棉花双隐性核雄性不育系花药及核辐射突变体雄性不育花药的细胞学研究
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摘要
棉花是重要的经济作物,在国民经济中占重要地位,具有较明显的杂种优势。杂种优势的利用是提高棉花产量与品质的重要措施之一。雄性不育是植物界普遍存在的现象,高等植物的雄性不育是杂种优势利用的基础。棉花的雄性不育主要分为两种主要类型：质核互作雄性不育和核雄性不育。质核互作雄性不育由于恢复系的局限性,在生产上应用较少。相比之下,核不育系已有较大应用规模。陆地棉双隐性核雄性不育系ms5ms6就是其中之一。虽然目前已利用该材料培育出多个杂交棉品种,但对其败育的细胞学研究却很少,由于棉花雄性不育败育的时期很长,且不局限于某一特定时期,败育特征也因材料而异。因此为了系统地研究该材料的败育时期及特征,本文利用石蜡切片技术和透射电镜技术,对其研究,取得的主要实验结果如下：
不育株花药的败育期是在花粉母细胞时期和小孢子时期,在造孢时期并无异常。在花粉母细胞时期约有20%的败育,其它都是在小孢子时期败育的,这就说明不育花药大多数能通过减数分裂时期。败育的主要特征是：在花粉母细胞时期,有少量的核仁穿壁现象,还有部分小孢子母细胞出现半月形。在小孢子时期,出现了小孢子粘连及少数五分体等不正常现象,小孢子都没有刺突长出,最后解体、退化。
陆地棉双隐性核雄性不育系mS5mS6的主要败育时期是在小孢子阶段,为了更进一步了解该材料更深层次的败育原因,采用透射电镜对其四分体时期及刚长出刺突的小孢子早期花药进行观察,发现败育花药在四分体时期,胼胝质未溶解,小孢子就已经开始收缩,且外璧加厚,而正常小孢子是在胼胝质溶解后,小孢子释放后才开始收缩的；在小孢子时期,小孢子只有一层内壁且萌发孔结构异常。绒毡层细胞在四分体时期就有降解,小孢子时期降解不同步。在整个花药发育过程中,不管是在小孢子还是绒毡层细胞中,内质网都缺乏及异常,脂肪积累少,这可能是导致小孢子败育最重要的原因。
核辐射也是创造雄性不育突变体的重要方式之一,本文利用20Gy的60Co-Y射线对陆地棉品种苏棉22号的花粉进行辐射处理,花粉给雌蕊授粉,并收获种子。从M1群体中选取三个雄性不育突变体,采用石蜡切片技术对花药发育过程进行研究.通过与对照比较,三个突变体各个时期的败育特征有一定差异,但它们的主要败育特征是一致的：从小孢子母细胞时期至小孢子发育阶段都有败育,小孢子母细胞、绒毡层和中层细胞以及花药的形态都受到不同程度的影响,包括小孢子母细胞有微核及双核现象、减数分裂阶段小孢子母细胞细胞质膨胀、绒毡层及中层细胞异常等。总体来说,γ射线辐射花粉授粉后形成的突变体在小孢子的整个发育过程都受到影响,从而产生各种败育现象。
Cotton is an important economic crop which has the obvious heterosis, play an important role in the national economy.The use of heterosis is one of the important measures to improve the quality and production of cotton. Male sterility is a widespread phenomenon in plants,which is also the basis of heterosis in higher plants. There are two main types in cotton:cytoplasmic-nuclear male sterility and nuclear male sterility. Because the limitations of cytoplasmic male sterility, the application in the production is limited. In contrast, application of nuclear male sterile lines have a larger scale, Upland cotton double-recessive nuclear male sterile line ms5ms6 is one of them. Although there were a number of hybrid cotton varieties has been produced using this material, but its study in cytology of abortion is rarely. Because there are long abortion period and not limited to a specific period and abortion characteristics vary according to the material of cotton, therefore, the author of this article using paraffin slice technique under light microscope and transmission electron microscopy to study the materials abortion periods and characteristics,the main experimental results as follows:
The result of embryology study shew the abortion stages were the development periods of pollen mother cells and microspore,and there was no abnormal in sporogenous period. There were 20% abortion in pollen mother cell and other abortion periods was all in microspore,those shows that the majority of the sterile anther can pass meiosis. The abortion characters of the double recessiveness nuclear male-sterile line ms5ms6 of upland cotton were:low frequency of nucleolus penetration through nucleus envelop and cell wall and part of pollen mother cells appeared in the shape of lunule; microspore adhesion and few of pentads were found during the periods of microspore; there were no spinescent protuberance on the pollen wall, finally the microspore appeared abnormal and breakdown; The anther of double recessiveness nuclear male-sterile line ms5ms6 of upland cotton.
