人工诱导贝类雄核发育的研究
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摘要
雄核发育(Androgenesis)是指后代的遗传物质完全来自父本的特殊的有性生殖方式。人工诱导雄核发育的原理是利用物理或化学方法等使卵子遗传失活，与正常精子受精，卵仅靠雄核发育成胚胎，而后通过抑制第一次卵裂使单倍体胚胎的染色体加倍发育成雄核二倍体个体。由于雄核发育后代的遗传物质完全来自父本，加倍后各基因位点均处于纯合状态，因而可以用于快速建立纯系，进行遗传分析。此外，雄核发育技术与精子冷藏技术相结合还可以成为物种保护的重要手段。γ射线、X射线和紫外线经常被用作失活卵子染色体的有效手段，其中紫外线更以其使用方便、安全廉价的优点被广泛应用。温度休克、静水压处理和化学药物诱导则是常见的加倍方法。在鱼类中，已有许多学者对失活卵子遗传物质和加倍雄核发育单倍体的有效措施进行了研究，并成功地诱导出雄核发育二倍体。然而人工诱导贝类雄核发育的研究在国内外还很少报道。本研究以栉孔扇贝(Chlamys farreri)和太平洋牡蛎(Crassostrea gigas)为研究对象，筛选出紫外线诱导雄核发育的适宜参数，并对诱导的受精细胞学过程进行了观察和探讨。
一、探讨了紫外线照射对栉孔扇贝和太平洋牡蛎卵子遗传失活的作用。用中心波长为254nm的不同紫外线剂量处理栉孔扇贝和太平洋牡蛎卵子，使其失去遗传活性，然后与正常精子授精，诱导雄核发育单倍体。受精率、D形幼虫发生率及染色体分布频率表明：在强度为2.8mW·cm~(-2)·s~(-1)的紫外线下分别照射20s和30s是获得栉孔扇贝和太平洋牡蛎雄核发育单倍体的适宜条件。研究发现受精率和D形幼虫发生率随照射时间的增加而下降，遗传失活的卵子与正常精子受精后其胚胎发育至D形幼虫前期停止。实验中各处理组均出现非整倍体。出现的原因可能由于紫外线照射剂量对卵子染色体遗传失活的作用程度不同以及光线对DNA的修复作用。
二、利用DAPI染色和荧光显微镜观察了栉孔扇贝、太平洋牡蛎正常卵子与雄核发育卵子在减数分裂、受精过程和卵裂早期中的核相变化。结果表明在两种贝类中，尽管紫外线照射并没有影响卵子的成熟分裂及雌性、雄性原核的形成，但使它们的发生过程滞后。在第1卵裂中期，雄核发育卵子中雌性原核并不像雄性原核一样形成染色体，而是形成1个浓缩的染色质小体(DCB)。第1卵裂后期，DCB不参与核分裂。第1卵裂结束时，DCB位于2个分裂球其中之一的细胞质内或在赤道板处被分割成
人工诱导贝类雄核发育的研究
两部分。实验结果提供了栉孔扇贝和太平洋牡蝠雄核发育的细胞学证据。
三、尝试了利用6一DMAP抑制第一卵裂以诱导栉孔扇贝雄核发育二倍体。结果
表明尽管部分单倍体胚胎的染色体能够加倍，却很难获得雄核发育二倍体D形幼虫。
相比较而言，当培养温度为19℃时，授精后80 min用浓度为60仁岁ml的6一DMAP
处理受精卵巧min诱导效果较好。细胞学观察表明，6一DMAP阻止了纺锤体的形成
和染色体的移动，导致一个融合的二倍性雄性原核的形成。
Androgenesis is defined as all-paternal inheritance. Viable androgenetic diploids can be generated by inhibiting the first cleavage to double paternally derived chromosomes, after fertilization of genetically inactivated eggs with normal sperm. It is a technique that could facilitate the production of completely homozygous isogenic lines, examine sex determination, make genetic analysis, and protect endangered species. Gamma, X-ray and ultraviolet irradiation inactivation are the usual methods used to inactivate egg nuclei. Among them, UV irradiation has been widely used for the convenient facility, safety and relatively low cost. In many fish species, efficient procedures of genetically inactivating of eggs and restoring diploidy have been studied and achievements of successful diploid androgenesis were reported. In contrast, in mollusks, artificial induction of androgenesis has rarely been studied. In this study, optimum inductive parameters in the scallop (Chlamys farreri), and the Pacific oyster (Crassos
trea gigas) were tested and cytological processes in induction were observed.
