性腺类器官在家畜干细胞体外育种中的应用与展望

doi:10.19543/j.cnki.1673-4556.20260812.003

Abstract

Abstract:

Conventional livestock breeding is constrained by long generation intervals and limited progeny quantity, resulting in delayed breeding progress that fails to satisfy expected breeding requirements. Therefore, novel breeding technical approaches are urgently required. As an emerging paradigm for animal breeding, the core of in vitro stem cell-based breeding technology lies in the targeted induction of pluripotent stem cells into functional gametes under in vitro conditions. Combined with high-throughput genomic selection, this technology will facilitate accelerated genetic improvement across generations, fundamentally breaking the dependence on the reproductive cycle of live animals in traditional breeding. Nevertheless, multiple bottlenecks remain in the research of livestock stem cells, including difficulties in establishing stable livestock stem cell lines, aberrant gametogenesis induced from stem cells, and the loss of germ cells during in vitro induction. These obstacles greatly restrict the practical application of stem cell-based breeding technologies in livestock industry. Currently, although functional gametes can be differentiated from pluripotent stem cells using mice as a research model, the induction procedure cannot be directly applied to livestock research due to species-specific differences. To address this bottlenecks, gonadal organoids, a three-dimensional culture system capable of mimicking the in vivo gonadal microenvironment and facilitating germ cell differentiation derived from embryonic stem cells, have been widely adopted in researches concerning in vitro gametogenesis in recent years. It serves as an ideal research platform to overcome the technical barriers restricting in vitro gametogenesis.This three-dimensional co-culture system can recapitulate the cell-cell interactions and cytokine gradients required for germ cell development, addressing the limitation that two-dimensional monolayer culture systems fail to reconstruct the full temporal sequence of gonadal development. Collectively, this paper here summarizes the application and progress of gonadal organoid models in stem cell breeding technologies, providing a theoretical basis for advancing stem cell-mediated breeding and the transformation of future livestock breeding systems.

Key words: In vitro stem cell-based breeding, Gonadal organoids, In vitro gametogenesis

CLC Number:

Ziqing Ge, Jiayue Wang, Panyin Gao, Lu Zhao, Chan Zhou, Xiaoying Liang, Xin Chen, Shuaipeng Zhu, Junjie Wang, Shunfeng Cheng, Wei Shen, Wei Ge. Gonadal organoids in livestock stem cell-based in vitro breeding: Applications and perspectives[J].Chinese Livestock and Poultry Breeding, 2026, 22(9): 66-77.

Figures/Tables 5

Fig. 1

Table 1

Research progress of porcine, bovine and ovine PSCs"

