Microsporogenesis, megasporogensis and development of male and female gametophytes of Torreya jiulongshanensis, a critically endangered plant
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摘要:
目的 九龙山榧Torreya jiulongshanensis为浙江特有的古老孑遗植物，数量极少，为国家Ⅱ级重点保护野生植物，也被列为浙江省极小种群保护植物。系统观测九龙山榧大、小孢子的发生和雌、雄配子体的发育过程，有助于发现其配子是否发育异常及导致结籽率低下的原因。 方法 以九龙山榧模式产地的雌、雄植株为研究对象，通过定期观测和取样，采用石蜡切片法对其大、小孢子的发生，雌、雄配子体的发育过程进行切片、染色和观察。 结果 ①小孢子叶球于8月中旬分化，8月下旬分化形成次生造孢细胞，11月中旬形成小孢子母细胞，翌年2月下旬进入减数分裂期，3月中旬形成游离小孢子细胞，于3月末逐渐成熟；成熟花粉粒为双核，无气囊。小孢子囊的发育类型为基本型，孢质分裂方式为同时型，四分体有四面体形和左右对称形。②雄配子体从3月中下旬开始发生，到7月底形成精子，整个过程历时约4个月。③翌年3月下旬，大孢子叶球每个苞叶的叶腋基部直立着生2枚胚珠；大孢子母细胞于4月下旬经过减数分裂Ⅰ和减数分裂Ⅱ形成4个直列大孢子，5月中旬珠孔端的3个大孢子退化，合点端的大孢子最终发育成为雌配子体。④九龙山榧有2个单生型椭圆形的颈卵器，从传粉到受精历时约7个月。 结论 九龙山榧雌雄生殖系统发育的周期均较红豆杉科Taxaceae其他种更长，其冗长的生殖周期、复杂的生殖过程和雌性生殖系统发育明显滞后于雄性生殖系统，加之种群个体数量极度稀少(1雌株、2雄株)是造成结籽率低下、自然更新困难的主要原因。图3表1参34 Abstract:Objective Torreya jiulongshanensis, an ancient relict plant endemic to Zhejiang Province, is a national Class Ⅱ key protected wild plant and also listed as a protected plant of extremely small population in Zhejiang Province. The purpose of this study is to systematically observe the occurrence of megaspores and microspores of T. jiulongshanensis, as well as the development of male and female gametophytes , so as to find out whether the gametes are abnormal and the reasons for low seed-setting rate. Method The male and female individuals of T. jiulongshanensis from the type locality were collected as the research objects. Through regular observation and sampling, the occurrence of megaspores and microspores and the development of female and male gametophytes were sectioned, stained and observed by paraffin sectioning method. Result (1) The microsporophyll differentiated in mid-August, differentiated into secondary sporogenesis cells in late August and formed microspore mother cells in mid-November. In late February of the next year, it entered the meiosis stage, formed free microspore cells in mid-March, and gradually matured at the end of March. The mature pollen grains were binucleate without air sacs. The development type of microsporangium was basic type, and the spore division model was simultaneous type. The tetrad had tetrahedral and zygomorphic forms. (2) Male gametophytes began to develop in mid-to-late March, and formed sperm at the end of July. The whole process lasted about 4 months. (3) In late March of the next year, 2 erect ovules were born at the base of leaf axils of each bract in megasporophyll. In late April, the megaspore mother cell underwent meiosis Ⅰ and Ⅱ, and formed 4 linear megaspores. In mid-May, 3 megaspores at the micropylar end degenerated and the megaspores at the chalazal end finally developed into female gametophytes. (4) T. jiulongshanensis had 2 solitary oval archegonia, which lasted about 7 months from pollination to fertilization. Conclusion The development cycle of male and female reproductive system of T. jiulongshanensis is longer than that of other species in the family Taxaceae. The main reasons for low seed-setting rate and difficult natural regeneration of T. jiulongshanensis include long reproductive cycle, complicated reproductive process, the development of female reproductive system which lags behind that of the male reproductive system, as well as the extremely limited population (1 female plant and 2 male ones). [Ch, 3 fig. 1 tab. 34 ref.] -
Key words:
- Torreya jiulongshanensis /
- microsporogenesis /
- male gametophyte /
- megasporogenesis /
- female gametophyte
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图 1 大、小孢子叶球(雌、雄球花)的发育
Figure 1 Development of microstrobilus and macrostrobilus (male and female strobili)
图 2 大孢子发生和雌配子体发育(含受精作用)
Figure 2 Megasporogenesis and female gametophyte development (including fertilization)
图 3 小孢子发生和雄配子体发育(含精子形成)
Figure 3 Microsporogenesis and male gametophyte development (including spermatogenesis)
表 1 九龙山榧的有性生殖过程
Table 1. Process of sexual reproduction in T. jiulongshanensis
发育时期(年-月-日) 小孢子和雄配子体发育 大孢子和雌配子体发育 2018-08-20 次生造孢细胞 2018-11-15 小孢子母细胞形成 2019-02-26—03-06 小孢子母细胞减数分裂期 2019-03-12—03-16 四分体时期 2019-03-16 小孢子从四分体中相互分离 2019-03-19 单核靠边期 2019-04-06 散粉与传粉 造孢组织 2019-04-23 减数分裂Ⅱ后期 2019-05-18 功能大孢子 2019-05-30 管细胞、生殖细胞和不育细胞 2019-06-06 游离核时期 2019-06-24 管核与不育核即将消失 2019-07-27 精原细胞分裂形成两个大小相同的精细胞 2019-08-11 细胞化阶段 2019-09-22 颈卵器母细胞 2019-09-22—11-29 颈卵器阶段 2019-11-29 受精 
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