生物多样性, 2019, 27(2): 159-167 doi: 10.17520/biods.2018317

研究报告

梅花个体内花柱长度的变异及其对繁殖成功的影响

王晓月1,2,3, 朱鑫鑫3, 杨娟3, 刘云静3, 汤晓辛,1,2,3,*

1 贵州师范大学国家林业局西南喀斯特山地生物多样性保护重点实验室, 贵阳 550025

2 贵州师范大学贵州省植物生理与发育调控重点实验室, 贵阳 550025

3 贵州师范大学生命科学学院, 贵阳 550025

Variation in style length and the effect on reproductive success in Chinese plums (Armeniaca mume)

Xiaoyue Wang1,2,3, Zhu Xinxin3, Yang Juan3, Liu Yunjing3, Tang Xiaoxin,1,2,3,*

1 Key Laboratory of State Forestry Administration on Biodiversity Conservation in Karst Mountainous Areas of Southwestern China, Guizhou Normal University, Guiyang 550025

2 Key Laboratory of Plant Physiology and Developmental Regulation, Guizhou Normal University, Guiyang 550025

3 School of Life Sciences, Guizhou Normal University, Guiyang 550025

通讯作者: * E-mail: txxfull@163.com

编委: 黄双全

责任编辑: 黄祥忠

收稿日期: 2018-11-27   接受日期: 2018-03-5   网络出版日期: 2019-02-20

基金资助: 国家自然科学基金(31560184)
国家自然科学基金(31300317)
贵州师范大学博士科研启动经费(11904/0518046)

Received: 2018-11-27   Accepted: 2018-03-5   Online: 2019-02-20

摘要

植物个体内花型的变异影响繁殖成功, 雌性繁殖性状的变异可能影响雌性的繁殖成功, 也可能作为花粉的受体影响雄性的繁殖成功。然而, 植物个体内不同花柱长度的花产生的花粉是否影响植物的繁殖成功却少有研究。梅(Armeniaca mume)是原产我国的重要木本花卉和经济果树, 我们的野外观察发现在同一植株内, 不同花的花柱长度有变异, 存在长柱型、短柱型和雄花型(雌蕊败育)三种花型, 是比较雌性繁殖性状的变异对两性繁殖成功的影响的理想材料。本文主要测量了不同花型的花部特征, 统计花期, 并开展体外花粉萌发以及人工控制授粉实验。结果表明: 长柱型的花冠直径、雌蕊长、单花花粉数、花粉体积显著大于短柱型和雄花型。长柱型的单花期以及雌花期显著长于短柱型。长柱型、短柱型以及雄花型花粉在活体柱头上的萌发率没有显著性差异, 雄花型的花粉管长度显著高于长柱型和短柱型。长柱型为母本的花粉萌发率以及花粉管的长度要显著高于短柱型。长柱型、短柱型、雄花型花粉授粉与自然对照处理的坐果率没有显著性差异, 而长柱型为母本的坐果率要显著高于短柱型为母本的坐果率。这些结果表明野生梅花的长柱型为母本有利于花粉的萌发和花粉管的伸长, 有高的坐果率; 但其作为花粉供体的雄性功能与其他花型没有差异。

关键词: ; 花柱长度变异 ; 个体内花变异 ; 萌发率 ; 花粉管长度 ; 坐果率 ; 花粉的受精率

Abstract

Style length might affect gametophyte competition during pollen germination and pollen tube elongation if pollen quality and quantity deposited on stigmas differs with style length. However, it is unknown whether pollen donors from flowers with different style lengths affect sexual reproduction. Field observation noted that style length of Armeniaca mume varied within individuals including those with long-style type, short-style type and male-flower type. Corolla diameters, pistil length, style length, pollen number and pollen volume of long-style type were significantly higher than those of short-style type. Flower longevity and female phase of long-style type were significantly longer than those of short-style type. Pollen germination rates among long-style, short-style and male-flower types were not significantly different. Pollen tube length of the male-flower type was significantly longer than that of long-style and short-style types. Pollen germination rates and pollen tube length of long-style type as pollen recipient were significantly higher than those of short-style type as pollen recipient. Fruit sets did not significantly differ between flowers pollinated by pollen from long-style, short-style and male-flower types with open-pollinated individuals. Fruit sets of long-style type as pollen recipient were significantly higher than those of short-style type as pollen recipient.

Keywords: Armeniaca mume ; variation in style length ; intraplant floral variation ; pollen germination rates ; pollen tube length ; fruit set ; pollen siring success

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引用本文

王晓月, 朱鑫鑫, 杨娟, 刘云静, 汤晓辛. 梅花个体内花柱长度的变异及其对繁殖成功的影响. 生物多样性, 2019, 27(2): 159-167 doi:10.17520/biods.2018317

Xiaoyue Wang. Variation in style length and the effect on reproductive success in Chinese plums (Armeniaca mume). Biodiversity Science, 2019, 27(2): 159-167 doi:10.17520/biods.2018317

同一种植株个体上不同花的结构和花型有着各种各样的变异(Bell, 1985; Barrett & Harder, 2005; Barrett & Hough, 2013)。例如榕属(Ficus)有雌雄同株或异株的物种, 花柱长度的变异往往在雌雄同株中比雌雄异株的大(Ma et al, 2009); 分布在青藏高原东缘的小花风毛菊(Saussurea parviflora), 花柱及其柱头分叉长度随着海拔的升高显著增长(王一峰等, 2008); 鸭跖草科鸭跖草(Commelina communis)同一个体上不同花的雌蕊有着明显不同的长度, 可分为两性花和雄花型(即结构上的雄花), 且两性花和雄花的花柱长度在不同居群间存在变异(Ushimaru et al, 2003)。这些植物同一个体上的不同花出现了性别分化, 即两性花植物中有些个体出现了雄花, 这种性系统为雄花两性花同株(andromonoecy)的种类约占被子植物种类的1.7% (黄双全和郭友好, 2000)。近期对欧洲千里光(Senecio vulgaris)的调查也表明花柱长度存在明显变异, 分为长柱型和短柱型(Love et al, 2016)。

花柱长度的变异意义可能与花大小、花粉体积、花粉管生长速率、花粉败育的比率等的变化有关(Plitmann, 1993; Johnson et al, 2010; Ramesha et al, 2011; Taylor & Williams, 2012)。Cresswell (2000)用细线替换油菜(Brassica napus)的部分雌蕊, 模拟了花柱长度的连续变化, 实验结果表明花柱长度并不影响花粉从花药中移出, 但长花柱落置了较多的花粉, 显示柱头长度影响了花粉落置的数量。Lankinen和Skogsmyr (2001)通过模型分析发现花柱的长度会影响花粉管生长的速率。Johnson等(2010)对柳叶菜科Oenothera属和Calylophus属70多种有性植物和功能上无性植物的分析表明, 由有性转变为无性的繁育系统时, 植株减少了对雌性(花柱)和雄性(花瓣、花冠管)功能的投资。这些研究表明花柱长度直接或间接地影响了花的雌雄两性功能。

花柱长度变异现象的研究已有许多推断, 相对于短花柱, 长花柱可能的功能包括: (1)柱头从花冠管中伸出利于接触传粉者身体, 促进柱头上花粉的落置(Bernasconi et al, 2007)。(2)提高花粉落置数量, 增加花粉萌发和花粉管伸长过程中配子体的竞争(Mulcahy, 1979; Erbar, 2003; Stephenson et al, 2003), 提高到达花柱和子房的花粉的基因质量, 也提高完成受精的胚珠的基因质量。Zhang等(2010)研究发现蔷薇科梨属沙梨(Pyrus pyrifolia)柱头表面落置更多的花粉提高了花粉萌发率、花粉管生长速率、种子的产量、果实的生长速率、坐果率以及果实的质量。然而, 增加柱头上花粉的落置数量并不一定提高种子的数量和后代的质量。Mazer等(2016)对花荵科克拉花属Clarkia unguiculataC. xantiana subsp. xantiana的研究表明: 花柱越长, 柱头落置的花粉数量越多; 柱头上花粉的数量与柱头表面到柱头与花柱的连接处、从连接处到花柱基部、从柱头表面到花柱基部的花粉衰败的比例显著正相关。花柱长度通过柱头堵塞或花粉之间的相互作用而增加花粉衰败的比率。(3)延长花粉管穿过花柱的距离, 花粉质量微小的差异会显著影响后代的繁殖成功(Mazer et al, 2010; Ramesha et al, 2011)。目前少有研究比较同一个体内不同花柱长度的花型对雌雄两性功能的影响。

