].呼和浩特:内蒙古农业大学,2023. TAO L.Study on the developmental pattern and regulation mechanism of the ovary in Mongolian horses[D].Hohhot:Inner Mongolia Agricultural University,2023.(in Chinese)
[9] 陶力,神英超,阿鲁斯,等.马卵巢石蜡切片制作体系的优化[J].黑龙江动物繁殖,2021,29(6):4-9. TAO L,SHEN Y C,ALUSI,et al.Optimization of paraffin section making system of horse ovary[J].Heilongjiang Journal of Animal Reproduction,2021,29(6):4-9.(in Chinese)
[10] PIOTROWSKA-TOMALA K K,JONCZYK A W,SZÓSTEK-MIODUCHOWSKA A,et al.Intrauterine devices influence prostaglandin secretion by equine uterus:In vitro and in vivo studies[J]. BMC Veterinary Research,2024,20(1):46.
[11] 陶力,神英超,伊敏娜,等.马属动物卵巢发育研究进展[J].中国畜牧杂志,2022,58(1):38-43. TAO L,SHEN Y C,YI M N,et al.Research progress on ovarian development of equine[J]. Chinese Journal of Animal Science,2022,58(1):38-43.(in Chinese)
[12] 张忠诚.家畜繁殖学[M].北京:中国农业出版社,2004. ZHANG Z C.Reproduction in Domestic Animals[M].Beijing:China Agriculture Press,2004.(in Chinese)
[13] 阿鲁斯.蒙古马自然排卵与促排排卵的对比研究[D].呼和浩特:内蒙古农业大学,2023. ALUSI.A comparative study of natural ovulation and induced ovulation in Mongolian mares[D].Hohhot:Inner Mongolia Agricultural University,2023.(in Chinese)
[14] WESSON J A,GINTHER O J.Influence of season and age on reproductive activity in pony mares on the basis of a slaughterhouse survey[J]. Journal of Animal Science,1981,52(1):119-129.
[15] ONO M,AKUZAWA H,NAMBO Y,et al.Analysis of the equine ovarian structure during the first twelve months of life by three-dimensional internal structure microscopy[J].Journal of Veterinary Medical Science,2016,77(12):1599-1603.
[16] O’SHEA J D.A histological study of non-follicular cysts in the ovulation fossa region of the equine ovary[J].Journal of Morphology,1968,124(3):313-320.
[17] 高宇. β神经生长因子促蒙古马排卵的初步应用[D]. 长春: 吉林大学, 2020. GAO Y. The application of nerve growth factor-beta in promoting ovulation of Mongolian mares[D]. Changchun: Jilin University, 2020. (in Chinese)
[18] BHINTZ H F,CYMBALUK N F.Nutrition of the horse[J].Annual Review of Nutrition,1994,14:243-267.
[19] 玉山江,罗永明,陈童.生殖激素在马、驴繁殖中的应用[J].黑龙江动物繁殖,2022,30(2):17-22. YU S J,LUO Y M,CHEN T.Reproductive hormones application in horses and donkey reproduction[J].Heilongjiang Journal of Animal Reproduction,2022,30(2):17-22.(in Chinese)
[20] 徐文慧,杨振,芦文圆,等.马繁殖季节和非繁殖季节变化观察[J/OL].畜牧兽医科学(电子版),2020,21:185-186. XU W H,YANG Z,LU W Y,et al.Observation on the changes of horse breeding season and non-breeding season[J/OL]. Veterinary and Animal Science (Electronic Edition),2020,21:185-186.(in Chinese)
[21] HEIDLER B,AURICH J E,POHL W,et al.Body weight of mares and foals,estrous cycles and plasma glucose concentration in lactating and non-lactating Lipizzaner mares[J].Theriogenology,2004,61(5):883-893.
[22] AURICH C.Reproductive cycles of horses[J].Animal Reproduction Science,2011,124(3-4):220-228.
[23] SEGABINAZZI L G T M,GILBERT R O,AMBROSIA R L,et al.Structural and functional dynamics of the ovary and uterus during the estrous cycle in donkeys in the Eastern Caribbean[J].Animals (Basel),2022,13(1):74.
