艳丽耳草的二型花柱及异型自交不亲和系统

doi:10.3724/SP.J.1003.2012.11243

生物多样性 ›› 2012, Vol. 20 ›› Issue (3): 337-347. DOI: 10.3724/SP.J.1003.2012.11243

所属专题：传粉生物学：理论探讨与初步实践；传粉生物学

艳丽耳草的二型花柱及异型自交不亲和系统

刘欣欣¹^,², 吴小琴¹^,^*(), 张奠湘¹

1 中国科学院植物资源保护与可持续利用重点实验室, 中国科学院华南植物园, 广州 510650
2 中国科学院研究生院, 北京 100049

收稿日期:2011-12-30 接受日期:2012-04-10 出版日期:2012-05-20 发布日期:2012-05-09
通讯作者: 吴小琴
作者简介:*E-mail: wxq@scbg.ac.cn
基金资助:
国家自然科学基金(31070161);国家自然科学基金(31170295);中国科学院知识创新工程资助项目(KSCX2-EW-J-28)

Distyly and heteromorphic self-incompatibility of Hedyotis pulcherrima (Rubiaceae)

Xinxin Liu¹^,², Xiaoqin Wu¹^,^*(), Dianxiang Zhang¹

1 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650
2 Graduate University of the Chinese Academy of Sciences, Beijing 100049

Received:2011-12-30 Accepted:2012-04-10 Online:2012-05-20 Published:2012-05-09
Contact: Xiaoqin Wu

摘要/Abstract

摘要：

异型花柱是一种受遗传因素控制的花型多态性现象, 包括二型花柱和三型花柱两种类型。本文以茜草科艳丽耳草(Hedyotis pulcherrima)为实验材料, 通过对其野外居群的花型、花部形态及花粉特征等观察, 发现艳丽耳草野外居群同时存在长花柱型花和短花柱型花, 长/短花柱型花的数量比例为1:1。两型花具有精确的交互式雌雄异位特征, 并且该特征与花冠长度相关性显著。长/短花柱型花的柱头裂片长度、花粉大小及淀粉含量等具有二型性。花粉体外培养时花粉萌发率及花粉管生长速率无显著性差异。人工授粉后, 艳丽耳草长/短花柱型花型间异交花粉管生长形态正常, 授粉24 h后花粉管均已进入子房。而长/短花柱型花在自交及型内异交下均表现为不亲和, 花粉管生长停止于柱头, 花粉管顶端累积胼胝质并膨大。艳丽耳草没有无融合生殖现象, 型间人工辅助异交授粉结实率为100%, 显著高于自然结实率。本研究结果表明, 艳丽耳草是典型的二型花柱植物, 并具有异型自交不亲和系统。

关键词: 二型花柱, Hedyotis pulcherrima, 异型自交不亲和, 繁育系统

Abstract

Heterostyly is a genetically controlled floral polymorphism, which includes both distyly and tristyly. We investigated morph ratio, floral and pollen morphology, and self-incompatibility of Hedyotis pulcherrima. Overall, the natural population of H. pulcherrima was isoplethic, containing long-styled and short-styled morphs with an equilibrium 1:1 ratio. Long-styled and short-styled morph exhibited a precise reciprocal herkogamy, which was significantly correlated with corolla length. Stigma-lobe length, pollen size, and starch content in pollen grains were dimorphic in the two morphs of H. pulcherrima, whereas pollen germination and pollen tube growth in vitro were not significantly different between the two morphs. Artificial pollination revealed that pollen tube shape was normal in both morphs where pollen tubes reached the ovary 24 h after pollination. However, pollen tube growth was arrested in the stigma with the accumulation of callose in the swollen tips in two morphs with self and intramorph pollination, indicating strict heteromorphic self-incompatibility in H. pulcherrima. No fruit was produced in emasculated netted flowers, suggesting the absence of apomixis. Artificial intermorph pollination resulted in 100% of fruit set, significantly higher than those with open pollination. Our results indicate that H. pulcherrima is a typically distylous species with heteromorphic self-incompatibility.

Key words: distyly, Hedyotis pulcherrima, heteromorphic self-incompatibility, breeding system

刘欣欣, 吴小琴, 张奠湘 (2012) 艳丽耳草的二型花柱及异型自交不亲和系统. 生物多样性, 20, 337-347. DOI: 10.3724/SP.J.1003.2012.11243.

