南京林业大学学报(自然科学版) ›› 2023, Vol. 47 ›› Issue (1): 154-162.doi: 10.12302/j.issn.1000-2006.202108013
收稿日期:
2021-08-08
接受日期:
2021-10-23
出版日期:
2023-01-30
发布日期:
2023-02-01
通讯作者:
陈赢男
基金资助:
XIANG Li(), YIN Tongming, CHEN Yingnan()
Received:
2021-08-08
Accepted:
2021-10-23
Online:
2023-01-30
Published:
2023-02-01
Contact:
CHEN Yingnan
摘要: 【目的】 柳树雌株性成熟后会产生严重的飞絮污染,阐明其性别分化的分子机制是控制飞絮污染的关键。研究柳树雌、雄花芽分化以及花序建成的关键时期,为性别分化分子调控机理研究奠定基础。【方法】 以木本植物簸箕柳(Salix suchowensis)雌株和雄株花芽及花序为材料,从当年6月至翌年2月分别在17个时间点采集样品,利用石蜡切片技术对其雌、雄花芽分化及花序建成过程进行观察,分析不同发育阶段的相关特征。【结果】 簸箕柳花序建成初期为6月中旬至7月初,小花原基分化期在7月中旬至8月下旬,雌、雄蕊分化期在9月上旬至11月初,11月中旬至翌年2月初雌雄花序进入休眠期,翌年2月中旬至3月为大小孢子发生及雌雄配子体发育期,随后雌雄花序开放。研究表明,花序外观形态变化与内部解剖结构变化之间具有一定的相关性。【结论】 簸箕柳雌、雄花均为单性花。花序生长趋势总体呈“S”形上升趋势,在整个发育过程中,雄花序比雌花序外形稍大。簸箕柳花序外观形态可以用于判断花发育的关键时期。研究结果为全面了解簸箕柳花芽分化以及花序建成的完整过程提供了详细信息,为进一步阐明不同关键发育阶段的分子调控机制奠定了基础。
中图分类号:
向丽,尹佟明,陈赢男. 南京地区簸箕柳雌、雄花序建成动态观察[J]. 南京林业大学学报(自然科学版), 2023, 47(1): 154-162.
XIANG Li, YIN Tongming, CHEN Yingnan. The dynamic observation of male and female inflorescence establishment of Salix suchowensis in Nanjing area[J].Journal of Nanjing Forestry University (Natural Science Edition), 2023, 47(1): 154-162.DOI: 10.12302/j.issn.1000-2006.202108013.
表1
簸箕柳雌、雄花不同发育阶段"
分化阶段 differentiation stage | 采样时期 sampling time |
---|---|
花序建成初期(Stage Ⅰ) early stage of inflorescence establishment | T1—T2 |
小花原基分化期(Stage Ⅱ) floret primordium differentiation stage | T3—T5 |
雌/雄蕊分化期(Stage Ⅲ) gynoecium or androecium differentiation stage | T5—T10 |
休眠期Stage IV dormancy stage | T11—T16 |
大/小孢子发生及雌配子体/雄配子体发育期(Stage V) female or male gametophyte development stage | T17 |
图4
簸箕柳雌花发育过程的解剖结构观察 T1—T2. 纵切面,花序分化,花序轴形成,出现小花原基longitudinal section, inflorescence primordium differentiation, inflorescence axis formation, floret primordium appearance;T3. 纵切面,小花原基分化发育,雌蕊原基形成longitudinal section, floret primordium differentiation and development, pistil primordium formation;T4—T8. 纵切面,雌蕊原基上部向内分两叉凹陷,子房膨大开始发育,形成多个胚珠原基longitudinal section, that the upper part of the pistil primordium is bifurcated inwardly, and the ovary expands and develops, forming multiple ovule primordia;T9—T10. 纵切面,子房逐渐膨大,胚珠渐渐成熟longitudinal section, that the ovary gradually expands, and the ovule gradually matures;T12、T16. 纵切面,雌蕊分化缓慢,进入休眠期longitudinal section, that the pistil differentiates slowly and enters the dormant period;T17. 纵切面,花序打破休眠,雌蕊继续快速分化,胚珠成熟形成胚囊longitudinal section, that the flower bud breaks dormancy, the pistil continues to differentiate rapidly, and the ovule matures to form an embryo sac.Br.小苞片 bracteoles;Fp.小花原基 floret primordium;Pp.雌蕊原基 pistil primordium;Ov.子房 ovary;Olp.胚珠原基 ovule primordium;Ol.胚珠 ovule;Sty.花柱 style;Ovw.子房壁 ovary wall;Gl.腺体 gland."