[24] GINTHER O J,GASTAL E L,GASTAL M O,et al.Comparative study of the dynamics of follicular waves in mares and women[J]. Biology of Reproduction,2004,71(4):1195-1201.
[25] EVANS A C.Characteristics of ovarian follicle development in domestic animals[J]. Reproduction in Domestic Animals,2003,38(4):240-246.
[26] GABRYS J,PIETRAS N,KOWAL-MIERZWA W,et al.Investigating the impact of extracellular vesicle addition during IVM on the fertilization rate of equine oocytes following ICSI[J].Reproductive Biology,2024,24(4):100967.
[27] PIERSON R A,GINTHER O J.Follicular population dynamics during the estrous cycle of the mare[J]. Animal Reproduction Science,1987,14(3):219-231.
[28] PIERSON R A,GINTHER O J.Ultrasonic evaluation of the preovulatory follicle in the mare[J]. Theriogenology,1985,24(3):359-368.
[29] TANG Z,ZHANG Z,LIN Q,et al.HIF-1α/BNIP3-mediated autophagy contributes to the luteinization of granulosa cells during the formation of Corpus luteum[J]. Frontiers in Cell and Developmental Biology,2020,8:619924.
[30] 郭少仙,张阳阳,姚利双.马超数排卵技术研究进展[J].中国畜牧业,2017,17:54-55. GUO S X,ZHANG Y Y,YAO L S.Research progress on superovulation techniques in horses[J]. China Animal Industry,2017,17:54-55.(in Chinese)
[31] 王振汉.蒙古母马发情规律调查[J].华北农学报,1965,4:47-52. WANG Z H.Investigation on the estrus patterns of Mongolian mares[J].Acta Agriculturae Boreali-Sinica,1965,4:47-52.(in Chinese)
[32] ISHAK G M,FEUGANG J M,PECHANOVA O,et al.Follicular-fluid proteomics during equine follicle development[J].Molecular Reproduction and Development,2022,89(7):298-311.
[33] GINTHER O J.Systemic and intrafollicular components of follicle selection in mares[J].Domestic Animal Endocrinology,2017,59:116-133.
[34] GINTHER O J.Contributions to mare reproduction research by the Ginther team[J].Journal of Equine Veterinary Science,2023,126:104295.
[35] 张永昌.非繁殖季节母驴诱导发情及排卵的研究[D].呼和浩特:内蒙古农业大学,2018. ZHANG Y C.Study on induction of oestrus and ovulation in female breeds during non-breeding season[D].Hohhot:Inner Mongolia Agricultural University,2018.(in Chinese)
[36] GINTHER O J,BERGFELT D R.Ultrasonic characterization of follicular waves in mares without maintaining identity of individual follicles [J]. Journal of Equine Veterinary Science, 1992,12(6):349-354.
[37] 简洪禹,张志鹏,麻朝晖,等.不同繁殖季节驴卵母细胞体外成熟与孤雌激活胚胎发育情况研究[J].黑龙江畜牧兽医,2019,20:60-63. JIAN H Y,ZHANG Z P,MA Z H,et al.Study on the in vitro maturation and parthenogenetic activation of donkey oocytes and embryonic development in different breeding seasons[J].Heilongjiang Animal Science and Veterinary Medicine,2019,20:60-63.(in Chinese)
[38] 张振威,黄炳舰,王永辉,等.驴繁殖生理与繁殖技术研究进展[J].中国畜牧杂志,2023,59(6):41-48. ZHANG Z W,HUANG B J,WANG Y H,et al.Advances in reproductive physiology and breeding technology of donkeys[J].Chinese Journal of Animal Science,2023,59(6):41-48.(in Chinese)
[39] GINTHER O J,SCRABA S T,BERGFELT D R.Reproductive seasonality of the jenney[J].Theriogenology,1987,27(4):587-592.
[40] TABERNER E,MEDRANO A,PEÑA A,et al.Oestrus cycle characteristics and prediction of ovulation in Catalonian jennies[J]. Theriogenology,2008,70(9):1489-1497.
[41] MIRAGAYA M H,NEILD D M,ALONSO A E.A review of reproductive biology and biotechnologies in donkeys[J].Journal of Equine Veterinary Science,2018,65:55-61.
