含笑花雄蕊发生与发育的基本特征

doi:10.12302/j.issn.1000-2006.202307066

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南京林业大学学报（自然科学版） ›› 2024, Vol. 48 ›› Issue (6): 34-40.doi: 10.12302/j.issn.1000-2006.202307066

含笑花雄蕊发生与发育的基本特征

刘思彤¹(), 聂堂杰¹, 吴庆贤¹, 靳乐妮¹, 万小霞², 尹增芳¹^,^*()

1.南京林业大学,南方现代林业协同创新中心,南京林业大学生命科学学院,江苏南京 210037
2.浙江大学冷冻电镜中心,浙江杭州 310058

收稿日期:2023-07-05 修回日期:2023-09-25 出版日期:2024-11-30 发布日期:2024-12-10
通讯作者: *尹增芳(zfyin@njfu.edu.cn),教授。
作者简介:
刘思彤(t13630359868@163.com)。
基金资助:
江苏省林业科技创新与推广项目(LYKJ[2019]44);江苏高校优势学科建设工程资助项目(PAPD)

The basic characteristics of stamen morphogenesis and development in Michelia figo

LIU Sitong¹(), NIE Tangjie¹, WU Qingxian¹, JIN Leni¹, WAN Xiaoxia², YIN Zengfang¹^,^*()

1. Co-Innovation Center for Sustainable Forestry in Southern China, College of Life Science, Nanjing Forestry University, Nanjing 210037, China
2. Cryo-electron Microscope Center, Zhejiang University, Hangzhou 310058, China

Received:2023-07-05 Revised:2023-09-25 Online:2024-11-30 Published:2024-12-10

摘要/Abstract

摘要：

【目的】观察含笑花(Michelia figo)雄蕊形态建成过程,探究其雄蕊发生和发育的基本特征,为木兰科植物繁育与进化生物学的研究积累基础资料。【方法】利用扫描电镜、显微成像及石蜡切片技术,对不同生长阶段含笑花花芽中的雄蕊进行解剖观察。【结果】含笑花雄蕊原基的发生方式为螺旋状排列、向心方式发生。随着雄蕊形态建成的完成,其表观形态特征发生系列变化,最终形成较长的花药与较短的花丝。花药组织包括花药隔和4个花药室。花药室内造孢组织分化形成的小孢子母细胞减数分裂不同步,最终形成左右对称型和正四面体型小孢子四分体,之后游离的小孢子发育成二细胞型花粉和少量的三细胞型花粉。花药壁由1层表皮细胞、1层药室内壁细胞、2~3层中层细胞及1层分泌绒毡层细胞组成。成熟花粉发育时期绒毡层解体消失,但部分中层细胞宿存,直至花药壁开裂时才完全解体消失。花丝外有1层表皮细胞包裹,内部由薄壁组织填充,中央贯穿着直达药隔的维管束。发育早期的花丝薄壁组织内具有气腔的结构,成熟期消失。【结论】含笑花雄蕊发生与发育的过程基本正常,雄蕊螺旋状向心发生、小孢子母细胞减数分裂不同步、花丝气腔结构的时序变化、多数左右对称型小孢子四分体及二细胞型花粉等基本特征,体现了木兰科植物的原始性。

关键词: 含笑花, 雄蕊发生, 花丝, 花药, 小孢子四分体

Abstract:

【Objective】The fundamental characteristics of stamen morphogenesis and development were described according to the observation of stamen morphogenesis in Michelia figo. The findings provide valuable basic data for future investigations in plant breeding and evolutionary biology within the Magnoliaceae family.【Method】Using the scanning electron microscopy and paraffin section technology, the stamens of flower buds in M. figo at different growth stages were observed.【Result】The formation pattern of stamen primordium in M. figo was found to be helical and centripetal. Upon the completion of stamen morphogenesis, distinct morphological changes were observed, leading to the formation of a longer anther and a shorter filament. The anther tissue consists of anther septa and four anther locules. Normally, the sporogenous tissue differentiates gradually into microspore mother cells in the anther locule. The microspore mother cell meiosis was not synchronized, and eventually the isobilateral and tertrahedroid microspore tetrads were formed. Then, the free microspore developed into two-celled pollen and a few three-celled pollen. The structure of the anther wall was composed of one layer of epidermis cells, one layer of endothecium cells, two to three layers of middle layer cells, and one layer of secretory tapetum cells. During the development of mature pollen, the tapetum cells disintegrated and disappeared, but some middle layer cells persisted until the anther wall underwent dehiscence. The filaments were surrounded by a single layer of epidermis cells on the outer surface. Internally, the filaments were composed of parenchyma tissue, and a vascular bundle ran through the center toward the connective tissue. There was the cavity structure in the parenchyma of the filaments during the earlier development stage, which later disappeared during the mature stage of the stamen.【Conclusion】The process of stamen morphorgenesis in M. figo is normal compared with other Michelia plant species. The fundamental characteristics observed, including the helical centripetal formation of stamens, asynchronous meiotic division of microspore mother cells, temporal changes of the cavity structure in filaments, predominance of isobilateral tetrads, and two-celled pollen reflected the inherent nature of the Magnoliaceae family.

Key words: Michelia figo, stamen morphorgenesis, filament, anther, microspore tetrad

中图分类号:

S718
Q944.33

刘思彤,聂堂杰,吴庆贤,等. 含笑花雄蕊发生与发育的基本特征[J]. 南京林业大学学报（自然科学版）, 2024, 48(6): 34-40.