Effects of ultraviolet irradiation on genetic inactivation of eggs were examined in these two species. UV irradiation of eggs for 20 s and 30 s respectively at a UV (254 nm) intensity of 2.8 mW- cm"2- s"1 were optimum doses to achieve haploid androgenesis of the scallop and the Pacific oyster on the basis of observations on the fertilization and development rates and chromosome constitutions. The rates of the fertilization and the development of D-shaped larvae decreased with increasing irradiation time, and the development of the genetically inactivated eggs fertilized with the normal sperm terminated before reaching the D-shaped stage. Aneuploids were found in these studies. Because ultraviolet irradiation is known to cause pyrimidine dimerization in the DNA helix, which prevents the replication of genome, the occurrence of these aneuploids is probably attributed to different degree of maternal chromosomal inactivation by UV irradiation and/or DNA repair.
Nuclear changes in normal and androgenetic eggs of the above two species were examined under a fluorescence microscope during meiosis, fertilization and early
development. Cytological processes in induction in the two species were almost the same. Although UV irradiation did not effect either meiotic maturation or the formation of the male and female pronuclei, their developmental progresses were delayed. At metaphase of the first cleavage, the female pronucleus in UV-irradiated eggs inseminated with the normal sperm did not form chromosome, unlike the male pronucleus, but became a dense chromatin body (DCB), which did not participate in the karyokinesis at anaphase of the first cleavage. At completion of cytokinesis of the first cleavage, the DCB was seen either in the cytoplasm of one of the two blastomeres or on the equatorial plate as two partitional parts. Cytological evidence of the induction of androgenesis was demonstrated.
The induction of androgenesis diploids by suppression of the first cleavage in the scallop, C.farreri was attemped using the treatment of 6-dimethylaminopurine (6-DMAP). The result showed that it was difficult to induce diploid androgenetic D-shaped larvae. By comparison, treating the zygotes with 6-DMAP (60 ng/ml) at 80 min postinsemination for 15 min at water temperature of 19癈 produced more expected D-shaped larvae. According to Cytological observations, 6-DMAP disrupted the spindle at mitosis and inhibited chromosome movement, resulting in the formation of one diploid male nucleus.
一、探讨了紫外线照射对栉孔扇贝和太平洋牡蛎卵子遗传失活的作用。用中心波长为254nm的不同紫外线剂量处理栉孔扇贝和太平洋牡蛎卵子，使其失去遗传活性，然后与正常精子授精，诱导雄核发育单倍体。受精率、D形幼虫发生率及染色体分布频率表明：在强度为2.8mW·cm~(-2)·s~(-1)的紫外线下分别照射20s和30s是获得栉孔扇贝和太平洋牡蛎雄核发育单倍体的适宜条件。研究发现受精率和D形幼虫发生率随照射时间的增加而下降，遗传失活的卵子与正常精子受精后其胚胎发育至D形幼虫前期停止。实验中各处理组均出现非整倍体。出现的原因可能由于紫外线照射剂量对卵子染色体遗传失活的作用程度不同以及光线对DNA的修复作用。
二、利用DAPI染色和荧光显微镜观察了栉孔扇贝、太平洋牡蛎正常卵子与雄核发育卵子在减数分裂、受精过程和卵裂早期中的核相变化。结果表明在两种贝类中，尽管紫外线照射并没有影响卵子的成熟分裂及雌性、雄性原核的形成，但使它们的发生过程滞后。在第1卵裂中期，雄核发育卵子中雌性原核并不像雄性原核一样形成染色体，而是形成1个浓缩的染色质小体(DCB)。第1卵裂后期，DCB不参与核分裂。第1卵裂结束时，DCB位于2个分裂球其中之一的细胞质内或在赤道板处被分割成
人工诱导贝类雄核发育的研究
两部分。实验结果提供了栉孔扇贝和太平洋牡蝠雄核发育的细胞学证据。
三、尝试了利用6一DMAP抑制第一卵裂以诱导栉孔扇贝雄核发育二倍体。结果
表明尽管部分单倍体胚胎的染色体能够加倍，却很难获得雄核发育二倍体D形幼虫。
相比较而言，当培养温度为19℃时，授精后80 min用浓度为60仁岁ml的6一DMAP
处理受精卵巧min诱导效果较好。细胞学观察表明，6一DMAP阻止了纺锤体的形成
和染色体的移动，导致一个融合的二倍性雄性原核的形成。