[42] 张强,王艳萍,郭旭川,等.新疆驴发情周期内卵泡生长发育规律的研究[A].中国畜牧兽医学会动物繁殖学分会第二十次学术研讨会论文集[C].2021. ZHANG Q,WANG Y P,GUO X C,et al.Study on the follicle growth and development law during estrus of Xinjiang donkey[A].Proceedings of the 20th Academic Seminar of Animal Reproduction Branch of Chinese Association of Animal Science and Veterinary Medicine[C].2021.(in Chinese)
[43] VANDEPLASSCHE G M,WESSON J A,GINTHER O J.Behavioral,follicular and gonadotropin changes during the estrous cycle in donkeys[J].Theriogenology,1981,16(2):239-249.
[44] MEIRA C,FERREIRA J C,PAPA F O,et al.Ovarian activity and plasma concentrations of progesterone and estradiol during pregnancy in jennies[J]. Theriogenology,1998,49(8):1465-1473.
[45] BLANCHARD T L,TAYLOR T S,LOVE C L.Estrous cycle characteristics and response to estrus synchronization in mammoth asses (Equus asinus americanus)[J].Theriogenology,1999,52(5):827-834.
[46] CARLUCCIO A,GLORIA A,VERONESI M C,et al.Factors affecting pregnancy length and phases of parturition in Martina franca jennies[J].Theriogenology,2015,84(4):650-655.
[47] PEREZ-MARIN C C,GALISTEO I,PEREZ-RICO A,et al.Effects of breed,age,season,and multiple ovulations on cyclic,PGF2α-induced,and postpartum estrus characteristics in Spanish jennies[J].Theriogenology,2016,85(6):1045-1052.
[48] DÍAZ-DURAN M,ZARCO L,BOETA A M.Ovarian dynamics and estrous cycle length in the donkey (Equus asinus)[J].Theriogenology,2017,103:1-8.
[49] 吴帅帅.马驴高效扩繁技术研究及应用[D].北京：中国农业大学,2019. WU S S.Development of high efficiency horse and donkey reproduction technology and their applications[D].Beijing:China Agricultural University,2019.(in Chinese)
[50] 张强.新疆驴发情周期内卵泡生长发育规律的研究[D].石河子：石河子大学,2020. ZHANG Q.Study on the follicle growth and development law during estrus of Xinjiang donkey[D].Shihezi:Shihezi University,2020.(in Chinese)
[51] MIGNÁCIO F S,MONTECHIESI D F,BERGFELT D R,et al.Ultrasound-guided aspiration of dominant follicles (≥25 mm) followed by luteinization and progesterone production during the estrous cycle in mares[J].Theriogenology,2021,160:128-133.
[52] 葛利江,徐常林,石瑞青,等.德州驴发情周期中卵泡动力学和激素变化规律的研究[J].畜牧兽医学报,2011,42(9):1256-1263. GE L J,XU C L,SHI R Q,et al.Ovarian follicular and hormonal dynamics in Dezhou jenny during estrous cycle monitored by Real-time ultrasonography[J].Acta Veterinaria et Zootechnica Sinica,2011,42(9):1256-1263.(in Chinese)
[53] 张新浩,李海静,姜桂苗,等.气温对德州驴繁殖规律的影响[J].黑龙江畜牧兽医,2017,8:74-76. ZHANG X H,LI H J,JIANG G M,et al.Effects of temperature on the reproductive patterns of Dezhou donkeys [J].Heilongjiang Journal of Animal Science and Veterinary Medicine,2017,8:74-76.(in Chinese)
[54] 曹波.奶牛不孕与子宫和卵巢超声形态学变化关系的研究[D].武汉：华中农业大学,2006. CAO B.Research on the relationships of sterility and ultrasonic morphologic changes of ovaries and uterus in dairy cow[D].Wuhan:Huazhong Agricultural University,2006.(in Chinese)
[55] 王新钧,刘春雨.家畜繁殖学[M].银川:阳光出版社,2013. WANG X J,LIU C Y.Animal Reproduction[M].Yinchuan:Sunshine Press,2013.(in Chinese)
[56] 王锋.动物繁殖学[M].北京:中国农业大学出版社,2022. WANG F.Animal Reproduction[M].Beijing:China Agricultural University Press,2022.(in Chinese)
