生物多样性 ›› 2018, Vol. 26 ›› Issue (5): 498-509. DOI: 10.17520/biods.2018046
所属专题: 传粉生物学; 昆虫多样性与生态功能
收稿日期:
2018-02-09
接受日期:
2018-05-15
出版日期:
2018-05-20
发布日期:
2018-09-11
通讯作者:
谭敦炎
作者简介:
# 共同第一作者
基金资助:
Jannathan Mamut1, Xiaojun Cheng1, Dunyan Tan1,2,*()
Received:
2018-02-09
Accepted:
2018-05-15
Online:
2018-05-20
Published:
2018-09-11
Contact:
Tan Dunyan
About author:
# Co-first authors
摘要:
异喙菊(Heteracia szovitsii)是分布于新疆北部荒漠、具短寿命花和多态果实的菊科一年生短命植物。本文对该物种的开花式样和花部特征以及繁育系统与昆虫传粉特性进行了研究, 以期明确其头状花序中小花的形态差异及其与多态果实间的关系、揭示其在短的花寿命内的繁殖特性。结果表明: 该物种头状花序中发育成多态果实的3类小花在数量、冠毛的有无以及舌片长/宽、子房宽、花柱裂片长和子房喙长之间均存在不同程度的差异, 且3类小花冠毛的有无以及子房宽和喙长与多态果实的形态一致, 说明多态果实的数量与形态在小花发育过程中就已完成分化。花序中的小花每天集中在上午开放, 使花序在功能上犹如一朵花, 可快速增大花展示和对传粉者的吸引, 以使异交在开花后短时间内完成; 3类小花的自花花粉均可在花柱裂片上萌发并长出花粉管, 说明该物种自交亲和。3类小花不去雄套袋后均能结实, 且自然授粉后的结实率显著高于不去雄套袋的, 说明该物种能自主自花授粉, 且异花授粉促进了结实。3类小花通过雄性先熟以及次级花粉呈现中的泵/刷机制, 将花粉呈现在花柱上部和未展开的柱头裂片外侧刷状毛上以及裂片顶端, 延长了花粉呈现(雄性阶段)持续的时间, 减少了花内雌雄功能间的干扰, 有利于传粉者异花传粉, 提高了雄/雌性适合度; 同时, 柱头裂片展开时可将自花花粉落置到裂片上, 以完成自主自花授粉。主要传粉者六显甜花蜂(Halictus sexnotatulus)的访花时间与花序开放时间、花粉活力最高期和柱头最佳授粉期高度吻合, 有利于雄性阶段花粉的输出以及雌性阶段柱头接受花粉, 保障授粉在开花后短时间内快速有效地完成以及异交成功。在新疆北部荒漠的春季环境中, 具短寿命花的异喙菊既可在低温多风导致传粉者不足或活动受限时, 通过快速自主自花授粉及自交亲和机制来提供繁殖保障, 也可在传粉者活动正常时, 通过集中开花、雄性先熟及次级花粉呈现等机制来快速完成异交, 提高雌/雄性及后代的适合度。
吉乃提汗·马木提, 成小军, 谭敦炎 (2018) 荒漠短命植物异喙菊的小花异形性及繁殖特性. 生物多样性, 26, 498-509. DOI: 10.17520/biods.2018046.
Jannathan Mamut, Xiaojun Cheng, Dunyan Tan (2018) Heteromorphism of florets and reproductive characteristics in Heteracia szovitsii (Asteraceae), a desert ephemeral annual herb. Biodiversity Science, 26, 498-509. DOI: 10.17520/biods.2018046.
图1 异喙菊头状花序与小花的形态、花粉呈现和访花昆虫。(A)头状花序纵剖面; (B)小花及其花粉呈现 (a: 外围小花, b: 过渡小花; c: 中央小花; d: 子房喙; e: 冠毛, f: 花粉呈现); (C)六显甜花蜂; (D)细腹食蚜蝇; (E)缘斑鼓额蚜蝇。
Fig. 1 The morphology of the capitulum and florets, its pollen presentation and flower visitors of Heteracia szovitsii. (A) Longitudinal section of the capitulum; (B) Florets and their pollen presentation (a, Peripheral floret; b, Intermediate floret; c, Central floret; d, Ovary beaks; e, Pappi; f, Pollen presentation); (C) Halictus sexnotatulus; (D) Sphaerophoria cylindrica; (E) Scaeva montana.