Xinxin Liu, Xiaoqin Wu, Dianxiang Zhang (2012) Distyly and heteromorphic self-incompatibility of Hedyotis pulcherrima (Rubiaceae). Biodiversity Science, 20, 337-347. DOI: 10.3724/SP.J.1003.2012.11243.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2012.11243

https://www.biodiversity-science.net/CN/Y2012/V20/I3/337

图/表 9

图1 艳丽耳草花部特征参数测量。(1)花冠长度; (2)柱头高度; (3)花柱长度; (4)柱头裂片长度; (5)花药高度; (6)花药长度; (7)花丝长度; (8)花丝基部至花冠基部距离; (9)花丝基部至花冠顶端距离; (10)子房高度。

Fig.1 Floral measurements of Hedyotis pulcherrima. (1) Corolla length; (2) Stigma height; (3) Style height; (4) Stigma lobe length; (5) Anther height; (6) Anther length; (7) Filament length; (8) From filament attachment to the base of corolla; (9) From filament attachment to the apex of corolla; (10) Ovary height.

图2 艳丽耳草的二型花。(A)长花柱型; (B)短花柱型; (C)长/短花柱花交互式雌雄异位, 左边是长花柱型花, 右边是短花柱型花。St: 柱头; An: 花药; 标尺 = 3 mm。

Fig. 2 Distylous flower of Hedyotis pulcherrima. (A) Long-styled flower; (B) Short-styled flower; (C) Reciprocal herkogamy, the left is long-styled flower, and the right is short-styled flower. St, Stigma; An, Anther. Bar = 3 mm.

表1 艳丽耳草长/短花柱型花部及花粉特征参数

Table 1 Floral and pollen attributes of long-styled and short-styled flowers of Hedyotis pulcherrima

花部和花粉特征 Floral and pollen attributes	长花柱型 Long-styled morph (n = 30)	短花柱型 Short-styled morph (n = 30)	t-检验 t-test
花冠长度 Corolla length (mm)	18.34±1.21	18.55±1.65	0.56
柱头高度 Stigma height (mm)	15.13±0.94	11.18±1.03	15.47^**
花柱长度 Style height (mm)	13.61±0.89	8.60±0.82	61.63^**
柱头裂片长度 Stigma lobe length (mm)	1.63±0.25	2.83±0.46	12.48^**
花药高度 Anther height (mm)	11.12±0.76	14.62±1.27	12.94^**
花药长度 Anther length (mm)	2.11±0.18	2.38±0.16	6.10^**
花丝长度 Filament length (mm)	0.42±0.11	1.22±0.14	24.93^**
花丝基部至花冠基部距离 From filament attachment to the base of corolla (mm)	9.84±0.70	12.42±1.28	9.69^**
花丝基部至花冠顶端距离 From filament attachment to the apex of corolla (mm)	8.31±0.65	6.25±0.66	12.15^**
子房高度 Ovary height (mm)	2.11±0.19	2.22±0.26	1.83^*
花粉数 No. of pollen grains per flower	8,980±144.05	7,988±181.85	9.56^**
胚珠数 No. of ovules per flower	79.2±6.75	83.6±5.78	2.09^*
花粉/胚珠比 P/O ratio	113.51±6.88	95.59±1.43	10.75^**
花粉极轴长度 Polar axis of pollen (P) (μm)	24.89±1.42	27.18±2.16	4.85^**
花粉赤道长度 Equatorial axis of pollen (E) (μm)	23.38±1.65	25.02±1.35	4.2^**
花粉极轴长/赤道轴长 (P/E)	1.05±0.06	1.03±0.07	1.19
网孔直径 Diameter of lumina (μm)	0.56±0.11	0.51±0.08	0.39

表2 艳丽耳草长/短花柱型花的花部特征间的Pearson相关性分析

Table 2 Pearson correlation coefficients for traits of long-styled and short-styled flowers in Hedyotis pulcherrima