[57] 赵明礼.同期发情及同期排卵-定时输精技术对奶牛繁殖效率的影响[D].北京：中国农业科学院,2016. ZHAO M L.Effects of estrus synchronization and ovsynch-timed artificial insemination on reproductive performance in Holstein cows and heifers[D].Beijing:Chinese Academy of Agricultural Sciences,2016.(in Chinese)
[58] 陈忠林,赵珈,曹中兴.几种家畜繁殖生产的要点[J].养殖技术顾问,2014,11:62. CHEN Z L,ZHAO J,CAO Z X.Key points of reproduction and production for several domestic animals[J].Modern Animal Husbandry Science & Technology,2014,11:62.(in Chinese)
[59] 蔡淑.奶牛养殖实用技术要点[J].湖北畜牧兽医,2017,38(9):2. CAI S.Management tips for poultry feeding [J]. Hubei Journal of Animal and Veterinary Sciences,2017,38(9):2.(in Chinese)
[60] 李会颖,吕桂英,贾琼.提高母牛繁殖力的技术措施[J].黑龙江动物繁殖,2009,17(2):2. LI H Y,LYU G Y,JIA Q.Technical measures to improve the reproductive performance of cows[J].Heilongjiang Animal Reproduction,2009,17(2):2.(in Chinese)
[61] 赵明礼,张珅,郝海生,等.同期发情和同期排卵-定时输精技术在奶牛繁殖中的应用[J].中国畜牧杂志,2015,51(17):7. ZHAO M L,ZHANG S,HAO H S,et al.Application of estrus synchronization and ovsynch-timed artificial insemination in dairy cattle[J]. Chinese Journal of Animal Science,2015,51(17):7.(in Chinese)
[62] SMITH M F,PERRY G A,ATKINS J A,et al.Physiological principles underlying synchronization of estrus[J].Tierarztl Prax Ausg G Grosstiere Nutztiere,2013,41(1):45-54.
[63] 李权武,武浩,秦鹏,等.影响奶牛情期受胎率的非特异因素调查与分析[J].动物医学进展,2000,S1:145-147. LI Q W,WU H,QIN P,et al.Investigation and analysis of nonspecific factors affecting the conception rate during estrus in dairy cows[J]. Progress in Veterinary Medicine,2000,S1:145-147.(in Chinese)
[64] 程耀斌.对黑白花奶牛发情、排卵及授精效果的比较观测[J].畜禽业,2016,7:42-43. CHENG Y B.Comparative observation of estrus,ovulation,and fertilization effects in Holstein dairy cows[J].Livestock and Poultry Industry,2016,7:42-43.(in Chinese)
[65] 李国富.母畜发情周期中的变化特点与鉴定方法[J].养殖技术顾问,2013,6:64. LI G F.Characteristics of changes and identification methods during the estrous cycle of female livestock[J].Modern Animal Husbandry Science & Technology,2013,6:64.(in Chinese)
[66] 邓立新,齐长明,贺丛.母羊生殖器官的B超探查[J].中国农业科技导报,2005,4:33-37. DENG L X,QI C M,HE C.Sonography of the reproductive organs in ewe[J].Journal of Agricultural Science and Technology,2005,4:33-37.(in Chinese)
[67] 郭亚军,柳苗苗,付德海,等.藏绵羊卵巢组织学及卵泡超微形态的观察[J].畜牧兽医学报,2021,52(2):389-398. GUO Y J,LIU M M,FU D H,et al.Observation of ovary histology and ultrastructure of follicles in Tibetan sheep[J]. Acta Veterinaria et Zootechnica Sinica,2021,52(2):389-398.(in Chinese)
[68] 杨利国.动物繁殖学[M].北京:中国农业出版社,2003. YANG L G.Animal Reproduction [M].Beijing:China Agriculture Press,2003.(in Chinese)
[69] 郝文,王成倩,晏航,等.FSH对多浪羊子宫和卵巢组织及血清生殖激素的影响[J].黑龙江畜牧兽医,2024,7:68-73. HAO W,WANG C Q,YAN H,et al.Effects of FSH on uterine and ovarian tissues and serum reproductive hormones in Duolang sheep[J].Heilongjiang Animal Science and veterinary Medicine,2024,7:68-73.(in Chinese)
[70] ABECIA J A,FORCADA F,GONZÁLEZ-BULNES A.Pharmaceutical control of reproduction in sheep and goats[J].Veterinary Clinics of North America-Food Animal Practice,2011,27(1):67-79.