我们在野外调查时发现, 同一植株的梅(Armeniaca mume, 也写为Prunus mume) (陈俊愉和包满珠, 1992), 花柱长度存在明显变异, 有花柱突出雄蕊群(定义为长柱型, long-style type), 花柱和雄蕊长度几乎一致(短柱型, short-style type), 以及花柱发育不良、极短且柱头呈黑色, 不具有可授性(雄花型, male-flower type) 3种类型。花柱长度变异从花没有开放至完全开放均容易区分, 例如长柱型的梅花, 其花柱在花蕾期就突出雄蕊群(图1), 方便我们判断花型。前人有研究表明植物个体内花型的变异影响繁殖成功, 雌性繁殖性状的变异可能影响雌性的功能(Lankinen & Skogsmyr, 2001; Bernasconi et al, 2007), 也可能作为花粉的受体影响雄性的功能。为了研究植物个体内不同花柱长度的花产生的花粉是否影响两性的繁殖成功, 本文以梅花为研究对象, 探讨以下问题: (1)不同花型哪些花部特征存在差异? 花柱长度是否和单花的花期变异有关? (2)不同花型的花的雄性功能(花粉的活力、花粉管的生长、坐果率)是否有显著性差异? (3)长柱型、短柱型花的雌性功能(作为母本进行授粉处理, 柱头上花粉的萌发率、花粉管在花柱中伸长以及授粉处理后的坐果情况)是否有显著性差异。

图1

图1   不同花柱长度花型的梅花。(A)长柱型、短柱型的花苞(花苞期即可区分); (B)长柱型; (C)短柱型; (D)雄花型。L: 长柱型; S: 短柱型; M: 雄花型。

Fig. 1   Flowers of Armeniaca mume showing different style lengths. (A) Buds of long-style type and short-style type (different type can be distinguished during the bud phase); (B) Long-style type; (C) Short-style type; (D) Male-flower type. L, Long-style type; S, Short-style type; M, Male-flower type.


1 材料与方法

梅为多年生乔木, 高达4-10 m, 通常分布在海拔1,700-3,100 m的林坡、疏林、河边、山坡、山地; 原产于四川省西部, 在我国大部分地区都有栽培。四川、贵州、云南交界的横断山区和云贵高原一带是梅的野生分布中心, 同时又是其遗传多样性中心(Lu & Bruce, 2003)。梅花是我国十大传统名花之一, 是冬季开花的木本花卉, 其果实可食用(梅子)、酿酒等(陈俊愉和包满珠, 1992)。花期在西南山区常为12月至翌年3月, 果实成熟期4-7月, 成熟时梅果黄绿色。本研究地点位于贵州省荔波县洞塘乡木朝村梅园(106°1¢11² E, 25°19¢47² N), 是梅花典型的野生分布区域。

1.1 比较不同花型的花部特征

为比较长柱型、短柱型和雄花型的花部特征, 随机选取30株梅花植株, 每株随机选择3种花型的花各1朵, 用游标卡尺测量(精确到0.01 mm)花萼长与宽、花冠直径、花瓣长与宽、雄蕊长、花药长、雌蕊长、花柱长, 并统计单花雄蕊数目。各个花部特征测量的具体方法参见附录1。采集每个植株的3种花型尚未开裂的花蕾各1朵, 保存在装有福尔马林-醋酸-酒精固定液(FAA, formalin-acetic acid-70% alcohol, 体积比为5 : 5 : 90)的离心管里。在实验室, 用镊子将花蕾的花药和子房分开, 分装在不同的离心管中, 对应标记。用镊子充分研磨花药并定容到2 mL, 充分摇匀后取20 mL液体滴于载玻片, 每花取3滴20 mL花粉溶液, 在光学显微镜下统计花粉的数量(a1, a2, a3), 花苞中花粉的总数量为(a1 + a2 + a3)/3 × 100。每个花粉液体样本用荧光显微镜拍照, 每张图片随机选择1粒花粉, 用图像分析软件(Digimizer Version 4.6.0)测量其直径(d)。梅花的花粉近球形, 体积(V) = 4p × (d/2)3/3。在解剖显微镜下观察花蕾中的胚珠, 并统计数量。

1.2 比较不同花型的花期

为探究花柱长度的变异与花所处的发育阶段以及不同花型的花期是否有关, 随机选择20个植株, 每个植株随机选择长柱型、短柱型、雄花型各1朵进行标记, 花蕾打开标记为雄花第1天, 每隔1天观察花的状态, 直至花萎蔫, 计为单花期。单花的雄期是从花蕾开放开始, 直到花药枯萎脱落的天数。收取不同时期的柱头, 用双氧水检测其活性。如果有活性, 柱头周围会有气泡(红雨等, 2006)。柱头活性实验表明, 梅花柱柱头膨大后才具有活性。单花的雌期是从柱头膨大到枯萎持续的天数。

1.3 比较不同花型花粉的萌发以及花粉管的生长

为比较不同的花型是否影响花粉的活力和花粉管的生长, 于2018年1月, 分别选择长柱型和短柱型单花作为花粉的受体, 长柱型、短柱型和雄花型的花粉作为供体进行授粉实验。随机选择30个植株, 每个植株随机标记6个花苞(长、短柱型花各3朵, 分别作为母本, 以不同颜色的尼龙线标记区分), 进行去雄套袋处理, 直至柱头成熟处于雌期。然后进行长柱型、短柱型和雄花型花粉授粉处理。从距离花粉受体植株至少30 m的植株上随机选择新鲜开裂的花药, 收集花粉并混合作为花粉源。选择晴朗的天气用小毛笔进行授粉, 每个雌期的柱头上涂满花粉(至少有100粒), 然后重新套袋。根据不同的花粉供体做出相应的标记。每个植株上共6种处理: (1)长柱型为母本, 授长柱型花粉; (2)长柱型为母本, 授短柱型花粉; (3)长柱型为母本, 授雄花型花粉; (4)短柱型为母本, 授长柱型花粉; (5)短柱型为母本, 授短柱型花粉; (6)短柱型为母本, 授雄花型花粉。每种处理1朵花。6 h后收集雌蕊, 储存在装有FAA固定液的1.5 mL离心管。在实验室中, 处理的雌蕊先在蒸馏水中清洗3次, 于室温下在8 mol/L的NaOH溶液中软化4 h。然后置于蒸馏水中漂洗3 min, 再在苯胺蓝溶液(0.1%质量浓度)中染色3 h。用荧光显微镜观察柱头, 统计柱头上花粉的总数量以及萌发的花粉数量(Wang et al, 2012)。花粉管先用荧光显微镜自带的软件拍照, 再用图像分析软件(Digimizer Version 4.6.0)测算花粉管的长度(Wang et al, 2018)。

1.4 人工控制授粉实验

为了比较不同父本(花粉供体)中长柱型、短柱型、雄花型的花粉是否导致结实的差异, 以及长柱型和短柱型作为母本(花粉受体)是否影响梅花的结实, 随机选择30个植株, 每个植株随机选择8个花苞, 长柱型和短柱型各4个, 分别进行4种授粉处理: 自然对照、长柱型花粉、短柱型花粉和雄花型花粉。对于每个植株已标记的花苞, 随机选择3个长柱型花苞, 3个短柱型花苞去雄套袋, 直至柱头膨大进入雌期, 然后分别进行长柱型、短柱型和雄花型花粉授粉处理, 剩余的1个长柱型和1个短柱型花苞作为自然对照处理。一个植株上共8种处理: (1)长柱型为母本, 自然对照; (2)长柱型为母本, 授长柱型花粉; (3)长柱型为母本, 授短柱型花粉; (4)长柱型为母本, 授雄花型花粉; (5)短柱型为母本, 自然对照; (6)短柱型为母本, 授长柱型花粉; (7)短柱型为母本, 授短柱型花粉; (8)短柱型为母本, 授雄花型花粉。每种处理1朵花。授粉后重新套袋以隔绝传粉者。2018年4月中下旬, 收集所有授粉操作处理的梅花果实, 在实验室中统计各种授粉处理的坐果情况。