Androgenesis is defined as all-paternal inheritance. Viable androgenetic diploids can be generated by inhibiting the first cleavage to double paternally derived chromosomes, after fertilization of genetically inactivated eggs with normal sperm. It is a technique that could facilitate the production of completely homozygous isogenic lines, examine sex determination, make genetic analysis, and protect endangered species. Gamma, X-ray and ultraviolet irradiation inactivation are the usual methods used to inactivate egg nuclei. Among them, UV irradiation has been widely used for the convenient facility, safety and relatively low cost. In many fish species, efficient procedures of genetically inactivating of eggs and restoring diploidy have been studied and achievements of successful diploid androgenesis were reported. In contrast, in mollusks, artificial induction of androgenesis has rarely been studied. In this study, optimum inductive parameters in the scallop (Chlamys farreri), and the Pacific oyster (Crassos
trea gigas) were tested and cytological processes in induction were observed.
Effects of ultraviolet irradiation on genetic inactivation of eggs were examined in these two species. UV irradiation of eggs for 20 s and 30 s respectively at a UV (254 nm) intensity of 2.8 mW- cm"2- s"1 were optimum doses to achieve haploid androgenesis of the scallop and the Pacific oyster on the basis of observations on the fertilization and development rates and chromosome constitutions. The rates of the fertilization and the development of D-shaped larvae decreased with increasing irradiation time, and the development of the genetically inactivated eggs fertilized with the normal sperm terminated before reaching the D-shaped stage. Aneuploids were found in these studies. Because ultraviolet irradiation is known to cause pyrimidine dimerization in the DNA helix, which prevents the replication of genome, the occurrence of these aneuploids is probably attributed to different degree of maternal chromosomal inactivation by UV irradiation and/or DNA repair.
Nuclear changes in normal and androgenetic eggs of the above two species were examined under a fluorescence microscope during meiosis, fertilization and early
development. Cytological processes in induction in the two species were almost the same. Although UV irradiation did not effect either meiotic maturation or the formation of the male and female pronuclei, their developmental progresses were delayed. At metaphase of the first cleavage, the female pronucleus in UV-irradiated eggs inseminated with the normal sperm did not form chromosome, unlike the male pronucleus, but became a dense chromatin body (DCB), which did not participate in the karyokinesis at anaphase of the first cleavage. At completion of cytokinesis of the first cleavage, the DCB was seen either in the cytoplasm of one of the two blastomeres or on the equatorial plate as two partitional parts. Cytological evidence of the induction of androgenesis was demonstrated.
The induction of androgenesis diploids by suppression of the first cleavage in the scallop, C.farreri was attemped using the treatment of 6-dimethylaminopurine (6-DMAP). The result showed that it was difficult to induce diploid androgenetic D-shaped larvae. By comparison, treating the zygotes with 6-DMAP (60 ng/ml) at 80 min postinsemination for 15 min at water temperature of 19癈 produced more expected D-shaped larvae. According to Cytological observations, 6-DMAP disrupted the spindle at mitosis and inhibited chromosome movement, resulting in the formation of one diploid male nucleus.
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