As the microstructure found that the major abortion period of double recessiveness nuclear male-sterile line mssms6 of upland cotton was in microspore stage, in order to further understand the causes of abortion, the anther of double recessiveness nuclear male-sterile line ms5ms6 of upland cotton (Gossypium hirsutumL.)in periods of tetrad and microspores which just has small spinescent protuberance was obseverd using transmission electron microscopy. The results shew that the abortion microspore shrink without corpus callosum solution during the period of tetrad stage while fertile microspore shrink after corpus callosum solution, male-sterile microspore has only one layers and abnormal aperture during the periods of microspore,Tapetum cells begin breakdown in period of tetrad, whether in the microspores or tapetum cells the lack and abnormal of endoplasmic reticulum and less of fat accumulation throughout the course of anther development, which may be the most important reason of microspore abortion.
Nuclear radiation is one of the important ways to create male-sterile mutant. The pollen of Sumian22 was irradiated by 60Co-y ray of 20Gy then fertilized to pistil and harvest its seed. Three male-sterile mutants were selected from M1 group.their anther researched by paraffin slice technique. Although there are certain different characteristics during the abortion anther development period, the main characteristics of abortion are consistent:the abortion stage from the development periods of pollen mother cells to microspore. pollen mother cells、tapetum、middle layer cells and the shape of anther were affected by the irradiation,the result contain micronucleus and double nucleus、cytoplasm expansion during the period of meiosis、tapetum and middle layer cells are abnormal.and so on. The microspores of mutants in the entire development process are affected and resulted in a variety of abortion phenomenon.
不育株花药的败育期是在花粉母细胞时期和小孢子时期,在造孢时期并无异常。在花粉母细胞时期约有20%的败育,其它都是在小孢子时期败育的,这就说明不育花药大多数能通过减数分裂时期。败育的主要特征是：在花粉母细胞时期,有少量的核仁穿壁现象,还有部分小孢子母细胞出现半月形。在小孢子时期,出现了小孢子粘连及少数五分体等不正常现象,小孢子都没有刺突长出,最后解体、退化。
陆地棉双隐性核雄性不育系mS5mS6的主要败育时期是在小孢子阶段,为了更进一步了解该材料更深层次的败育原因,采用透射电镜对其四分体时期及刚长出刺突的小孢子早期花药进行观察,发现败育花药在四分体时期,胼胝质未溶解,小孢子就已经开始收缩,且外璧加厚,而正常小孢子是在胼胝质溶解后,小孢子释放后才开始收缩的；在小孢子时期,小孢子只有一层内壁且萌发孔结构异常。绒毡层细胞在四分体时期就有降解,小孢子时期降解不同步。在整个花药发育过程中,不管是在小孢子还是绒毡层细胞中,内质网都缺乏及异常,脂肪积累少,这可能是导致小孢子败育最重要的原因。
核辐射也是创造雄性不育突变体的重要方式之一,本文利用20Gy的60Co-Y射线对陆地棉品种苏棉22号的花粉进行辐射处理,花粉给雌蕊授粉,并收获种子。从M1群体中选取三个雄性不育突变体,采用石蜡切片技术对花药发育过程进行研究.通过与对照比较,三个突变体各个时期的败育特征有一定差异,但它们的主要败育特征是一致的：从小孢子母细胞时期至小孢子发育阶段都有败育,小孢子母细胞、绒毡层和中层细胞以及花药的形态都受到不同程度的影响,包括小孢子母细胞有微核及双核现象、减数分裂阶段小孢子母细胞细胞质膨胀、绒毡层及中层细胞异常等。总体来说,γ射线辐射花粉授粉后形成的突变体在小孢子的整个发育过程都受到影响,从而产生各种败育现象。
Cotton is an important economic crop which has the obvious heterosis, play an important role in the national economy.The use of heterosis is one of the important measures to improve the quality and production of cotton. Male sterility is a widespread phenomenon in plants,which is also the basis of heterosis in higher plants. There are two main types in cotton:cytoplasmic-nuclear male sterility and nuclear male sterility. Because the limitations of cytoplasmic male sterility, the application in the production is limited. In contrast, application of nuclear male sterile lines have a larger scale, Upland cotton double-recessive nuclear male sterile line ms5ms6 is one of them. Although