1.5 数据分析

利用广义线性模型(generalized linear model, GLM)中的正态分布模型-恒定函数比较分析3种花型的花部特征、花粉体积、单花期、雌期及雄期,其中花部特征、花粉体积、单花期、雌期、雄期作为自变量, 3种花型作为因变量。采用泊松分布模型-对数线性函数分析比较3种花型的花粉数以及胚珠数, 其中花粉数以及胚珠数作为自变量, 3种花型作为因变量。采用二项式分布-Logistic关联函数分析不同花型柱头花粉萌发率, 其中柱头上萌发的花粉数量作为自变量, 柱头落置的总花粉数作为一次实验中总的变量, 不同的花粉来源、不同的母本以及二者的交互作用作为因变量。采用正态分布-恒定函数比较分析不同处理的花粉管长度, 花粉管的长度作为自变量, 不同的花粉来源、不同的母本以及二者的交互作用作为因变量。采用二项式分布-Logistic关联函数分析比较不同授粉处理的梅花坐果率, 果实的数量作为自变量, 授粉的花朵数作为一次实验中总的变量, 不同的授粉处理、不同的母本以及二者的交互作用作为因变量。

所有的数据分析在SPSS 20.0 (IBM Inc., New York, NY)中进行。

2 结果

2.1 三种花型花部特征和花期的比较

广义线性模型(GLM)对长柱型、短柱型和雄花型花部特征分析结果表明: 长柱型的花冠直径、雌蕊长、花柱长、单花的花粉数量以及花粉的体积显著高于短柱型以及雄花型(P < 0.05, 表1)。短柱型的雄蕊长度显著大于长柱型和雄花型(Wald χ2 = 33.193, df = 2, P < 0.001), 而雄花型的雄蕊数目显著高于长柱型和短柱型(Wald χ2 = 8.460, df = 2, P = 0.015)。其他花部特征的比较分析结果见表1。长柱型的花柱在花蕾期即突出雄蕊群, 持续整个花期。短柱型的花柱在花蕾期和雄蕊群长度基本一致, 并持续整个花期。雄花型从花蕾期就已雌蕊败育。长柱型和雄花型的单花期没有显著性差异, 但均显著高于短柱型(Wald χ2 = 6.620, df = 2, P = 0.037)。长柱型和短柱型的雄花期没有显著差异(Wald χ2 = 0.140, df = 2, P = 0.079)。长柱型的雌花期显著长于短柱型 (Wald χ2 = 4.670, df = 2, P = 0.031)(表1)。

表1   长柱型、短柱型、雄花型梅花花部特征(平均值 ± 标准误)的比较(广义线性模型)。表中数字加粗表示长柱型的指标显著大于短柱型和雄花型相应的指标。花部特征样本量为30, 花期样本量为20。不同小写字母表明不同花型对应的花部特征有显著性差异。

Table 1  Comparisons of floral traits (mean ± SE) among long-style, short-style and male-flower types within individuals of Armeniaca mume using generalized linear model (GLM). Floral traits of long-style type significantly larger than those of short-style and male-flower types are in bold. The sample size of floral characters and floral phase was 30 and 20, respectively. Different letters indicate significant differences in flower traits among three types in the same row.

花部特征
Flower character
长柱型
Long-style type
短柱型
Short-style type
雄花型
Male-flower type
Wald χ2df
P
花萼长 Spur length (mm)5.30 ± 0.09a5.38 ± 0.08a4.99 ± 0.10b8.97020.011
花萼宽 Spur width (mm)3.62 ± 0.05a3.78 ± 0.07a3.76 ± 0.12a2.75520.252
花冠直径 Corolla diameter (mm)24.42 ± 0.44a23.43 ± 0.23b21.64 ± 0.41c27.6562< 0.001
花瓣长 Petal length (mm)10.44 ± 0.18a10.03 ± 0.16a9.17 ± 0.18b22.1602< 0.001
花瓣宽 Petal width (mm)9.70 ± 0.20a9.31 ± 0.16a8.05 ± 0.25b32.4492< 0.001
雄蕊数 Stamen Number53.71 ± 0.71b53.56 ± 0.79b56.93 ± 1.10a8.46020.015
雄蕊长 Stamen length (mm)9.25 ± 0.21b10.39 ± 0.18a8.82 ± 0.22b33.1932< 0.001
花药长 Anther length (mm)1.01 ± 0.02a1.02 ± 0.01a1.01 ± 0.02a0.50020.779
雌蕊长 Pistil length (mm)14.46 ± 0.20a12.34 ± 0.21b3.42 ± 0.16c8.46020.015
花柱长 Style length (mm)12.20 ± 0.19a10.18 ± 0.18b/58.3941< 0.001
花粉数 Pollen Number37,522.22 ± 1,305.85a30,386.67 ± 1,412.62b32,000.00 ± 2,003.29b15.13320.001
花粉体积 Pollen volume (mm3)1,213.07 ± 28.87a1,102.47 ± 31.08b1,067.13 ± 49.45b10.22620.006
胚珠数 Ovule Number2.00 ± 0.00a2.00 ± 0.00a0.90 ± 0.35b46.6202< 0.001
单花期 Flower longevity (days)10.43 ± 0.24a9.29 ± 0.56b10.70 ± 0.30a6.62020.037
雄花期 Male phase (days)5.46 ± 0.35a5.23 ± 0.52a/0.14010.079
雌花期 Female phase (days)4.96 ± 0.30a4.06 ± 0.30b/4.67010.031

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2.2 三种花型的花粉萌发率以及花粉管生长的比较

整体上, 不同的花粉供体, 长柱型、短柱型、雄花型的花粉在活体柱头的萌发率(44.35% ± 5.08%, 59.81% ± 4.66%, 51.42% ± 5.39%)没有显著性差异(Wald χ2 = 1.010, df = 2, P = 0.604)(表2, 图2)。长柱型为母本(即花粉授到长柱型的柱头)的花粉萌发率(56.41% ± 4.43%)显著高于短柱型为母本的萌发率(46.50% ± 3.51%)(Wald χ2 = 4.110, df = 1, P = 0.043), 且花粉供体和花粉受体对梅花的花粉萌发率的影响没有交互作用(Wald χ2 = 3.860, df = 2, P = 0.145)(表2), 不同花粉供体对梅花花粉萌发率的影响在不同的花粉受体间是一致的。长柱型、短柱型、雄花型花粉授到长柱型的柱头, 花粉的萌发率(36.19% ± 7.80%, 72.08% ± 5.97%, 59.31% ± 7.89)没有显著性差异(Wald χ2 = 2.828, df = 2, P = 0.243)。长柱型、短柱型、雄花型花粉授到短柱型柱头, 花粉的萌发率(53.62% ± 5.84%, 43.46% ± 5.84%, 41.34% ± 6.54%)没有显著性差异(Wald χ2 = 3.570, df = 2, P = 0.168)。

表2   授粉处理(长柱型、短柱型和雄花型的花粉)时花粉受体(长柱型和短柱型分别为母本)及其交互作用对梅花柱头花粉的萌发率、花粉管长度的影响(广义线性模型)

Table 2  Effect of pollination treatments (long-style, short-style and male-flower types) and pollen recipient type (long-style vs short-style), and their interaction on pollen germination rates and pollen tube length in Armeniaca mume (generalized linear model)

变量 Source of variationWald χ2dfP
萌发率 Germination rates
授粉处理 Pollination treatments1.01020.604
花粉受体 Pollen recipient type4.11010.043
交互作用 Interaction3.86020.145
花粉管长度 Pollen tube length
授粉处理 Pollination treatments147.1782< 0.001
花粉受体 Pollen recipient type26.6151< 0.001
交互作用 Interaction19.5262< 0.001

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图2

图2   不同处理间梅花的花粉萌发率(A)和花粉管长度(B)的比较(平均值±标准误)。长柱型花粉、短柱型花粉、雄花型花粉分别为花粉供体, 短柱型和长柱型梅花分别作为花粉受体。相同字母表示处理间没有显著差异。条形图内部的数字表示处理的样本量。

Fig. 2   Comparisons of in vivo pollen germination rates (A) and pollen tube length (mean ± SE) (B) of pollen donors from long-style, short-style and male-flower types, with the long- style and short-style types in Armeniaca mume as pollen recipients. The same letters indicate no significant difference within the flower type. Numbers of sample size are shown in the bars.


