中国畜禽种业 ›› 2026, Vol. 22 ›› Issue (9): 66-77.doi: 10.19543/j.cnki.1673-4556.20260812.003cstr: 32418.14.j.cnki.1673-4556.20260812.003

• 干细胞工程与种质资源创新 • 上一篇    下一篇

性腺类器官在家畜干细胞体外育种中的应用与展望

戈紫晴1(), 王佳悦2, 高盼银2, 赵璐2, 周婵2, 梁小滢2, 陈欣1, 朱帅鹏1, 王俊杰1, 程顺峰1, 沈伟1, 葛伟1()   

  1. 1. 青岛农业大学动物科技学院，山东 青岛 266109
    2. 青岛农业大学生命科学学院，山东 青岛 266109
  • 收稿日期:2026-06-18 出版日期:2026-09-26 发布日期:2026-09-10
  • 通讯作者: 葛伟
  • 作者简介:
    戈紫晴（2005—），女，山东济宁人，研究方向：动物遗传育种与繁殖，E-mail：。
  • 基金资助:
    国家自然科学基金(32570626)

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

Ziqing Ge1(), Jiayue Wang2, Panyin Gao2, Lu Zhao2, Chan Zhou2, Xiaoying Liang2, Xin Chen1, Shuaipeng Zhu1, Junjie Wang1, Shunfeng Cheng1, Wei Shen1, Wei Ge1()   

  1. 1. College of Animal Science and Technology, Qingdao Agricultural University, Qingdao, 266109, Shandong
    2. College of Life Sciences, Qingdao Agricultural University, Qingdao, 266109, Shandong
  • Received:2026-06-18 Online:2026-09-26 Published:2026-09-10
  • Contact: Wei Ge

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摘要：

传统大家畜育种进程受限于世代间隔长以及后代数量有限等问题，导致育种进程缓慢，短期内难以满足预期的育种需求，因此亟需开展新的育种技术路径。干细胞体外育种技术作为新兴动物育种范式，其核心在于将多能干细胞（Pluripotent stem cells，PSCs）在体外定向诱导为功能性配子（即体外配子发生，In vitro gametogenesis，IVG），结合高通量基因组选择（Genomic selection，GS），实现跨越个体世代的加速遗传改良，从根本上突破传统育种对活体动物繁殖周期的依赖。然而目前，家畜干细胞研究还存在诸多困难，例如家畜干细胞建系困难、干细胞诱导配子发生异常以及体外诱导过程中生殖细胞容易发生丢失等，这些问题严重限制了家畜干细胞育种技术的应用。此外，虽然小鼠上已经实现了完全体外条件下利用干细胞诱导获得功能性的配子，但是由于不同物种之间的差异性，小鼠上的诱导方案还无法直接应用到家畜研究中。针对上述瓶颈，性腺类器官作为一种能够模拟体内性腺微环境、辅助胚胎干细胞（Embryonic stem cells，ESCs）完成生殖细胞分化的三维培养体系，近年来在体外配子发生研究中得到了广泛应用，为攻克体外配子发生障碍提供了理想的研究平台。该三维共培养体系能够复刻生殖细胞发育所需的细胞互作与细胞因子梯度，填补二维单层培养体系无法重现性腺发育完整时序的短板。鉴于此，本文总结了近年来性腺类器官模型在干细胞育种技术中的应用及发展，以期为推动干细胞育种发展及未来家畜育种体系的变革提供理论基础。

关键词: 干细胞体外育种, 性腺类器官, 体外配子发生

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

中图分类号: 

  • S81

图1

干细胞体外育种的流程 注：图片使用Microsoft PowerPoint2021绘制，下图同。"

表1

猪、牛、羊PSCs的研究进展"

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

表2

干细胞体外分化配子获得存活后代的研究进展"

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

图2

性腺类器官的形成与功能"

表3

性腺类器官的构建与应用"

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
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