there were a number of hybrid cotton varieties has been produced using this material, but its study in cytology of abortion is rarely. Because there are long abortion period and not limited to a specific period and abortion characteristics vary according to the material of cotton, therefore, the author of this article using paraffin slice technique under light microscope and transmission electron microscopy to study the materials abortion periods and characteristics,the main experimental results as follows:
The result of embryology study shew the abortion stages were the development periods of pollen mother cells and microspore,and there was no abnormal in sporogenous period. There were 20% abortion in pollen mother cell and other abortion periods was all in microspore,those shows that the majority of the sterile anther can pass meiosis. The abortion characters of the double recessiveness nuclear male-sterile line ms5ms6 of upland cotton were:low frequency of nucleolus penetration through nucleus envelop and cell wall and part of pollen mother cells appeared in the shape of lunule; microspore adhesion and few of pentads were found during the periods of microspore; there were no spinescent protuberance on the pollen wall, finally the microspore appeared abnormal and breakdown; The anther of double recessiveness nuclear male-sterile line ms5ms6 of upland cotton.
As the microstructure found that the major abortion period of double recessiveness nuclear male-sterile line mssms6 of upland cotton was in microspore stage, in order to further understand the causes of abortion, the anther of double recessiveness nuclear male-sterile line ms5ms6 of upland cotton (Gossypium hirsutumL.)in periods of tetrad and microspores which just has small spinescent protuberance was obseverd using transmission electron microscopy. The results shew that the abortion microspore shrink without corpus callosum solution during the period of tetrad stage while fertile microspore shrink after corpus callosum solution, male-sterile microspore has only one layers and abnormal aperture during the periods of microspore,Tapetum cells begin breakdown in period of tetrad, whether in the microspores or tapetum cells the lack and abnormal of endoplasmic reticulum and less of fat accumulation throughout the course of anther development, which may be the most important reason of microspore abortion.
Nuclear radiation is one of the important ways to create male-sterile mutant. The pollen of Sumian22 was irradiated by 60Co-y ray of 20Gy then fertilized to pistil and harvest its seed. Three male-sterile mutants were selected from M1 group.their anther researched by paraffin slice technique. Although there are certain different characteristics during the abortion anther development period, the main characteristics of abortion are consistent:the abortion stage from the development periods of pollen mother cells to microspore. pollen mother cells、tapetum、middle layer cells and the shape of anther were affected by the irradiation,the result contain micronucleus and double nucleus、cytoplasm expansion during the period of meiosis、tapetum and middle layer cells are abnormal.and so on. The microspores of mutants in the entire development process are affected and resulted in a variety of abortion phenomenon.
引文
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