对于花粉管长度而言, 雄花型为供体的花粉管长度(1,041.43 ± 87.37 mm)显著大于长柱型(386.70 ±31.03 mm)和短柱型(345.37 ± 23.71 mm), 长柱型和短柱型为供体的花粉管长度没有显著性差异(Wald χ2 = 147.178, df = 2, P < 0.001)(表2, 图2)。长柱型为母本的花粉管长度(559.78 ± 31.39 mm)显著高于短柱型为母本的花粉管长度(344.35 ± 29.98 mm)(Wald χ2 = 26.615, df = 1, P < 0.001)(表2, 图2)。花粉供体和花粉受体交互影响梅花的花粉管长度(Wald χ2 = 19.526, df = 1, P < 0.001)(表2), 不同花粉供体对梅花花粉管生长的影响在不同的花粉受体上不同。

2.3 人工控制授粉实验

对于不同供体的授粉处理的梅花坐果率而言, 长柱型(8.33% ± 3.43%)、短柱型(12.56% ± 4.31%)、雄花型(16.29% ± 5.70%)和自然对照(20.97% ± 7.16%)没有显著性差异(Wald χ2 = 0.704, df = 3, P = 0.872)。长柱型为母本的坐果率(20.45% ± 4.57%)显著高于(Wald χ2 = 3.914, df = 1, P = 0.048)短柱型为母本的坐果率(10.31% ± 2.93%)(表3, 图3), 且不同的授粉处理与不同的花粉受体对梅花坐果率没有显著的交互作用(Wald χ2 = 5.920, df = 3, P = 0.116), 授粉处理对梅花坐果率的影响在不同的花粉受体间是一致的(表3)。长柱型为花粉受体时, 授以长柱型(14.58% ±7.28%), 短柱型(19.44% ± 7.53%), 雄花型花粉(26.43% ±11.27%)和自然对照(22.73% ± 11.90%)的梅花的坐果率没有显著差异(Wald χ2 = 3.005, df = 3, P = 0.391)。短柱型为花粉受体时, 授以长柱型(4.17% ± 2.88%)、短柱型(9.38% ± 5.24%)、雄花型花粉(9.52% ± 5.58%)和自然对照(20.00% ± 9.18%)的梅花的坐果率没有显著差异(Wald χ2 = 2.920, df = 3, P = 0.404)。

表3   授粉处理(自然对照、长柱型、短柱型和雄花型的花粉)时花粉受体(长柱型和短柱型分别为母本)及其交互作用对梅花坐果率的影响(广义线性模型)

Table 3  Effect of pollination treatments (control, long-style type, short-style type, male-flower type) and pollen recipient type (long-style vs short-style), and their interaction on fruit set of Armeniaca mume (generalized linear model)

变量 Source of variationWald χ2dfP
坐果率 Fruit set
授粉处理 Pollination treatments0.70430.872
花粉受体 Pollen recipient type3.91410.048
交互作用 Interaction5.92030.116

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图3

图3   不同处理梅花坐果率比较。花粉供体为长柱型、短柱型、雄花型花粉和自然对照, 短柱型和长柱型梅花分别作为花粉受体。相同字母表示处理间没有显著的差异。条形图内部的数字表示处理的样本量。

Fig. 3   Comparison of fruit set between pollen donors from long-style, short-style and male-flower types, with the long-style and short-style types in Armeniaca mume as pollen recipients. Bars with the same letters are not significantly different. Numbers of sample size are shown in the bars.


3 讨论

本研究结果表明, 梅花同一植株内不同花的花柱长度有变异, 可分为长柱型、短柱型和雄花型。长柱型的花、单花花粉数量和花粉体积以及单花期与雌性期显著大于短柱型和雄花型。长柱型为母本有利于花粉在柱头上的萌发以及花粉管的伸长, 并能显著提高坐果率。长柱型为花粉供体的雄性功能(例如花粉的萌发率、坐果率)与短柱型和雄花型没有差异。

前人做了大量植物个体内花特征变异、性功能特化的研究(Lloyd, 1980; Diggle, 1995, 1997; Granado-Yela et al, 2017)。例如, Lloyd (1980)提出不同时期的花调整不同的资源配置, 主要包括花的性别决定、子房的发育和果实的成熟3个时期。其中, 对花性别决定的调节有利于植物雌性投入的调整, 保持雌性投入和雄性投入之间恒定的比例; Diggle (1995)提出结构效应假说, 即不同的花调整两性资源的配置, 是因为处在个体的不同位置。对实验居群不同花型数目统计结果显示梅花中的雄花型数目在2018年1月6日为0, 在1月10日为244朵, 1月18日为621朵, 雄花型在开花后期增多(未发表的数据)。观察显示雄花型随机分布在树枝上, 后期花雄性增多可能是因为植物随时间调配两性资源(Liu & Huang, 2013)。

长花柱可能通过促进花粉的落置或者促进花柱中花粉管的生长而提高植物的繁殖成功(Campbell, 1989; Bernasconi et al, 2007)。花荵科Ipomopsis aggregata的长雌蕊个体单花的雌期延长, 提高了柱头花粉的落置(Campbell, 1989)。Bernasconi等(2007)研究表明唇形科大花夏枯草(Prunella grandiflora)花柱的长度与花粉管的数量正相关, 花粉管的数量与结实率呈正相关, 花柱长度与种子产量间接正相关。然而长的花柱并不总是具有生殖上的优势, 例如欧洲千里光(Senecio vulgaris), 其结实率取决于柱头上花粉的落置量, 长花柱的柱头几乎不落置或者只落置很少量自花的花粉, 且萌发率低, 因此长柱型结实率低; 而短柱型能成功落置自身的花粉且亲和, 结实率高(Love et al, 2016)。我们的观察结果表明, 长柱型梅花的单花期和雌期显著高于短柱型, 花期延长增加了传粉者访问的概率, 柱头可能落置更多的花粉。长柱型梅花为母本有利于提高花粉的萌发率、花粉管长度和坐果率, 说明梅花的长花柱有利于结实。

在物种水平上, 花柱长度和花粉大小呈正相关(Roulston et al, 2000; Jürgens et al, 2012), 可能个体较小的花粉不能到达长花柱的胚珠从而导致了种子的败育。梅花长短柱花型内(长柱型为母本, 接受长柱型的花粉; 短柱型为母本, 接受短柱型的花粉)及花型间异交授粉(长柱型为母本, 接受短柱型的花粉; 短柱型为母本, 接受长柱型的花粉)的坐果率没有显著性差异, 说明长短柱花型之间是亲和的。雄花型的花粉管长度显著大于长柱型和短柱型, 可能是因为当单花变异为雄花时, 提高了对雄性功能资源的投入, 花粉粒大有利于花粉管的伸长, 但是花粉管的生长速率的不同还没有造成坐果率显著的差异。梅花的坐果率应该不仅受到花粉萌发和花粉管生长过程的影响, 可能还受到授粉后果实生长发育的环境因素以及资源投入的影响。虽然不同花型花粉管的生长速率不同, 但是坐果率没有差异。

梅花是冬季开花的木本花卉, 对其繁殖特性的了解还很缺乏。我们的野外观察表明梅花有多种访花动物, 但传粉贡献受天气影响(未发表的数据)。不同长度的花柱是否影响传粉者的访花行为以及传粉效率, 还有待进一步研究。

致谢: 感谢两位审稿专家及编委提出修改意见并指出原稿中的不足。

附录 Supplementary Material

附录1 梅花花部特征测量

Appendix 1 Measurements of floral traits in Armeniaca mume

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New Phytologist, 173, 393-400.

DOI:10.1111/j.1469-8137.2006.01920.x      URL     PMID:17204085      [本文引用: 4]

090004 Intraspecific variation in pollen deposition and number of pollen tubes per style is rarely quantified, but is essential for assessing the occurrence of pollen limitation and pollen competition and their evolutionary implications. Moreover, pollen deposition, pollen tube growth, and the fate of fertilized ovules are rarely distinguished in field studies. Here we present such a study in eight natural populations of Prunella grandiflora . 090004 We quantified microgametophyte population sizes and inferred pollen limitation when the number of fertilizable ovules exceeded pollen tubes, and assessed seed set and fate after open pollination. 090004 Two and three populations had on average significantly fewer pollen grains and pollen tubes per flower, respectively, than the fixed number of fertilizable ovules per fruit, while one population experienced significant pollen competition. Style length was positively correlated with the number of pollen tubes. While pollen availability was very variable, seed abortion was significantly less frequent in denser populations, and in one population the proportion of well-developed seeds was significantly, positively correlated with the number of pollen tubes in the style. 090004 Less pollen deposition, lower numbers of pollen tubes reaching the base of the style, lower pollen quality and therefore increased abortion of fertilized ovules can all reduce seed set in natural P. grandiflora stands. Substantial intraspecific variability implies that microgametophyte competition also occurs in this species. Finally, style morphology may affect pollen receipt.

Campbell DR ( 1989)

Measurements of selection in a hermaphroditic plant: Variation in male and female pollination success

Evolution, 43, 318-334.