	花冠长度 Corolla length	柱头高度 Stigma height	花柱长度Style height	柱头裂片长度Stigma lobe length	花药高度 Anther height	花药长度 Anther length	花丝长度Filament length	花丝基部至花冠基部距离 From filament attachment to the base of corolla	花丝基部至花冠顶端距离From filament attachment to the apex of corolla
长花柱型 Long-styled morph
柱头高度 Stigma height	0.566^**
花柱长度 Style height	0.501^**	0.935^**
柱头裂片长度 Stigma lobe length	0.373^*	0.540^**	0.206
花药高度 Anther height	0.796^**	0.452^*	0.417^*	0.257
花药长度 Anther length	0.278	0.344	0.196	0.483^**	0.248
花丝长度 Filament length	0.475^**	0.261	0.286	0.040	0.368^*	0.184
花丝基部至花冠基部距离 From filament attachment to the base of corolla	0.836^**	0.391^*	0.327	0.301	0.934^**	0.231	0.323
花丝基部至花冠顶端距离 From filament attachment to the apex of corolla	0.723^**	0.442^*	0.429^*	0.202	0.377^*	0.319	0.325	0.411^*
子房高度 Ovary height	0.510^**	0.533^**	0.498^**	0.290	0.263	0.203	0.492^**	0.199	0.495^**
短花柱型 Short-styled morph
柱头高度 Stigma height	0.723^**
花柱长度 Style height	0.606^**	0.873^**
柱头裂片长度 Stigma lobe length	0.490^**	0.623^**	0.162
花药高度 Anther height	0.825^**	0.620^**	0.583^**	0.319
花药长度 Anther length	-0.206	0.056	0.058	0.020	-0.109
花丝长度 Filament length	0.135	0.080	0.085	0.025	0.059	-0.379^*
花丝基部至花冠基部距离 From filament attachment to the base of corolla	0.902^**	0.732^**	0.620^**	0.486^**	0.875^**	-0.238	0.154
花丝基部至花冠顶端距离 From filament attachment to the apex of corolla	0.748^**	0.492^**	0.391^*	0.369^*	0.426^*	-0.276	0.297	0.493^**
子房高度 Ovary height	0.336	0.307	0.250	0.221	0.046	0.091	0.125	0.097	0.501^**

表2 艳丽耳草长/短花柱型花的花部特征间的Pearson相关性分析

Table 2 Pearson correlation coefficients for traits of long-styled and short-styled flowers in Hedyotis pulcherrima

	花冠长度 Corolla length	柱头高度 Stigma height	花柱长度Style height	柱头裂片长度Stigma lobe length	花药高度 Anther height	花药长度 Anther length	花丝长度Filament length	花丝基部至花冠基部距离 From filament attachment to the base of corolla	花丝基部至花冠顶端距离From filament attachment to the apex of corolla
长花柱型 Long-styled morph
柱头高度 Stigma height	0.566^**
花柱长度 Style height	0.501^**	0.935^**
柱头裂片长度 Stigma lobe length	0.373^*	0.540^**	0.206
花药高度 Anther height	0.796^**	0.452^*	0.417^*	0.257
花药长度 Anther length	0.278	0.344	0.196	0.483^**	0.248
花丝长度 Filament length	0.475^**	0.261	0.286	0.040	0.368^*	0.184
花丝基部至花冠基部距离 From filament attachment to the base of corolla	0.836^**	0.391^*	0.327	0.301	0.934^**	0.231	0.323
花丝基部至花冠顶端距离 From filament attachment to the apex of corolla	0.723^**	0.442^*	0.429^*	0.202	0.377^*	0.319	0.325	0.411^*
子房高度 Ovary height	0.510^**	0.533^**	0.498^**	0.290	0.263	0.203	0.492^**	0.199	0.495^**
短花柱型 Short-styled morph
柱头高度 Stigma height	0.723^**
花柱长度 Style height	0.606^**	0.873^**
柱头裂片长度 Stigma lobe length	0.490^**	0.623^**	0.162
花药高度 Anther height	0.825^**	0.620^**	0.583^**	0.319
花药长度 Anther length	-0.206	0.056	0.058	0.020	-0.109
花丝长度 Filament length	0.135	0.080	0.085	0.025	0.059	-0.379^*
花丝基部至花冠基部距离 From filament attachment to the base of corolla	0.902^**	0.732^**	0.620^**	0.486^**	0.875^**	-0.238	0.154
花丝基部至花冠顶端距离 From filament attachment to the apex of corolla	0.748^**	0.492^**	0.391^*	0.369^*	0.426^*	-0.276	0.297	0.493^**
子房高度 Ovary height	0.336	0.307	0.250	0.221	0.046	0.091	0.125	0.097	0.501^**