家畜Livestock	年份 Year	作者 Author	方法 Method	发现 Finding	问题 Limitation
猪Porcine	2009	Ezashi等^[13]、Wu等^[14]和Esteban等^[15]	病毒载体的体细胞重编程技术	得到了3种iPSCs	多能性完全依赖外源因子的持续表达，无法完全重编程
	2022	Zhi等^[16]	“3i/LAF”培养基	从E8～E10胚胎中建立猪EpiSCs	依然不具备真正的原始态多能性
	2024	Xiang等^[17]	培养基4FIXY衍生ESCs与三维两步分化	从ESCs生成猪类囊胚	无法像真实囊胚一样，完成后续的着床与妊娠发育
	2024	Choi等^[18]	无血清条件下开发了一种由FGF2、IWR-1和WH-4-023组成的简化培养基	抑制WNT和SRC能够建立单细胞传代的多能性干细胞，可以在无血清条件下实现无饲养层扩增。	需要依赖商用外源耗材，同时还缺失体内发育功能验证
	2025	Wang等^[19]	WNT小分子抑制剂的化学培养条件操控法	发现WNT抑制剂能够维持猪ESCs的多能性，抑制ESCs转化为XEN细胞	没有形成具有种系嵌合体和传代特性的高质量猪胚胎干细胞
	2026	Shi等^[20]	新型复合培养基Cocktail LACID	建立了一套能稳定获得高质量猪诱导多能干细胞的改良方案	难以形成嵌合胚胎，无法通过核移植发育成囊胚
牛 Bovine	2011	Han等^[21]	逆转录病毒作为载体	成功建立了牛iPSC样细胞	多能性完全依赖外源转基因的持续表达
	2018	Bogliotti等^[22]	含FGF2和WNT抑制剂的培养体系	成功建立稳定Primed型的牛ESCs	研究未进行胚胎移植，无法验证其能否发育获得完整个体
	2021	Soto等^[23]	优化了该培养体系	简化了牛ESCs的构建流程	依然无法突破处于Primed多能状态的限制
	2024	Shirasawa等^[24]	革新了植入前胚胎和ESCs多能性培养技术	成功开发出一种从新构建的牛ESCs中稳定诱导PGCLCs的新方法	尚未被证明具备发育为功能性配子的能力
	2024	Zhi等^[25]	利用“3i/LAF”培养条件	成功构建了牛EpiSCs培养系统	尚未实现由EpiSCs体外诱导获得具备受精能力的功能性配子
	2024	Smith等^[26]	构建牛孤雄以及孤雌单倍体胚胎干细胞体系，筛选体外优良基因组	实现了雌雄单倍体细胞组合定向培育成指定基因组的牛后代	规避减数分裂基因随机重组，未实现足月妊娠
	2025	Yang等^[27]	将新开发的“鱼精蛋白辅助的单倍体胚胎干细胞胞质内注射技术”与先进的基因编辑方法相结合	首次成功建立了牛和羊的单倍体胚胎干细胞系，成功培育出存活以及正常生长繁殖的牛的基因修饰个体	着床后发育仍存在明显障碍，其发育稳定性和规模化应用效率仍有待提高
羊 Ovine	2020	Malik等^[28]	孤雌激活观察到了类似体内胚胎发育的过程	进一步证实了ESCs衍生的卵母细胞样细胞具有全能性	无法证明诱导产生的卵母细胞样细胞具备功能性配子的完整发育潜能
	2025	Saadeldin等^[29]	结合无饲养层和子宫内膜类器官培养基培养	实现胚胎体外发育至受精后第20 d	仅完成体外20 d胚胎培养的形态与分子表征，缺乏体内移植验证
	2025	Wang等^[30]	导入多能转录因子，将黑骨绵羊成纤维细胞重新组合而获得iPSCs	发现该细胞具备完整三胚层分化潜能，同时可以作为核供体提升体细胞核移植胚胎卵裂与囊胚发育效率	必须依赖外源转基因才能维持多能性，无法制备无转基因干细胞

Table 1

Table 2

Research progress on viable offspring generated from in vitro differentiated stem cell-derived gametes"

类型 Type	年份Year	作者 Author	干细胞类型 Stem cell type	诱导分化方式 Induction and differentiation method
利用干细胞诱导分化精子 In vitro derivation of sperm from stem cells	2011	Hayashi等^[33]	ESCs、iPSCs	将ESCs和iPSCs诱导EpiLCs，之后通过细胞因子，将EpiLCs诱导为PGCLCs
	2016	Zhou等^[34]	ESCs	完全体外条件下完成减数分裂，分化为功能性圆形精子细胞，且使用胞质内单精子注射
	2016	Ishikura等^[35]	ESCs	将ESCs诱导为PGCLCs，后与胚胎睾丸体细胞共培养，成功分化为精原细胞样细胞，并进一步扩增获得具有SSCs活性的生殖干细胞样细胞
	2021	Ishikura等^[36]	ESCs	将小鼠ESCs诱导为PGCLCs，该类细胞完成表观遗传重编程后，能够在重构睾丸中分化为精原细胞样细胞并扩增为GSCLCs，在体内移植或离体培养后通过ICSI获得后代
	2022	Oikawa等^[37]	ESCs	大鼠ESCs诱导为上胚层样细胞后，通过3D技术分化为PGCLCs，移植入无生殖系小鼠睾丸完成精子发生，经ICSI获得后代
	2025	Li等^[38]	ESCs	利用转录因子在体外诱导小鼠PGCLCs，与睾丸体细胞共培养后经ROSI获得后代
	2026	Yoshino等^[39]	ESCs、iPSCs	将小鼠ESCs/iPSCs重构为睾丸体细胞样细胞，之后组装形成睾丸类器官，使PGCLCs在该类器官中分化为GSCLCs，移植后产生成熟精子并经ICSI获得后代
利用干细胞诱导分化卵子 In vitro derivation of oocytes from stem cells	2012	Hayashi等^[40]	ESCs、iPSCs	将ESCs与iPSCs诱导分化为PGCLCs，之后与雌性胚胎性腺体细胞混合聚集构建重构卵巢，共培养过程中完成X染色体激活、基因组印记擦除并启动减数分裂相关程序，移植小鼠体内获得卵泡期卵母细胞
	2016	Hikabe等^[41]	ESCs、iPSCs	先将小鼠ESCs/iPSCs诱导为PGCLCs，再经重构卵巢培养、体外成熟后受精获后代
	2021	Yoshino等^[42]	ESCs	将ESCs诱导分化为PGCLCs，并与胎儿卵巢体细胞样细胞共聚集来组装体外重构卵巢卵泡结构，胎儿卵巢体细胞样细胞作为颗粒细胞支持PGCLCs完成减数分裂并发育为功能卵母细胞
	2023	Murakami等^[43]	iPSCs	将小鼠iPSCs中XY染色体转换为XX染色体，再诱导分化为功能性卵母细胞，受精产生小鼠后代