DOI:10.1111/j.1558-5646.1989.tb04230.x      URL     PMID:28568547      [本文引用: 2]

I measured phenotypic selection of floral traits through both male and female functions of the hermaphroditic flowers of Ipomopsis aggregata (Pursh) V. Grant su...

Cresswell JE ( 2000)

Manipulation of female architecture in flowers reveals a narrow optimum for pollen deposition

Ecology, 81, 3244-3249.

DOI:10.2307/177415      URL     [本文引用: 1]

The invariability of animal-pollinated flowers relative to vegetative organs, such as leaves, suggests that these flowers may have been finely tuned for an optimum pollination performance. The evolutionary mechanism that could optimize and conserve a species' flower architecture is stabilizing selection, which occurs when intermediate phenotypes attain the highest fitness. Evidence for the operation of such a mechanism is rare in this context, however. Here, I report an experiment that used oil-seed rape (Brassica napus cv. Westar) as the basis of a model system in which the spatial disposition of flowers' sexual organs was manipulated in order to characterize the relationship between form and function. Specifically, individual flowers were manipulated by replacing a section of the style with a length of fine wire in order to produce flowers with pistils of various lengths. The pollination performance of these virgin, model flowers was then assessed by measuring the amount of pollen removed and deposited during a single bumblebee visit. There was no evidence that pollen removal was affected by pistil length. Variation in pistil length, however, was associated with a pronounced optimum in pollen deposition, which was attained by only a narrow range of pistil variants of intermediate length. These results suggest that stabilizing selection may maintain the architectural invariability of animal-pollinated flowers.

Chen JY, Bao MZ ( 1992)

Botanical classification and horticultural classification of Chinese Mei (Prunus mume) resources

Journal of Zhejiang Forest College, 9, 119-132. (in Chinese with English abstract)

URL     [本文引用: 2]

[ 陈俊愉, 包满珠 ( 1992)

中国梅(Prunus mume)的植物学分类和园艺学分类

浙江林学院学报, 9, 119-132.]

URL     [本文引用: 2]

Diggle PK ( 1995)

Architectural effects and the interpretation of patterns of fruit and seed development

Annual Review of Ecology and Systematics, 26, 531-552.

DOI:10.1146/annurev.es.26.110195.002531      URL     [本文引用: 2]

The commonly observed proximal-to-distal decrease within inflorescences of fruit and/or seed maturation per flower has frequently been attributed to competition among developing fruits for resources. The research summarized in this review suggests, however, that the observed variation can also be due to architecture--that is, to sources of variation inherent in plant axes. Thus, the fate of a developing flower depends not only on the reproductive events that have preceded it during the ontogeny of the organism, but on where it occurs within the architecture of an individual. The effects of architecture are separable experimentally from the effects of differential resource allocation, and careful experimental analysis of these two factors will enhance our understanding of the physiological, developmental, and evolutionary controls of fruit and seed production in flowering plants.

Diggle PK ( 1997)

Ontogenetic contingency and floral morphology: The effects of architecture and resource limitation

International Journal of Plant Sciences, 158, 99-107.

DOI:10.1086/297510      URL     [本文引用: 1]

Floral form and function commonly vary quantitatively and qualitatively within inflorescences. Overall floral size, the sizes of individual floral organs, and the frequency of successful fruit and/or seed maturation may decline distally within inflorescences. These frequently observed patterns of variation have been attributed to competition for limited resources among developing flowers and fruits. The variation, however, can also be due to architecture, that is, to sources of variation inherent in plant axes. Floral morphology may change with position on an inflorescence axis, even in the absence of resource competition. Experimental analyses of Solanum hirtum and Arabidopsis thaliana identify the separate effects of architectural and resource limitation on floral development and morphology. The effects of architecture may mimic, mask, or lead to misinterpretation of the effects of resource competition on floral form. Floral morphology and function may be characterized as ontogenetically contingent: the morphology and fate of a developing flower depend on the events that have preceded it during the ontogeny of the organism and on where it occurs within the architecture of the individual. Careful consideration of the role of ontogenetic contingency in diverse taxa will be required to fully understand patterns of variation in floral morphology and function within individuals and may ultimately enhance our understanding of morphological diversification among taxa.

Erbar C ( 2003)

Pollen tube transmitting tissue: Place of competition of male gametophytes

International Journal of Plant Sciences, 164, 265-277.

DOI:10.1086/377061      URL     [本文引用: 1]

In angiosperm reproduction, pollen tubes elongate from the stigma through the stylar transmitting tissue to the ovary to deliver the male gametes for fertilization. This article reviews different forms of pollen tube transmitting tracts within the style as well as possibilities for bridging the gap from the transmitting tissue to the ovule. The pollen tube transmitting tissue is the place of possible pollen tube competition. Often the number of pollen grains deposited onto the stigma or stigmata of a flower exceeds the number of ovules in the ovary of the pollinated flower. In the styles of most taxa investigated, a reduction in number of the pollen tubes occurs from the stigma to the ovary (pollen tube attrition): the pollen tubes compete for entry into the embryo sac of the ovules within the ovary. A prediction of the extent of pollen tube competition can be partially equated to the P-S/O ratio. This ratio relates the total number of pollen grains (P) deposited onto the stigmata (S) at the end of the female phase of anthesis (stigma receptivity) to the number of ovules (O) present in the ovary. Some quantitative data on pollen load on the stigma or stigmata and resulting pollen tube competition will be demonstrated especially in the choricarpous gynoecium of Geum rivale (Rosaceae) and in the coenocarpous gynoecium of Cichorium intybus (Asteraceae). Data from this study are compared with data of other taxa obtained from the literature.

Granado-Yela C, Balaguer L, Cayuela L, Méndez M ( 2017)

Unusual positional effects on flower sex in an andromonoecious tree: Resource competition, architectural constraints, or inhibition by the apical flower?

American Journal of Botany, 104, 608-615.

DOI:10.3732/ajb.1600433      URL     PMID:28428197      [本文引用: 1]

Abstract PREMISE OF THE STUDY: Two, nonmutually exclusive, mechanisms-competition for resources and architectural constraints-have been proposed to explain the proximal to distal decline in flower size, mass, and/or femaleness in indeterminate, elongate inflorescences. Whether these mechanisms also explain unusual positional effects such as distal to proximal declines of floral performance in determinate inflorescences, is understudied. METHODS: We tested the relative influence of these mechanisms in the andromonoecious wild olive tree, where hermaphroditic flowers occur mainly on apical and the most proximal positions in determinate inflorescences. We experimentally increased the availability of resources for the inflorescences by removing half of the inflorescences per twig or reduced resource availability by removing leaves. We also removed the apical flower to test its inhibitory effect on subapical flowers. KEY RESULTS: The apical flower had the highest probability of being hermaphroditic. Further down, however, the probability of finding a hermaphroditic flower decreased from the base to the tip of the inflorescences. An experimental increase of resources increased the probability of finding hermaphroditic flowers at each position, and vice versa. Removal of the apical flower increased the probability of producing hermaphroditic flowers in proximal positions but not in subapical positions. CONCLUSIONS: These results indicate an interaction between resource competition and architectural constraints in influencing the arrangement of the hermaphroditic and male flowers within the inflorescences of the wild olive tree. Subapical flowers did not seem to be hormonally suppressed by apical flowers. The study of these unusual positional effects is needed for a general understanding about the functional implications of inflorescence architecture. 2017 Botanical Society of America.

Hong Y, Fang HT, Na R ( 2006)

Pollen viability and stigma receptivity of Prunus mongolica Maxim

Guihaia, 26, 589-591. (in Chinese with English abstract)

DOI:10.3969/j.issn.1000-3142.2006.06.003      URL     [本文引用: 1]

用过氧化氢酶法测定蒙古扁桃花粉的活力和寿命,用联苯胺-过氧化氢法测定柱头可授性.结果表明,蒙古扁桃 花粉活力在散粉后第2天最高,3 d后活力迅速下降,三种类型的植株的花粉活力下降速度存在差异,其活力可持续30 d左右.短柱头植株柱头在开花后1 d可分泌黏液,长柱头和中柱头花植株的柱头在开花后2 d才分泌黏液,短柱头分泌黏液的持续时间较后者长.柱头可授性持续时间同种之间也存在差异,长柱头和中柱头植株可达6 d,而短柱头花植株可持续10 d左右.

[ 红雨, 方海涛, 那仁 ( 2006)

濒危植物蒙古扁桃花粉活力和柱头可授性研究

广西植物, 26, 589-591.]