日开放动态 Diurnal flowering dynamics | 外围小花 Peripheral floret | 过渡小花 Intermediate floret | 中央小花 Central floret | F | P |
---|---|---|---|---|---|
花冠舌片展开到柱头从聚药雄蕊筒中伸出持续时间 Duration time from corolla ligules expanding to stigma exposure from sygensious stamen tube (min) | 19.13 ± 0.60a | 12.20 ± 0.59b | 10.53 ± 0.45c | 67.718 | < 0.001 |
柱头伸出到柱头裂片完全开展持续时间 Duration time from stigma exposure until its lobes are completely expanded (min) | 51.40 ± 0.98a | 45.10 ± 1.28b | 33.23 ± 0.80c | 78.218 | < 0.001 |
柱头裂片完全开展到花冠舌片萎蔫闭合持续时间 Duration time from stigma lobes completely expanded until corolla ligules wilt and close (min) | 110.20 ± 1.71a | 57.40 ± 1.35b | 18.83 ± 0.71c | 119.43 | < 0.001 |
单花开放持续时间 Anthesis duration time per floret (min) | 173.81 ± 2.33a | 109.77 ± 1.37b | 74.47 ± 1.23c | 855.503 | < 0.001 |
表1 异喙菊3类小花在头状花序中的开放动态(平均值 ± 标准误差)
Table 1 Flowering dynamics of the three floral morphs in the capitula of Heteracia szovitsii (mean ± SE)
日开放动态 Diurnal flowering dynamics | 外围小花 Peripheral floret | 过渡小花 Intermediate floret | 中央小花 Central floret | F | P |
---|---|---|---|---|---|
花冠舌片展开到柱头从聚药雄蕊筒中伸出持续时间 Duration time from corolla ligules expanding to stigma exposure from sygensious stamen tube (min) | 19.13 ± 0.60a | 12.20 ± 0.59b | 10.53 ± 0.45c | 67.718 | < 0.001 |
柱头伸出到柱头裂片完全开展持续时间 Duration time from stigma exposure until its lobes are completely expanded (min) | 51.40 ± 0.98a | 45.10 ± 1.28b | 33.23 ± 0.80c | 78.218 | < 0.001 |
柱头裂片完全开展到花冠舌片萎蔫闭合持续时间 Duration time from stigma lobes completely expanded until corolla ligules wilt and close (min) | 110.20 ± 1.71a | 57.40 ± 1.35b | 18.83 ± 0.71c | 119.43 | < 0.001 |
单花开放持续时间 Anthesis duration time per floret (min) | 173.81 ± 2.33a | 109.77 ± 1.37b | 74.47 ± 1.23c | 855.503 | < 0.001 |
花类型 Type of floret | 外围小花 Peripheral floret | 过渡小花 Intermediate floret | 中央小花 Central floret | F | P | ||
---|---|---|---|---|---|---|---|
每花序花数目 No. of florets per capitulum | 8.02 ± 0.13a | 10.10 ± 0.35b | 15.60 ± 0.63c | 100.09 | < 0.05 | ||
冠毛 Pappi | 数目 Number | - | - | 44.85 ± 1.21 | - | - | |
长 Length (mm) | - | - | 3.01 ± 0.04 | - | - | ||
花冠 Corolla | 舌片 Ligule | 长 Length (mm) | 4.89 ± 0.10a | 4.28 ± 0.08b | 3.92 ± 0.05c | 37.82 | < 0.05 |
宽 Width (mm) | 1.84 ± 0.06a | 1.56 ± 0.03b | 1.37 ± 0.02c | 46.02 | < 0.05 | ||
花冠筒长 Length of corolla tubes (mm) | 1.49 ± 0.05a | 1.37 ± 0.07a | 1.35 ± 0.06a | 1.71 | 0.19 | ||
雄蕊 Stamen | 花丝长 Length of filaments (mm) | 1.07 ± 0.05a | 1.02 ± 0.07a | 1.04 ± 0.06a | 0.17 | 0.85 | |
花药长 Length of anthers (mm) | 1.10 ± 0.04a | 1.15 ± 0.03a | 1.17 ± 0.04a | 1.23 | 0.30 | ||
雌蕊 Pistil | |||||||
子房 Ovary | 长 Length (mm) | 1.18 ± 0.06a | 1.03 ± 0.05b | 0.91 ± 0.03b | 8.62 | < 0.05 | |
宽 Width (mm) | 0.93 ± 0.04a | 0.57 ± 0.03b | 0.39 ± 0.02c | 70.91 | < 0.05 | ||
子房喙长 Length of ovary beaks (mm) | 0.30 ± 0.02a | 0.32 ± 0.02a | 0.36 ± 0.01b | 4.34 | < 0.05 | ||