[71] 王海利,肖喜东,胡军.山羊的生殖生理特点及在生产中的合理应用[J].养殖与饲料,2022,7:21. WANG H L,XIAO X D,HU J.Reproductive physiological characteristics of goats and their rational application in production[J].Animals Breeding and Feed,2022,7:21.(in Chinese)
[72] 徐益兵.提高山羊生产性能关键技术的研究与推广[J].农业开发与装备,2018,10:2. XU Y B.Research and promotion of key technologies for improving goat production performance[J].Agricultural Development & Equipments,2018,10:2.(in Chinese)
[73] 侯衍猛,曹洪防,徐云华,等.莱芜黑山羊发情周期中FSH、LH、E₂和P的分泌规律[J].中国兽医学报,2006,26(3):340-343. HOU Y M,CAO H F,XU Y H,et al.The secretory characteristics of FSH,LH,E₂ and P of Laiwu Black goat during the oestrus cycle[J].Chinese Journal of Veterinary Science,2006,26(3):340-343.(in Chinese)
[74] 阿尔曼·海热,艾克拜尔·艾合麦提,古丽沙热·吾甫尔,等.不同季节吐鲁番黑羊血清中生殖激素的变化规律研究[J].中国畜牧杂志,2024,60(1):255-259. ARMAN H,EKBER A,GULISHARE W,et al.Study on the variation of reproductive hormones in the serum of Turpan Black sheep in different seasons[J].Chinese Journal of Animal Science,2024,60(1):255-259.(in Chinese)
[75] 刘莉莉,何玉英,杨亚文,等.甘加藏羊发情周期和乏情期血浆T₃和T₄浓度动态变化及HPOA轴DIO2和DIO3的表达[J].西北农业学报,2022,31(5):544-551. LIU L L,HE Y Y,YANG Y W,et al.Dynamic changes of T₃ and T₄ in plasma and expression of DIO2 and DIO3 in HPOA axis of Ganjia Tibetan sheep during estrus cycle and anestrus[J].Acta Agriculturae Boreali-occidentalis Sinica,2022,31(5):544-551.(in Chinese)
[76] 向泽宇,王长庭.青藏高原藏羊遗传资源的现状、存在问题及对策[J].中国畜牧兽医文摘,2011,27(2):1-4. XIANG Z Y，WANG C T，Current status,existing problems,and countermeasures of genetic resources of Tibetan sheep on the Qinghai-Tibet plateau[J]. Chinese Journal of Animal Science and Veterinary Medicine Abstracts,2011,27(2):1-4.(in Chinese)
[77] 吴艳芳,仲长胜,陈玉林,等.滩羊发情周期内4种生殖激素动态变化规律研究[J].中国畜牧杂志,2022,58(9):184-187. WU Y F，ZHONG C S,CHEN Y L,et al.Study on the dynamic changes of four reproductive hormones during the estrous cycle in Tan sheep[J].Chinese Journal of Animal Science,2022,58(9):184-187.(in Chinese)
[78] 贺建宁,王金鑫,狄冉,等.常年发情和季节性发情绵羊在不同季节生殖激素变化规律[J].畜牧兽医学报,2013,44(10):1547-1553. HE J N，WANG J X，DI R，et al.Seasonal variations of reproductive hormone level in year-round and seasonal estrus sheep[J].Acta Veterinaria et Zootechnica Sinica,2013,44(10):1547-1553.(in Chinese)
[79] 刘欣.辽宁绒山羊的繁殖特点及提高繁殖成活率的有效方法[J].畜禽业,2020,31(9):38. LIU X.Reproductive characteristics of Liaoning cashmere goats and effective methods for improving reproductive survival rate[J].Livestock and Poultry Industry,2020,31(9):38.(in Chinese)
[80] 王宝珍.绵羊卵巢卵母细胞体外成熟培养的研究[J].甘肃畜牧兽医,2006,36(5):3. WANG B Z.Study on in vitro maturation culture of ovarian oocytes in sheep[J]. Gansu Animal Husbandry and Veterinary Medicine,2006,36(5):3.(in Chinese)
[81] 丁建云.母猪的生殖器官解剖和中挑花阉割术[J].畜牧兽医科技信息,2017,33(8):82. DING J Y.Anatomy of the reproductive organs of sows and the technique of mid-ventral laparotomic spaying[J].Chinese Journal of Animal Husbandry and Veterinary Medicine,2017,33(8):82.(in Chinese)
[82] KYZEKOVÁ P,BELKOVÁ L,MACÁK V,et al.Ovarian activity and the size of oocytes in ovarian follicles in sexually mature gilts in individual seasons[J]. Folia Veterinaria,2019,63(3):53-59.