DOI:10.3969/j.issn.1000-3142.2006.06.003      URL     [本文引用: 1]

用过氧化氢酶法测定蒙古扁桃花粉的活力和寿命,用联苯胺-过氧化氢法测定柱头可授性.结果表明,蒙古扁桃 花粉活力在散粉后第2天最高,3 d后活力迅速下降,三种类型的植株的花粉活力下降速度存在差异,其活力可持续30 d左右.短柱头植株柱头在开花后1 d可分泌黏液,长柱头和中柱头花植株的柱头在开花后2 d才分泌黏液,短柱头分泌黏液的持续时间较后者长.柱头可授性持续时间同种之间也存在差异,长柱头和中柱头植株可达6 d,而短柱头花植株可持续10 d左右.

Huang SQ, Guo YH ( 2000)

New advances in pollination biology

Chinese Science Bulletin, 45, 225-237. (in Chinese)

DOI:10.3321/j.issn:0023-074X.2000.03.001      [本文引用: 1]

传地生物学是研究与传粉事件有关的种种生物学特性及其规律的一门学科,是植物生殖生态学和进化生物学关注的焦点之一,概述了近年来的主要研究热点及其进展,包括植物性表达的多样性及其进化、传粉者与花部特征的相互作用及其进化机制、传粉系统的进化、花粉竞争的生物学含义、花粉流的标记方法及散布规律的研究等5个方面,介绍了中国传粉生物学研究的现状及不足之处,结合国际上的发展趋势已由观察的、静态的描述传粉现象的阶段过

[ 黄双全, 郭友好 ( 2000)

传粉生物学的研究进展

科学通报, 45, 225-237.]

DOI:10.3321/j.issn:0023-074X.2000.03.001      [本文引用: 1]

传地生物学是研究与传粉事件有关的种种生物学特性及其规律的一门学科,是植物生殖生态学和进化生物学关注的焦点之一,概述了近年来的主要研究热点及其进展,包括植物性表达的多样性及其进化、传粉者与花部特征的相互作用及其进化机制、传粉系统的进化、花粉竞争的生物学含义、花粉流的标记方法及散布规律的研究等5个方面,介绍了中国传粉生物学研究的现状及不足之处,结合国际上的发展趋势已由观察的、静态的描述传粉现象的阶段过

Johnson MTJ, Smith SD, Rausher MD ( 2010)

Effects of plant sex on range distributions and allocation to reproduction

New Phytologist, 186, 769-779.

DOI:10.1111/j.1469-8137.2010.03201.x      URL     PMID:20180909      [本文引用: 2]

090004Despite an abundance of theory, few empirical studies have explored the ecological and evolutionary consequences of sex. We used a comparative phylogenetic approach to examine whether transitions between sexual and asexual reproduction are associated with changes in the size and distribution of species' geographical ranges, and their investment in reproduction.090004Here, we reconstructed the phylogeny of the genus Oenothera sections Oenothera and Calylophus (Onagraceae), which contain 35 sexual and 30 functionally asexual species. From each species, we collected data on the geographical distribution and variation in plant traits related to reproduction.090004Functionally asexual species occurred at higher latitudes, but did not differ in range size, compared with sexual species. Transitions to asexuality were associated with decreased investment in floral structures, including the length of petals, floral tubes and styles. Decreased anther size and increased seed size within asexual species also suggest altered allocation to male and female fitness.090004The observed range shifts are consistent with superior colonization of environments by asexual species following glaciation, and the observed changes in reproductive allocation support predictions made by models relating to the evolution of selfing. Our results suggest that the evolutionary consequences of asexual reproduction might be less restrictive than previously thought.

Jürgens A, Witt T, Gottsberger G ( 2012)

Pollen grain size variation in Caryophylloideae: A mixed strategy for pollen deposition along styles with long stigmatic areas?

Plant Systematics and Evolution, 298, 9-24.

DOI:10.1007/s00606-011-0518-z      URL     [本文引用: 1]

AbstractThe literature supports a positive correlation between pollen size and style length and that pollen size may be affected by a trade-off with pollen number. A correlation between pollen size and pollinator type has been hypothesized by some authors but not confirmed. To assess these associations, pollen grain number, pollen grain diameter and style length were measured in 53 species of Caryophylloideae that had been classified beforehand as being diurnal, nocturnal or self-pollinated. Surprisingly, many species showed two, and a few even three, distinct pollen grain size classes per individual anther. Therefore, further analyses were based on the different pollen size classes and respective pollen grain numbers. Firstly, we found no indication of a trade-off between pollen size and number. Secondly, a positive linear correlation between style length and pollen grain diameter was found using conventional analysis methods and phylogenetically independent contrasts. Thirdly, when using conventional analysis methods a strong relationship between style length and pollination mode was found, with nocturnal species having significantly longer styles and larger pollen grains than diurnal and selfing species. These results were not supported, however, when calculating comparisons of phylogenetically independent pairs. We discuss our findings in relation to intra-specific pollen polymorphism and pollen degeneration, both of which have been associated with different pollen grain sizes in Caryophylloideae. Furthermore, we discuss the possibility that pollen polymorphism might be a mixed strategy to optimise the success of pollen grains deposited at different pollen vule distances on long styles.

Lankinen Å, Skogsmyr I ( 2001)

Evolution of pistil length as a choice mechanism for pollen quality

Oikos, 92, 81-90.

DOI:10.1034/j.1600-0706.2001.920110.x      URL     [本文引用: 2]

During the fertilisation process in plants, pollen tube growth rate may be selected as a trait important in male to male competition. Since female morphology provides the necessary selective arena for such competition, we investigate if sexual selection theory can be used to explain the evolution of pistil length as a female choice mechanism. This choice is performed by direct interference with male to male competition. Furthermore, the sessile nature of plants limits the number of mates a female can choose between, which could limit the benefit a female can gain from distinguishing between donors. To mirror these circumstances, we model a situation when there are only two competitors at a time. Using a game theoretical approach we show that if pollen tube growth rate can be used as an indication of heritable quality, pistil length can be selected in response to variation of this trait. We further find that length of the pistil affects selection of pollen tube growth rate. Thus female preference and male competitive ability co-evolve, but this does not necessarily lead to a positive relationship between the two. Under certain circumstances we find a negative relation instead. Given realistic differences in male quality, the model indicates that there is a potential for evolution of female morphology as a choice mechanism for pollen quality.

Liu CQ, Huang SQ ( 2012)

Does the relative importance of resource competition and architectural effect in floral variation vary with stages of floral ontogeny?

Journal of Systematics and Evolution, 50, 119-124.

DOI:10.1111/j.1759-6831.2011.00175.x      URL     [本文引用: 1]

Abstract Variation in floral allocation within inflorescences has been attributed to resource competition and/or architectural effect. The two hypotheses were extensively studied and both hypotheses were partly supported by previous studies. However, the relative importance of resource competition and architectural effect may vary with stages of floral ontogeny in a species. To detect the effects of ontogenetic stages on floral variation, we manipulated the 6-flowered inflorescences of Adenophora jasionifolia (Campanulaceae) by early, late, and non-thinning flowers. Our results indicated that pollen and ovule production of the remaining flowers were not significantly different between early and late thinning manipulations, suggesting that floral sex allocation was determined far before flowering, in support of the architectural effect hypothesis. Under hand-pollination treatments, early thinning but not late thinning resulted in a significant increase in seed number and seed set of the remaining flowers. Therefore, the increase in seed production of the remaining flowers related to the floral ontogeny. The resource competition for floral allocation was significant under early thinning rather than late thinning manipulation. Our results suggest that studies on floral variation within inflorescences should take into account the stages of floral ontogeny.

Lloyd DG ( 1980)

Sexual strategies in plants. I. An hypothesis of serial adjustment of maternal investment during one reproductive session

New Phytologist, 86, 69-79.