图3 艳丽耳草长/短花柱型花粉、花冠及柱头扫描电镜图。(A, B)长花柱型花粉赤道面; (C, D)长花柱型花粉极面; (E, F)短花柱型花粉赤道面; (G, H)短花柱型花粉极面; (I)长花柱型花冠喉部的毛; (J)短花柱型花冠喉部的毛; (K)长花柱型湿性柱头, 2裂; (L)短花柱型湿性柱头, 2裂。

Fig. 3 Morphology of pollen grains, corolla tubes and stigmas of Hedyotis pulcherrima under scanning electron microscopy. Equatorial (A, B) and polar view (C, D) of pollen grains from long-styled flowers; equatorial (E, F) and polar (G, H) view of pollen grains from short-styled flowers; pubescence at the throat of the corolla tube of a long-styled (I) and a short-styled (J) flower; bilobed stigma of the wet papillate type in a long-styled (K) and short-styled (L) flower.

图4 艳丽耳草长/短花柱型花粉组织化学染色。(A)长花柱型花粉KI-I2染色; (B)短花柱型花粉KI-I2染色; (C)长花柱型花粉苏丹IV染色; (D)短花柱型花粉苏丹IV染色。标尺= 50 μm。

Fig. 4 Histochemical staining of Hedyotis pulcherrima pollen grains. (A) KI-I2 staining of long-styled pollen grains; (B) KI-I2 staining of short- styled pollen grains; (C) Sudan IV staining of long-styled pollen grains; (D) Sudan IV staining of short-styled pollens. Bar =50 μm.

图5 艳丽耳草人工授粉处理后花粉管体内生长图。(A)长花柱型花自交处理24 h后, 红框内示花粉管顶端膨大; (B)短花柱型花自交处理24 h后; (C)长花柱型花型内异交处理24 h后; (D)短花柱型花型内异交处理24 h后, 红框内示花粉管顶端膨大; (E)短花柱型花型间异交处理24 h后; (F)长花柱型花型间异交处理24 h后; 标尺=1 mm。

Fig. 5 Pollen tube growth in vivo of Hedyotis pulcherrima following self-, intramorph and inter- morph pollination. (A) Long-styled morph stigma lobes and style 24 h after self-pollination; red ellipse showing accumulation of callose in the swollen tube; (B) Short-styled morph stigma lobes and style 24 h after self-pollination; (C) Long-styled morph stigma lobes and style 24 h after intramorph pollination; (D) Short-styled morph stigma lobes and style 24 h after intramorph pollination; red ellipse showing accumulation of callose in the swollen tube; (E) Short-styled morph stigma lobes and style 24 h after intermorph pollination; (F) Long-styled morph stigma lobes and style 24 h after intermorph pollination. Bar = 1 mm.

图6 艳丽耳草花粉管体内生长速度。(A)型间异交下花粉管生长速度; (B)自交及型内异交下花粉管生长速度。L: 长花柱型; S: 短花柱型。

Fig. 6 Pollen tube growth rate of Hedyotis pulcherrima in vivo. (A) In intermorph pollination; (B) In self- and intramorph pollination. L, Long-styled morph; S, Short-styled morph.

表3 艳丽耳草人工辅助授粉结实率

Table 3 Fruit sets of Hedyotis pulcherrima under artificial pollination

处理方式 Treatment	结实率 Fruit sets (%)		t-检验 t-test
处理方式 Treatment	长花柱型 Long-styled morph	短花柱型 Short-styled morph	t-检验 t-test
自交 Self-pollination	0 (n = 35)	0 (n = 52)	–
型内异交 Intramorph pollination	0 (n = 37)	0 (n = 30)	–
型间异交 Intermorph pollination	100 (n = 31)	100 (n = 29)	–
去雄套袋 Netting with emasculation	0 (n = 83)	0 (n = 63)	–
自然套袋 Netting without emasculation	0 (n = 53)	0 (n = 54)	–
自然 Open pollination	70.2±16.2 (n = 58)	68.2±21.3 (n = 93)	0.08

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艳丽耳草的二型花柱及异型自交不亲和系统

Distyly and heteromorphic self-incompatibility of Hedyotis pulcherrima (Rubiaceae)

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