生物多样性 ›› 2019, Vol. 27 ›› Issue (11): 1236-1244.doi: 10.17520/biods.2019160

• 研究报告 • 上一篇    下一篇

普通针毛蕨颈卵器和卵的发育

詹臻, 张剑锋, 曹建国, 戴锡玲()   

  1. 上海师范大学生命科学学院, 上海 200234
  • 收稿日期:2019-05-10 接受日期:2019-08-12 出版日期:2019-11-20
  • 通讯作者: 戴锡玲 E-mail:daixiling2010@shnu.edu.cn
  • 基金项目:
    上海植物种质资源工程技术研究中心项目(17DZ2252700)

Development of archegonium and oogenesis of the fern Macrothelypteris torresiana

Zhen Zhan, Jianfeng Zhang, Jianguo Cao, Xiling Dai()   

  1. College of Life Sciences, Shanghai Normal University, Shanghai 200234
  • Received:2019-05-10 Accepted:2019-08-12 Online:2019-11-20
  • Contact: Dai Xiling E-mail:daixiling2010@shnu.edu.cn

蕨类植物卵发生是有性生殖研究的重要内容。真水龙骨类II的卵发生尚无人研究, 本文对该类群中的普通针毛蕨(Macrothelypteris torresiana)卵发生过程进行了光镜和透射电镜研究。结果显示: 卵细胞刚形成时, 与腹沟细胞紧密相连, 随着卵进一步发育, 卵细胞和腹沟细胞之间逐渐形成分离腔, 但孔区处卵细胞和腹沟细胞始终相连。随后, 卵细胞上表面有不定型物质堆积在质膜外, 形成一层加厚的卵膜, 孔区处没有卵膜覆盖的位置最后形成受精孔。在卵发育后期, 卵细胞核变得不规则, 近成熟时卵核产生大量核外突。卵发育过程中卵膜的出现、受精孔的产生以及核外突等特征, 与进化的真水龙骨类中其他蕨类植物的卵发生研究结果相似, 与薄囊蕨类中原始基部类群的卵发生现象有显著差别。由此可见, 普通针毛蕨属于进化类型。依据卵发生中卵膜和受精孔等特征推测原始薄囊蕨类经过里白类、桫椤类, 最终演化为水龙骨类。

关键词: 蕨类植物, 颈卵器, 卵发生, 超微结构, 普通针毛蕨

Fern oogenesis is a vital process in sexual reproduction, yet Polypodia II oogenesis has not been studied. This paper studies sexual reproduction oogenesis of Macrothelypteris torresiana of Polypodiaceae. Our findings document oogenesis and egg movement throughout sexual reproduction. The appearance of egg envelope, the formation of fertilization pore and nuclear evaginations during the development of Macrothelypteris torresiana are similar to other pteridophytes in eupolypods, and are significantly different from those of the primitive basal groups in leptosporangiates. This indicates that Macrothelypteris torresiana may be a evolutionary fern. Based on the characteristics of the egg envelope and the fertilization pore, we infer that primitive leptosporangiates evolved into polypods after passing through gleicheniales and cyatheales.

Key words: ferns, archegonium, oogenesis, ultrastructure, Macrothelypteris torresiana

图1

普通针毛蕨颈卵器发育的光学显微镜图。 (A-B): 初生细胞(PC); (C): 中央细胞(CC)和单核颈沟细胞(MNC); (D-E): 幼卵细胞(EC)、腹沟细胞(VCC)与双核颈沟细胞(NCC); (F): 卵细胞发育早期; (G): 卵细胞(EC)与腹沟细胞(VCC)之间分离腔(SC)逐渐形成; (H): 卵细胞与腹沟细胞之间完全分离; (I): 受精孔(FP)形成; (J): 成熟卵时期的颈卵器; (K): 颈卵器壁细胞向基部弯曲; (L): 成熟卵后期, 双核颈沟细胞、腹沟细胞完全退化。FP: 受精孔; SC: 分离腔。"

图2

普通针毛蕨颈卵器发育和卵发生的电镜图。 (A): 中央细胞(CC)和单核颈沟细胞(MNC); (B): 单核颈沟细胞(MNC)放大及其细胞器; (C): 双核颈沟细胞(NCC)与中央细胞(CC); (D): 幼卵阶段, 颈卵器颈部4-5层细胞高; (E): 腹沟细胞(VCC)呈凸透镜状; (F): 腹沟细胞(VCC)和卵细胞(EC)内细胞器发达。M: 线粒体; N: 核; P: 质体; Ve: 囊泡; Pd: 胞间连丝。"

图3

普通针毛蕨卵发生电镜图。 (A): 腹沟细胞(VCC)与双核颈沟细胞(NCC)和卵细胞(EC)之间有胞间连丝相连; (B): 卵细胞细胞核规则, 腹沟细胞的细胞核位于中央; (C): 双核颈沟细胞完全退化, 细胞内出现大量不定型物质; (D): 卵细胞细胞核(N)变形, 卵膜(EE)开始堆积; (E): 卵膜下方线粒体、质体发达; (F): 卵细胞内出现圆形内质网(ER)和嗜锇性囊泡(Veo)。M: 线粒体; N: 核; P: 质体; Ve: 囊泡; ER: 内质网; SC: 分离腔。"

图4

普通针毛蕨卵发生电镜图。 (A): 孔区与腹沟细胞(VCC)分离, 出现受精孔(FP); (B): 卵细胞弓形; (C): 腹沟细胞(VCC)和双核颈沟细胞(NCC)完全退化; (D): 成熟卵细胞内的细胞器; (E): 卵细胞(EC)侧面; (F): 卵细胞与周围颈卵器壁细胞分离。M: 线粒体; N: 核; P: 质体; SC: 分离腔; FP: 受精孔; EE: 卵膜。"

图5

真水龙骨类卵发生可能的演化路径。 (A): 真水龙骨类I, 卵发育后期形成卵膜和受精孔(侧方), 卵核不规则且产生大量核外突; (B): 真水龙骨类II, 卵发育后期形成卵膜和受精孔(侧方), 卵核不规则且产生大量核外突; (C): 桫椤类, 卵发育后期形成卵膜和受精孔(中央), 卵核不规则且产生大量核外突; (D): 里白类, 卵发育后期形成卵膜和受精孔(中央), 卵核外产生少量核外突; (E): 原始薄囊蕨类, 卵发育后期无卵膜、受精孔, 卵核规则且不产生核外突。VCC: 腹沟细胞; EC: 卵细胞; N: 核; SC: 分离腔; FP: 受精孔; EE: 卵膜。"

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