DOI:10.1111/j.1469-8137.1980.tb00780.x      URL     [本文引用: 2]

It is postulated that in one reproductive session the level of maternal expenditure of an angiosperm plant is determined by a temporal series of controls on the number of potential fruit in which an investment is made. The serial adjustment hypothesis has three parts: A. The amount of maternal expenditure is regulated at many developmental units, particularly single flowers and fruit, at three principal sequential stages the determination of flowers, the development of ovaries and the maturation of fruit. B. At each stage, the initiation or continuation of an investment requires an amount of available resources above a certain threshold. Hence maternal expenditure is continually adjusted to the resources available at each developmental site. C. The pattern of controls that maximizes the maternal fitness of a plant is selected. Many factors affect the relative advantages of regulation at the three stages, therefore species and sexual morphs appear to vary widely in their proportional use of the three stages. Regulation of flower determination has the general advantages of offering bidirectional adjustment of maternal investment, maintaining a constant ratio of maternal to paternal investment, and reducing wasted expenditure. The later stages, ovary and fruit regulation, allow secondary adjustments of maternal investment in unpredictable circumstances and permit adjustment of the relative numbers of polliniferous and seminiferous flowers. The relative advantages of restricting the numbers of developing ovaries, or of maturing fruit, depend principally on the extent to which differences in the capacity of flowers to mature fruit are evident before anthesis.

Love J, Graham SW, Irwin JA, Ashton PA, Bretagnolle F, Abbott RJ ( 2016)

Self-pollination, style length development and seed set in self-compatible Asteraceae: Evidence from Senecio vulgaris L

Transactions of the Botanical Society of Edinburgh, 9, 371-379.

[本文引用: 2]

Lu LD, Bruce B ( 2003)

Armeniaca. In: Flora of China (eds Wu ZY, Raven PH, Hong DY)

Vol. 9, pp. 396-401. Science Press, Beijing & Missouri Botanical Garden Press St. Louis.

[本文引用: 1]

Ma WJ, Zhang FP, Peng YQ, Yang DR ( 2009)

Comparison of style length and reproduction success in Ficus of different breeding systems

Chinese Journal of Plant Ecology, 33, 911-918. (in Chinese with English abstract)

DOI:10.3773/j.issn.1005-264x.2009.05.010      URL     [本文引用: 1]

多年来,不同繁育系统的榕树(Ficus spp.)的进化问题引起生物学家们极大的兴趣。前人通过对不同繁育系统榕树雌花的花柱长度、传粉榕小蜂产卵器长度和繁殖率的比较,推测榕树的雌雄异株是由雌雄同株进化而来的。为验证这一推论,选取雌雄同株的垂叶榕(Ficus benjamina)和钝叶榕(F.curtipes)以及雌雄异株的斜叶榕(F.tinctoria)和鸡嗉子榕(F.semicordata),进行了雌花花柱长度、传粉榕小蜂产卵器长度及繁殖率的比较。研究结果显示:1)两种雌雄同株榕树的传粉榕小蜂(Eupristina koningsbergeri和Eupristina sp.)的产卵器长度,显著长于两种雌雄异株榕树的传粉榕小蜂(Liporrhopalum gibbosae和Ceratosolen gravelyi)产卵器的长度,且雌雄同株榕树雌花花柱长度的变异大于雌雄异株雌花花柱长度的变异;2)两种雌雄同株榕树的雌花花柱长度呈单峰分布,且花柱长度的变异大于传粉榕小蜂产卵器长度的变异;两种雌雄异株榕树花序的雌花花柱长度呈双峰分布,雌花花柱长度的变异也大于传粉榕小蜂产卵器的长度变异;3)两种雌雄同株榕树的传粉榕小蜂产卵器长度能到达雌花子房的比例更高,可更有效地产生瘿花,而雌雄异株榕树产生种子的优势更强。研究表明:在传粉榕小蜂产卵器长度及花序雌花花柱分布方面,雌雄异株榕树有着明显的优势,但在繁殖率方面没有绝对优势。因此,前人从雌花花柱分布及繁殖率比较而做出榕树进化的推测并不正确。要真正解决榕树的进化问题,需要多学科、多指标的综合分析和深入研究。

[ 马文娟, 张凤萍, 彭艳琼, 杨大荣 ( 2009)

不同繁育系统的榕树雌花的花柱长度和繁殖率比较

植物生态学报, 33, 911-918.]

DOI:10.3773/j.issn.1005-264x.2009.05.010      URL     [本文引用: 1]

多年来,不同繁育系统的榕树(Ficus spp.)的进化问题引起生物学家们极大的兴趣。前人通过对不同繁育系统榕树雌花的花柱长度、传粉榕小蜂产卵器长度和繁殖率的比较,推测榕树的雌雄异株是由雌雄同株进化而来的。为验证这一推论,选取雌雄同株的垂叶榕(Ficus benjamina)和钝叶榕(F.curtipes)以及雌雄异株的斜叶榕(F.tinctoria)和鸡嗉子榕(F.semicordata),进行了雌花花柱长度、传粉榕小蜂产卵器长度及繁殖率的比较。研究结果显示:1)两种雌雄同株榕树的传粉榕小蜂(Eupristina koningsbergeri和Eupristina sp.)的产卵器长度,显著长于两种雌雄异株榕树的传粉榕小蜂(Liporrhopalum gibbosae和Ceratosolen gravelyi)产卵器的长度,且雌雄同株榕树雌花花柱长度的变异大于雌雄异株雌花花柱长度的变异;2)两种雌雄同株榕树的雌花花柱长度呈单峰分布,且花柱长度的变异大于传粉榕小蜂产卵器长度的变异;两种雌雄异株榕树花序的雌花花柱长度呈双峰分布,雌花花柱长度的变异也大于传粉榕小蜂产卵器的长度变异;3)两种雌雄同株榕树的传粉榕小蜂产卵器长度能到达雌花子房的比例更高,可更有效地产生瘿花,而雌雄异株榕树产生种子的优势更强。研究表明:在传粉榕小蜂产卵器长度及花序雌花花柱分布方面,雌雄异株榕树有着明显的优势,但在繁殖率方面没有绝对优势。因此,前人从雌花花柱分布及繁殖率比较而做出榕树进化的推测并不正确。要真正解决榕树的进化问题,需要多学科、多指标的综合分析和深入研究。

Mazer SJ, Dudley LS, Hove AA, Emms SK, Verhoeven AS ( 2010)

Physiological performance in Clarkia sister taxa with contrasting mating systems: Do early-flowering autogamous taxa avoid water stress relative to their pollinator-dependent counterparts?

International Journal of Plant Sciences, 171, 1029-1047.

DOI:10.1086/656305      URL     [本文引用: 1]

Mazer SJ, Moghaddasi A, Bello AK, Hove AA ( 2016)

Winning in style: Longer styles receive more pollen, but style length does not affect pollen attrition in wild Clarkia populations

American Journal of Botany, 103, 408-422.

DOI:10.3732/ajb.1500192      URL     PMID:26933011      [本文引用: 1]

Abstract PREMISE OF THE STUDY: One proposed function of long styles is to intensify selection among male gametophytes relative to short styles. If so, given sufficient competition, longer styles will have higher rates of pollen tube attrition (failure to reach the style base) within the style than shorter ones. Alternatively, style length may influence pollen receipt, which itself may affect attrition rates. METHODS: We tested these predictions by collecting senescing styles from wild populations of two insect-pollinated Clarkia species. We examined the number of pollen grains adhering to the stigma, length of styles, and rates of attrition from the stigma surface to the stigma-style junction (SSJ), from the SSJ to the style base, and from the stigma surface to the style base. Multivariate analyses estimated the independent effects of pollen grains per stigma, the number of pollen tubes at the SSJ, and style length on attrition. KEY RESULTS: Style length was generally positively correlated with pollen receipt, and the number of pollen grains per stigma was positively correlated with all three attrition rates. In neither species was any attrition rate affected by style length independent of the number of pollen grains per stigma. CONCLUSIONS: Pollen attrition was mediated by style length, but the function of style length was primarily to increase the number of germinating pollen grains, which affected attrition rates either through stigma clogging or pollen-pollen interactions. Style length may have a direct effect on pollen receipt due to the stigma's position relative to pollinator body parts, but traits correlated with style length may also directly affect pollen receipt. 2016 Botanical Society of America.

Mulcahy DL ( 1979)

The rise of the angiosperms: A genecological factor

Science, 206, 20-23.

DOI:10.1126/science.206.4414.20      URL     [本文引用: 1]

Plitmann U ( 1993)

Pollen tube attrition as related to breeding systems in Brassicaceae

Plant Systematics and Evolution, 188, 65-72.