[83] 梁学超,赵秋云,卿素珠.猪卵巢发育的组织学观察及卵泡闭锁规律研究[A].中国畜牧兽医学会动物解剖及组织胚胎学分会第十九次学术研讨会论文集[C].2016. LING X C，ZHAO Q Y，QIN S Z，Histological observation of porcine ovarian development and study on the law of follicular atresia[A].Proceedings of the 19th Academic Symposium of the Animal Anatomy and Histology and Embryology Branch of the Chinese Association of Animal Science and Veterinary Medicine[C].2016.(in Chinese)
[84] NONNEMAN D J,SCHNEIDER J F,LENTS C A,et al.Genome-wide association and identification of candidate genes for age at puberty in swine[J].BMC Genomic Data,2016,17:1-9.
[85] 刘宇超.西藏藏猪卵巢卵泡期和黄体期的特征及差异lncRNA的筛选与分析[D].林芝：西藏农牧学院,2024. LIU Y C.Characterization of follicular and luteal phases in the ovaries of Tibetan pigs in Tibet and screening and analysis of differential lncRNAs[D].Linzhi:Tibet Agriculture and Animal Husbandry College,2024.(in Chinese)
[86] 李文杰.猪的繁殖特点与人工授精[J].畜牧兽医科技信息,2018,32(5):116. LI W J.Reproductive characteristics of pigs and artificial insemination[J]. Chinese Journal of Animal Husbandry and Veterinary Medicine,2018,32(5):116.(in Chinese)
[87] 谭冰冰,袁依依,杨日旺,等.德保猪的繁殖性能研究[J].黑龙江畜牧兽医,2023,8:43-47. TAN B B，YUAN Y Y，YANG R W，et al.Study on reproductive performances of Debao pigs[J].Heilongjiang Animal Science and Veterinary Medicine，2023,8:43-47.(in Chinese)
[88] 徐广鹤,胡成波,栾华东,等.辽宁黑猪、二花脸猪、莱芜猪繁殖性能比较研究[A].东北养猪研究会2010学术年会论文集[C].2010. XU G H，HU C B，LUAN H D，et al.Comparative study on reproductive performance of Liaoning Black pigs,Erhualian pigs,and Laiwu pigs[A].Proceedings of the 2010 Academic Annual Meeting of the Northeast Swine Production Research Society[C].2010.(in Chinese)
[89] 娜仁花,刘海蓉,仲永盛,等.大白猪的特征特性与生产性能分析[J].畜牧与饲料科学,2007,2:29-30. NA R H，LIU H R，ZHONG Y S，et al.Characteristics and production performance analysis of Large White pigs[J]. Animal Husbandry and Feed Science,2007,2:29-30.(in Chinese)
[90] 卢晟盛,罗龙兴,兰宗宝,等.广西巴马小型猪雄性初情期和精液性状的分析[J].实验动物科学,2007,6:30-32. LU S S，LUO L X，LAN Z B，et al.Analysis of prepuberty and semen characteristics of male Guangxi Bama Minipig[J]. Laboratory Animal Science,2007,6:30-32.(in Chinese)
[91] 毕台飞,苗飞.香猪繁殖性能测定[J].家畜生态学报,2007,5:82-85. BI T F，MIAO F.Study on the reproductive characters of Xiang pig[J].Journal of Domestic Animal Ecology,2007,5:82-85.(in Chinese)
[92] 王宏辉,王昆山,李瑜鑫,等.藏猪在舍饲条件下的行为观察[J].家畜生态学报,2002,23(2):32-34. WANG H H,WANG K S,LI Y X,et al.Study on Tibet hog’s behaviour characteristics under the condition of feeding[J].Journal of Domestic Animal Ecology,2002,23(2):32-34.(in Chinese)
[93] 杨延彪,曹国顺,郭宏伟,等.甘南蕨麻猪的生态分布及其生产性能[J].家畜生态,2003,24(1):2. YANG Y B,CAO G S,GUO H W,et al.Ecological distribution and productive performance of Juema pig in Southern part of Gansu province[J].Ecology of Domestic Animal,2003,24(1):2.(in Chinese)
[94] 刘祥,王法恩,高洪敏,等.鲁西黑猪繁殖性能的研究[J].畜禽业,2016,11:32-35. LIU X,WANG F E,GAO H M,et al.Study of reproductive performance in Luxi Black pig[J].Livestock and Poultry Industry,2016,11:32-35.(in Chinese)
[95] SOEDE N M,LANGENDIJK P,KEMP B.Reproductive cycles in pigs[J]. Animal Reproduction Science,2011,124(3-4):251-258.