Table 2

Fig. 2

Table 3

Construction and application of gonadal organoids"

性腺 Gonad	物种 Species	年份 Year	作者 Author	培养技术 Culture technique	应用 Application
睾丸 Testis	小鼠	2011	Sato等^[47]	气-液平面培养法	实现了具有受精功能的小鼠精子体外生成
		2017	Alves-Lopes等^[48]	三层梯度系统	成功构建了睾丸类器官
		2019	Sakib等^[49]	微孔培养法	构建出具有器官型结构的睾丸类器官
		2020	Majidi-Gharenaz等^[50]	睾丸脱细胞支架	小鼠精原干细胞进行三维培养
		2021	Richer等^[51]	三维打印技术	实现了早期生殖细胞的长期维持及减数分裂启动
		2024	Stopel等^[52]	气-液平面培养法	构建可复刻体内睾丸生精小管样结构的小鼠胚胎期类器官
		2026	Yoshino等^[39]	三维共培养体系	重构了睾丸的发育全过程，复刻了体内性别决定机制，产生了形成生精小管和相邻间质组织的细胞类型
	家畜	2019	Sakib等^[53]	微孔培养法	构建出具有睾丸特异结构的猪睾丸类器官
		2021	Cham等^[54]	气-液平面培养法	建出包含血管结构、高度模拟天然未成熟睾丸组织的高仿生类器官
		2022	Cortez等^[55]	三维自组装技术	成功诱导组装出牛睾丸类器官
		2024	Cortez等^[56]	三维自组装技术	优化类器官三维形态与结构稳定性
		2025	Elsenhans等^[57]	微孔聚集培养技术	开发了一套简便、可重复的标准化睾丸类器官批量制备体系
卵巢 Ovary	小鼠	2016	Morohaku等^[58]	气-液平面培养法	优化卵泡发育微环境，实现冻存性腺组织分化获得成熟卵母细胞
		2017	Laronda等^[59]	三维打印技术	调控支架孔隙的前进角优化卵泡-支架，显著提升了卵泡的存活率与发育潜能
		2021	Li等^[60]	三维培养技术	模拟了体内卵巢的结构与功能，可支持生殖细胞完成减数分裂并生成功能性卵母细胞
		2024	Dipali等^[61]	三维培养技术	自主开发出一种可自发组装的新型卵巢体细胞类器官模型
		2025	Zhao等^[62]	3D-GCs体系	体外培养小鼠未成熟的生发泡期卵母细胞
		2026	Kang等^[63]	三维打印技术	精准制备出动态三维体外培养平台
	家畜	2025	Gómez-Álvarez等^[64]	机械-酶法	利用牛卵巢表面上皮细胞成功构建了牛卵巢类器官

Table 3

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Gonadal organoids in livestock stem cell-based in vitro breeding: Applications and perspectives

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