DOI:10.1007/BF00937836      URL     [本文引用: 1]

In order to estimate pollen tube attrition and to relate it to the breeding system, numbers of microgametophytes within the stigma and at the bottom of the style were scored in species of the Brassicaceae, 17 sporophytically self-incompatible, 11 predominant inbreeders, 4 facultative outcrossers. Attrition was evident in almost all species. The mean number of tubes at the bottom of the style, and mean number of stylar tubes per ovule were significantly lower in outbreeders. The degree of attrition, measured as "attrition rate" (the proportion of tubes in the stigma that do not reach the bottom of the style) was higher in outbreeders; it was positively correlated with the pollen: ovule ratios as estimates of the degree of outbreeding in the species involved. In outbreeders, sexual selection seems to be rather effective in inhibiting certain microgametophytic genotypes from reaching the ovules. Because of the low number of tubes per ovule in most cases, we assume that, in the Brassicaceae, female choice may be more effective in sexual selection and in ensuing pollen tube attrition.

Ramesha BT, Yetish MD, Ravikanth G, Ganeshaiah KN, Ghazoul J, Shaanker RU ( 2011)

Stylish lengths: Mate choice in flowers

Journal of Biosciences, 36, 229-234.

DOI:10.1007/s12038-011-9057-6      URL     PMID:21654077      [本文引用: 2]

The styles of flowers may represent an arena for pollen competition in the race to fertilize ovules. Accordingly, selection should favour a longer ‘race’ to better discriminate among variable pollen by increasing style length. Sampling across a taxonomically diverse range of wild and outcrossed species, we found that the distribution of style lengths within plants were skewed towards longer styles, as predicted. In self-pollinated domesticated species, where discrimination among pollen is less important, we found no such pattern. We conclude that style length is under directional selection towards longer styles as a mechanism for mate choice among pollen of variable quality.

Roulston TH, Cane JH, Buchmann SL ( 2000)

What governs protein content of pollen: Pollinator preferences, pollen- pistil interactions, or phylogeny?

Ecological Monographs, 70, 617-643.

[本文引用: 1]

Stephenson AG, Travers SE, Mena-Ali JI, Winsor JA ( 2003)

Pollen performance before and during the autotrophic- heterotrophic transition of pollen tube growth

Philosophical Transactions of the Royal Society of London B: Biological Sciences, 358, 1009-1018.

DOI:10.1098/rstb.2003.1290      URL     [本文引用: 1]

Taylor ML, Williams JH ( 2012)

Pollen tube development in two species of Trithuria (Hydatellaceae) with contrasting breeding systems

Sexual Plant Reproduction, 25, 83-96.

DOI:10.1007/s00497-012-0183-6      URL     PMID:22367232      [本文引用: 1]

AbstractTrithuriasubmersa. . . . submersa has been accompanied by predictable modifications to reproductive development, which, because of the ancient relationship between Hydatellaceae and all other angiosperms, suggests that traits underlying the lability of flowering plant post-pollination biology were present early in their history.

Ushimaru A, Itagaki T, Ishii HS ( 2003)

Variation in floral organ size depends on function

Evolutionary Ecology Research, 5, 615-622.

[本文引用: 1]

Wang XF, Armbruster WS, Huang SQ ( 2012)

Extra-gynoecial pollen-tube growth in apocarpous angiosperms is phylogenetically widespread and probably adaptive

New Phytologist, 193, 253-260.

DOI:10.1111/j.1469-8137.2011.03912.x      URL     [本文引用: 1]

http://doi.wiley.com/10.1111/j.1469-8137.2011.03912.x

Wang XY, Quan QM, Wang B, Li YX, Huang SQ ( 2018)

Pollen competition between morphs in a pollen-color dimorphic herb and the loss of phenotypic polymorphism within populations

Evolution, 72, 785-797.

DOI:10.1111/evo.13445      URL     PMID:29399790      [本文引用: 1]

Abstract Flower color polymorphism is relatively uncommon in natural flowering plants, suggesting that maintenance of different color morphs within populations is difficult. To address the selective mechanisms shaping pollen-color dimorphism, pollinator preferences and reproductive performance were studied over three years in Epimedium pubescens in which some populations had plants with either green or yellow pollen (and anthers). Visitation rate and pollen removal and receipt by the bee pollinator ( Andrena emeishanica ) did not differ between the two color morphs. Compared to the green morph, siring success of the yellow morph's pollen was lower, but that of mixtures of pollen from green and yellow morphs was lowest. This difference, corresponding to in and ex vivo experiments on pollen performance, indicated that pollen germination, rather than tube growth, of the green morph was higher than that of the yellow morph and was seriously constrained in both morphs if a pollen competitor was present. A rare green morph may invade a yellow-morph population, but the co-existence of pollen color variants is complicated by the reduced siring success of mixed pollinations. Potential pollen competition between morphs may have discouraged the maintenance of multiple phenotypes within populations, a cryptic mechanism of competitive exclusion. This article is protected by copyright. All rights reserved

Wang YF, Gao HY, Shi HY, Wang JH, Du GZ ( 2008)

Adaptive significance of Saussurea parviflora's sexual organs, Qinghai-Tibetan Plateau, China

Journal of Plant Ecology (Chinese Version), 32, 379-384. (in Chinese with English abstract)

DOI:10.3773/j.issn.1005-264x.2008.02.015      URL     [本文引用: 1]

小花风毛菊(Saussurea paniflora)是菊科风毛菊属的一种多年生草本植物,是该属在青藏高原东缘的一个优势种。该文研究了小花风毛菊11个居群性器官的变异与11个海拔高度(2100~3500m)之间的相关性,以及花丝、花药长和花柱、花柱分枝长与花粉数之间的相关性;探讨了小花风毛菊的性器官是如何随海拔高度发生变异,如何与胁迫环境相适应,实现交配成功,提高繁殖成功率,使其成为该属植物“现代分布中心”的优势种。结果表明:1)花丝、花药长与海拔高度呈极显著的正相关关系(P〈0.01);2)花粉数与海拔高度呈极显著的负相关关系(P〈0.01);3)花柱、花柱分枝长及花柱分枝长和雌蕊长的比与海拔高度呈极显著的正相关关系(P〈0.01);4)花柱、花柱分枝长和花丝长与花粉数呈显著的负相关关系(P〈0.05);5)种子的成熟率与海拔高度呈显著的正相关关系(P〈0.05)。说明小花风毛菊种内性器官在特定的生存环境下发生了变异,特别是在花粉数随海拔的升高明显减少以及昆虫的多样性、丰富性和活动性明显降低的情况下,它能够通过花柱分枝长度的变异来提高柱头对传粉昆虫的感受性,能通过种子成熟率的提高来保证高海拔数目显著减少的种子的成功发育,进而在一定程度上导致了该物种繁殖成功率的上升,使它成为青藏高原这一胁迫环境中的优势种。

[ 王一峰, 高宏岩, 施海燕, 王剑虹, 杜国祯 ( 2008)

小花风毛菊的性器官在青藏高原的海拔变异

植物生态学报, 32, 379-384.]

DOI:10.3773/j.issn.1005-264x.2008.02.015      URL     [本文引用: 1]

小花风毛菊(Saussurea paniflora)是菊科风毛菊属的一种多年生草本植物,是该属在青藏高原东缘的一个优势种。该文研究了小花风毛菊11个居群性器官的变异与11个海拔高度(2100~3500m)之间的相关性,以及花丝、花药长和花柱、花柱分枝长与花粉数之间的相关性;探讨了小花风毛菊的性器官是如何随海拔高度发生变异,如何与胁迫环境相适应,实现交配成功,提高繁殖成功率,使其成为该属植物“现代分布中心”的优势种。结果表明:1)花丝、花药长与海拔高度呈极显著的正相关关系(P〈0.01);2)花粉数与海拔高度呈极显著的负相关关系(P〈0.01);3)花柱、花柱分枝长及花柱分枝长和雌蕊长的比与海拔高度呈极显著的正相关关系(P〈0.01);4)花柱、花柱分枝长和花丝长与花粉数呈显著的负相关关系(P〈0.05);5)种子的成熟率与海拔高度呈显著的正相关关系(P〈0.05)。说明小花风毛菊种内性器官在特定的生存环境下发生了变异,特别是在花粉数随海拔的升高明显减少以及昆虫的多样性、丰富性和活动性明显降低的情况下,它能够通过花柱分枝长度的变异来提高柱头对传粉昆虫的感受性,能通过种子成熟率的提高来保证高海拔数目显著减少的种子的成功发育,进而在一定程度上导致了该物种繁殖成功率的上升,使它成为青藏高原这一胁迫环境中的优势种。

Zhang CX, Tateishi N, Tanabe K ( 2010)

Pollen density on the stigma affects endogenous gibberellin metabolism, seed and fruit set, and fruit quality in Pyrus pyrifolia

Journal of Experimental Botany, 61, 4291-4302.

DOI:10.1093/jxb/erq232      URL     [本文引用: 1]

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