[96] SCHWARZ T,KOPYRA M,NOWICKI J.Physiological mechanisms of ovarian follicular growth in pigs—A review[J].Acta Veterinaria Hungarica,2008,56(3):369-378.
[97] 张国庭.马卵巢卵泡发育波的动态变化观察及不同激素处理效果的研究[D].乌鲁木齐：新疆农业大学,2008. ZHANG G T.Study on the dynamic changes of follicular waves in mare ovaries and the effects of different hormone treatments[D].Urumqi:Xinjiang Agricultural University.2008.(in Chinese)
[98] GINTHER O J.Reproductive biology of the mare.Basic and applied aspects[J].Canadian Veterinary Journal,1992,21(10):289.
[99] 郑丕留.驴生殖器官的解剖[J].解剖学报,1954,2:63-72. ZHENG P L.Anatomy of the reproductive system of mule[J]. Acta Anatomica Sinica,1954,2:63-72.(in Chinese)
[100] 冯玉龙,陈永广,曲洪磊,等.驴卵泡发育规律的研究[J].中国畜牧杂志,2017,53(8):3. FENG Y L,CHEN Y G,QU H L,et al.Study on the law of follicular development in donkeys[J].Chinese Journal of Animal Science,2017,53(8):3.(in Chinese)
[101] FRATTARELLI J L.A prospective analysis of the changes in ovarian morphology during hormonal pituitary suppression before in vitro fertilization [J]. Fertility and Sterility,2006,86(3):577-582.
[102] 刘源壹,李昕俞,巴音那木拉,等.单细胞转录组测序技术及其在动物繁殖中的应用进展[J].畜牧兽医学报,2023,54(2):421-433. LIU Y Y,LI X Y,BAYINNAMULA,et al.Single-cell transcriptome sequencing technology and its application in animal reproduction[J].Acta Veterinaria et Zootechnica Sinica,2023,54(2):421-433.(in Chinese)
[103] 韦炳耐,俸祥仁,何立贵,等.努比亚山羊人工授精繁殖效果试验[J].贵州畜牧兽医,2017,41(6):4-6. WEI B N,FENG X G,HE L G,et al.The effect of artificial insemination on reproductive performances in Nubian goats[J]. Guizhou Journal of Animal Husbandry & Veterinary Medicine,2017,41(6):4-6.(in Chinese)
[104] CHEN R,WAN R,PENG K,et al.Equol alleviates the in vitro aging-induced disruption of porcine oocytes[J]. Reproduction in Domestic Animals,2025,60(1):e70007.
[105] MIRO J.Ovarian ultrasonography in the mare[J].Reproduction in Domestic Animals,2012,47:30-33.
[106] MEIRA C,FERREIRA J,PAPA F,et al.Study of the estrous cycle in donkeys (Equus asinus) using ultrasonography and plasma progesterone concentrations[J].Biology of Reproduction,1995,52(monograph_series1):403-410.