图5
簸箕柳雄花发育过程的解剖结构观察 T1—T2. 纵切面,花序分化,花序轴形成,出现小花原基 longitudinal section, inflorescence primordium differentiation, inflorescence axis formation, floret primordium appearance;T3—T4.纵切面,小花原基不断分化发育并增大,形成雄蕊原基longitudinal section, floret primordium continuously differentiated and enlarged, forming stamen primordium;T5—T10. 纵切面,雄蕊原基增大发育,分化出花药原始体,分裂形成4个角隅小花粉囊的花药longitudinal section, the stamen primordium enlarges and develops, differentiates into anther primitives, and splits to form anthers of four corner small pollen sacs;T11—T16. 纵切面,雄蕊发育缓慢,进入休眠期,小孢子紧密排列在花粉囊longitudinal section that the stamens develop slowly and enter the dormant period, and the microspores are tightly arranged in the pollen sac;T17. 纵切面,雄蕊饱满圆润的花粉粒松散密布花粉囊内longitudinal section, that the stamens are full and round pollen grains are loose and densely packed in the pollen sac.Sp.雄蕊原基 stamen primordium;St.雄蕊 stamen;Anp.花药原基 anther primordia;Pos.花粉囊 pollen sac;Pog.花粉粒 pollen grain;Fi.花丝 filament;Gl.腺体gland;En.药室内壁 endothecium."
表2
不同时期簸箕柳雌、雄花序外部形态变化和内部解剖结构的比较"
采样时间 sampling time | 外部形态 external morphology | 解剖结构anatomical structure | |
---|---|---|---|
雌female | 雄male | ||
2020-06-15 (T1) | 雌花芽瘦小较瘪,长约5.2 mm;雄花芽稍大,长约7.9 mm。雌、雄花芽均上窄下宽,外披嫩绿色鳞片 | 雌花序和苞片快速分化,苞片腋部细胞密集分化成小花原基突起 | 雄花序和苞片快速分化,苞片腋部细胞密集分化成小花原基突起 |
2020-08-01 (T4) | 花序逐渐膨大,雌花序长约8.0 mm,雄花序长约11.2 mm,鳞片颜色嫩绿色 | 苞片不断伸长,小花原基逐渐分化增大,形成雌蕊原基 | 苞片不断伸长,小花原基逐渐分化增大,形成雄蕊原基 |
2020-09-01 (T6) | 花序继续膨大,逐渐饱满,雌花序长约9.2 mm,雄花序长约12.0 mm,鳞片颜色变成浅绿色 | 雌蕊原基从上往下分化,形成两叉凹陷的2心皮 | 雄蕊原基开始从中间向两边分化 |
2020-10-01 (T8) | 花序膨大,趋向饱满,雌花序长约10.1 mm,雄花序长约12.8 mm,鳞片浅黄色 | 子房膨大,基部出现1~2个胚原基突起 | 雄蕊原基两侧增大分化成4个角隅,角隅处细胞分化成花药原始体 |
2020-11-01 (T10) | 花序饱满,雌花序长约10.5 mm,雄花序长约13.1 mm,鳞片浅黄色且芽体两侧微红 | 子房继续膨大,胚原基分化成多个胚珠 | 角隅处花药雏形开始分化 |
2020-12-01 (T12) | 花序缓慢增大,雌花序鳞片变深黄色,长约10.7 mm。雄花序鳞片浅褐色间淡红色,长约13.6 mm | 子房膨大变化幅度小,胚珠呈扁椭圆形,进入休眠期 | 花粉囊增大缓慢,内部花药原基密集排列,进入休眠期 |
2021-01-01 (T14) | 花序生长停滞,雄花序比雌花序稍大,雌花序长约11.0 mm,雄花序长14.2 mm,其鳞片均为褐色 | 胚珠排列紧凑,发育停滞 | 花药原基密集排列,发育停滞 |
2021-02-16 (T17) | 雌、雄花序快速伸长膨大,雌花序长约19.8 mm,雄花序长约21.0 mm,鳞片脱落,花序露出,花序绒毛出现 | 子房膨大成椭圆形,胚珠饱满,趋向成熟的雌蕊形成胚囊 | 雄蕊花粉成熟,花粉粒松散密布分散在花粉囊里 |
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