LIU Sitong, NIE Tangjie, WU Qingxian, JIN Leni, WAN Xiaoxia, YIN Zengfang. The basic characteristics of stamen morphogenesis and development in Michelia figo[J].Journal of Nanjing Forestry University (Natural Science Edition), 2024, 48(6): 34-40.DOI: 10.12302/j.issn.1000-2006.202307066.

图/表 4

表1

含笑花花芽及雄蕊发育时序"

时间(月-日) time (month-day)	花芽长/mm bud length	花芽宽/mm bud width	雄蕊表观形态 stamen apparent morphology	雄蕊发育状态 stamen developmental status
时间(月-日) time (month-day)	花芽长/mm bud length	花芽宽/mm bud width	雄蕊表观形态 stamen apparent morphology	雄配子male gametophyte	花药壁anther wall
06-01—06-15	2.68±0.85	1.79±0.62	半球形雄蕊原基 hemispherical stamen primordium	花药原基 anther primordium	表皮原细胞 dermatogen cell
07-01	3.42 ±0.89	1.88 ±0.63	雄蕊淡黄色、半透明状,花药和花丝开始分化light yellow translucent stamens, begin to differentiate into anthers and filaments	孢原细胞 archesporial cell	1层表皮原细胞 one layer of dermatogen cell
11-20	6.16±0.56	3.22±0.19	雄蕊黄绿色、透明状,花药和花丝区别明显yellow-green transparent stamens, anthers distinct with filaments obviously	造孢细胞 sporogenous cell	周缘细胞分裂 peripheral cells division
12-01	7.61±0.45	3.76±0.92	雄蕊开始积累黄色色素stamens begin to accumulate yellow pigment	小孢子母细胞 microspore mother cell	分化出药室内壁、中层、绒毡层 endothecium, middle layerand tapetum layer differentiated
12-10—01-05	8.02±0.81	4.22±0.67	雄蕊淡黄色yellowish stamens	小孢子母细胞减数分裂 microspore mother cell meiosis	中层细胞开始解体 middle layer cells begin to disintegrate
01-05—02-20	9.85±0.74	5.06±0.87	雄蕊黄色,花药先端积累紫色色素 yellow stamens, the tips of anthers begin to accumulate purple pigment	小孢子四分体 microspore tetrad	表皮细胞扁平方形;药室内壁径向伸长squarish epidermal cells; endothecium elongated radially
02-25—03-10	11.83±1.46	4.24±1.04	雄蕊紫色色素逐渐消退stamens losing purple pigment gradually	小孢子 microspore	未完全消失的中层细胞 middle layer cells haven’t completely disappeared
03-20	16.17±1.59	9.24±1.29	花药积累黄色色素,花丝积累紫色色素anthers accumulating yellow pigment, filaments accumulating purple pigment.	二细胞型花粉 two-celled pollen	药室内壁细胞壁栓质增厚,绒毡层细胞空泡化endothecium cell wall thickening with cork, the tapetum layer cells vacuolated
04-01	21.28±3.41	12.02±2.38	花药黄色带紫色条纹,花丝紫红色 yellow anthers with purple stripes, purple-red filaments	成熟花粉粒 mature pollen grain	中层、绒毡层完全解体消失 middle layer and tapetum layer disintegrated and disappeared completely
04-05	22.13±2.36	21.62±2.43	花药纵向开裂anthers dehiscent longitudinally	散粉与传粉pollen dispersal and pollination	花药壁开裂anther wall dehiscence

表1

图1

图2

含笑花雄蕊花药壁和雄配子体的发育过程 ac.孢原细胞archesporial cell;pp.初生壁细胞primary parietal cell;ps.初生造孢细胞primary sporogenous cell;ep.表皮epidermis;en.药室内壁endothecium;ml.中层middle layer;ta.绒毡层tapetum layer;mmc.小孢子母细胞microspore mother cell;te.四分体tetrad stage;tc.二细胞型花粉two-celled pollen;tcp.三细胞型花粉three-celled pollen;ct.木栓质加厚的药室内壁cork thicking wall of the endothecium。 a.花芽横切面,示花药原基the transverse section of flower bud, showing anther primordium;b.花药原基,示孢原细胞the anther primordium, showing the archesporial cell;c.发育早期花药壁与初生造孢组织the anther wall and the primary sporogenous tissue in early development stages;d.发育中期花药壁与初生造孢组织the anther wall and the primary sporogenous tissue in middle development stages;e.初生造孢组织与绒毡层primary sporogenous tissue and tapetum layer;f.花药壁与次生造孢组织anther wall and secondary sporogenous tissue;g.小孢子母细胞microspore mother cell;h.二分体与四分体dyad and tetrad;i.二分体dyad;j.四面体型小孢子四分体tertrahedroid tetrad;k.左右对称型小孢子四分体iIisobilateral tetrad;l.二细胞型花粉和三细胞型花粉the two-celled pollen and three-celled pollen;m.花药横切,示不同步发育的花药室the transverse section of anther, showing the asynchronous development of anther locule;n.小孢子单核居中期mononuclear in the center of microspore;o.小孢子单核靠边期mononuclear in the margin of microspore;p.花药横切,示花药壁the transverse section of anther, showing the anther wall;q.花药横切,示二细胞型花粉和三细胞型花粉the transverse section of anther, showing the two-celled pollen and three-celled pollen;r.散粉期,示花药开裂the pollen dispersal stage, showing dehiscent anther。"

图2

图3

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含笑花雄蕊发生与发育的基本特征

The basic characteristics of stamen morphogenesis and development in Michelia figo

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