[107] 王必强.安格斯母牛同期发情技术应用示范与推广[J].中国牛业科学,2018,44(1):81-83. WANG B Q.Demonstration and extension of application of Angus cow estrous synchronization technology[J].China Cattle Science,2018,44(1):81-83.(in Chinese)
[108] WOODS D C,TILLY J L.Revisiting claims of the continued absence of functional germline stem cells in adult ovaries[J].Stem Cells,2023,41(2):200-204.
[109] RICHARD S,ANDERSON N J,ZHOU Y,et al.Mouse primary follicles experience slow growth rates after activation and progressive increases that influence the duration of the primary follicle phase[J].Biology of Reproduction,2023,109(5):684-692.
[110] SARTORI R,FRICKE P M,FERREIRA J C,et al.Follicular deviation and acquisition of ovulatory capacity in bovine follicles[J]. Biology of Reproduction,2001,65(5):1403-1409.
[111] PEARODWONG P,TRETIPSKUL C,PANYATHONG R,et al.Factors influencing pre-ovulatory follicle diameter and weaning-to-ovulation interval in spontaneously ovulating sows in tropical environment[J].Reproduction in Domestic Animals,2020,55(12):1756-1763.
[112] 王欢,邹惠影,朱化彬,等.CRISPR/Cas9基因编辑技术在家畜育种新材料创制中的研究进展[J].畜牧兽医学报,2021,52(4):851-861. WANG H,ZOU H Y,ZHU H B,et al.Advances in evaluation of livestock breeding new materials by using the CRISPR/Cas9 gene editing technology[J].Acta Veterinaria et Zootechnica Sinica,2021,52(4):851-861.(in Chinese)
[113] 潘东霞,王辉,熊本海,等.CRISPR-Cas9基因编辑技术在牛、羊生产中的应用研究进展[J].中国畜牧兽医,2024,51(11):4880-4889. PAN D X,WANG H,XIONG B H,et al.Research progress on CRISPR-Cas9 gene editing technology in cattle and sheep production[J].China Animal Husbandry & Veterinary Medicine,2024,51(11):4880-4889.(in Chinese)
[114] 刘婧,郑积敏.CRISPR-Cas9基因编辑技术的应用与展望[J].化学教育(中英文),2025,46(4):1-8. LIU J,ZHENG J M.Application and prospect of CRISPR-Cas9[J].Chinese Journal of Chemical Education,2025,46(4):1-8.(in Chinese)
[115] HAN A,QAMAR A Y,BANG S,et al.Effect of extracellular vesicles derived from oviductal and uterine fluid on the development of porcine preimplantation embryos[J].Theriogenology,2025,234:216-224.
[116] JAWAD A,OH D,CHOI H,et al.Myo-inositol improves developmental competence and reduces oxidative stress in porcine parthenogenetic embryos[J].Frontiers in Veterinary Science,2024,11:1475329.
[117] 周荣,石俊松,周秀,等.猪卵丘包裹层数与猪卵母细胞质量的相关性研究[J].广东畜牧兽医科技,2014,39(5):27-29. ZHOU R,SHI J S,ZHOU X,et al.Relevance analysis between the layers of cumulus cells and the quality of porcine oocyte[J].Guangdong Journal of Animal and Veterinary Science,2014,39(5):27-29.(in Chinese)
[118] BELLIDO-QUISPE D K,ARCCE I M L,PINZÓN-OSORIO C A,et al.Chemical activation of mammalian oocytes and its application in camelid reproductive biotechnologies:A review[J]. Animal Reproduction Science,2024,266:107499.
[119] 曹云霞.高通量测序技术在生殖医学领域的研究与应用[J].安徽医科大学学报,2024,59(6):929-934. CAO Y X.Research and application of high-throughput sequencing technology in reproductive medicine[J].Acta Universitatis Medicinalis Anhui,2024,59(6):929-934.(in Chinese)
[120] ABDOON A S S,SOLIMAN S S,HUSSEIN N S,et al.Metabolomic profile of dromedary camel follicular fluid during the breeding and non-breeding seasons[J].Scientific Reports,2025,15(1):8923.
[121] ZHU Z,MIAO Z,CHEN H,et al.Ovarian transcriptomic analysis of Shanma ducks at peak and late stages of egg production[J]. Asian-Australasian Journal of Animal Sciences,2017,30(9):1215-1224.