花柱长 Length of style (mm) | 4.06 ± 0.15a | 4.21 ± 0.14a | 3.84 ± 0.15a | 1.64 | 0.20 | ||
柱头裂片长 Length of stigma lobes (mm) | 0.81 ± 0.03a | 0.77 ± 0.03ab | 0.72 ± 0.01b | 3.13 | < 0.05 |
表2 异喙菊头状花序中小花的数量特征(平均值 ± 标准误)
Table 2 The quantitative characteristics of florets in capitulum of Heteracia szovitsii (mean ± SE)
花类型 Type of floret | 外围小花 Peripheral floret | 过渡小花 Intermediate floret | 中央小花 Central floret | F | P | ||
---|---|---|---|---|---|---|---|
每花序花数目 No. of florets per capitulum | 8.02 ± 0.13a | 10.10 ± 0.35b | 15.60 ± 0.63c | 100.09 | < 0.05 | ||
冠毛 Pappi | 数目 Number | - | - | 44.85 ± 1.21 | - | - | |
长 Length (mm) | - | - | 3.01 ± 0.04 | - | - | ||
花冠 Corolla | 舌片 Ligule | 长 Length (mm) | 4.89 ± 0.10a | 4.28 ± 0.08b | 3.92 ± 0.05c | 37.82 | < 0.05 |
宽 Width (mm) | 1.84 ± 0.06a | 1.56 ± 0.03b | 1.37 ± 0.02c | 46.02 | < 0.05 | ||
花冠筒长 Length of corolla tubes (mm) | 1.49 ± 0.05a | 1.37 ± 0.07a | 1.35 ± 0.06a | 1.71 | 0.19 | ||
雄蕊 Stamen | 花丝长 Length of filaments (mm) | 1.07 ± 0.05a | 1.02 ± 0.07a | 1.04 ± 0.06a | 0.17 | 0.85 | |
花药长 Length of anthers (mm) | 1.10 ± 0.04a | 1.15 ± 0.03a | 1.17 ± 0.04a | 1.23 | 0.30 | ||
雌蕊 Pistil | |||||||
子房 Ovary | 长 Length (mm) | 1.18 ± 0.06a | 1.03 ± 0.05b | 0.91 ± 0.03b | 8.62 | < 0.05 | |
宽 Width (mm) | 0.93 ± 0.04a | 0.57 ± 0.03b | 0.39 ± 0.02c | 70.91 | < 0.05 | ||
子房喙长 Length of ovary beaks (mm) | 0.30 ± 0.02a | 0.32 ± 0.02a | 0.36 ± 0.01b | 4.34 | < 0.05 | ||
花柱长 Length of style (mm) | 4.06 ± 0.15a | 4.21 ± 0.14a | 3.84 ± 0.15a | 1.64 | 0.20 | ||
柱头裂片长 Length of stigma lobes (mm) | 0.81 ± 0.03a | 0.77 ± 0.03ab | 0.72 ± 0.01b | 3.13 | < 0.05 |
图2 荧光显微镜下异喙菊花粉在柱头上落置、萌发及花 粉管生长情况。(A)花粉落置; (B)花粉萌发(a: 花粉; b: 花粉管)。
Fig. 2 Pollen deposition, pollen germination and pollen tube growth on stigma of Heteracia szovitsii under the fluorescence microscope. (A) Pollen deposition; (B) Pollen germination (a, Pollen; b, Pollen tube)
花类型 Type of florets | 自然传粉 Natural pollination | 不去雄套纸袋 Bagged florets without emasculation | F1 | P |
---|---|---|---|---|
外围小花 Peripheral floret | 90.16 ± 1.79Aa | 79.05 ± 2.42Ab | 6.08 | < 0.001 |
过渡小花 Intermediate floret | 88.14 ± 1.81Aa | 74.60 ± 3.01Ab | 7.90 | < 0.001 |
中央小花 Central floret | 87.87 ± 1.56Aa | 77.58 ± 1.71Aa | 8.16 | < 0.001 |
F2 | 0.53 | 0.86 | ||
P | 0.592 | 0.426 |
表3 异喙菊3类小花在不同处理下的结实率(%, 平均值 ± 标准误)
Table 3 Fruit-set in the three floral morphs of Heteracia szovitsii under different treatments (mean ± SE)
花类型 Type of florets | 自然传粉 Natural pollination | 不去雄套纸袋 Bagged florets without emasculation | F1 | P |
---|---|---|---|---|
外围小花 Peripheral floret | 90.16 ± 1.79Aa | 79.05 ± 2.42Ab | 6.08 | < 0.001 |
过渡小花 Intermediate floret | 88.14 ± 1.81Aa | 74.60 ± 3.01Ab | 7.90 | < 0.001 |
中央小花 Central floret | 87.87 ± 1.56Aa | 77.58 ± 1.71Aa | 8.16 | < 0.001 |
F2 | 0.53 | 0.86 | ||
P | 0.592 | 0.426 |
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