红树植物胎生现象研究进展

doi:10.17521/cjpe.2016.0087

摘要/Abstract

摘要：

植物胎生是指有性繁殖产生的后代在母体上直接萌发的现象, 在红树植物中最为常见。红树植物生长在热带亚热带海岸潮间带, 耐受高盐、高温、淹水缺氧和海浪冲击等复杂环境。胎生被认为是红树植物对这种特殊生境的重要适应方式。该文从形态发育、生理生化、分子水平、生态适应4个层次讨论红树植物胎生现象对复杂生境的适应性, 并指出现有研究存在的不足, 对将来的研究方向进行了展望。与非胎生胚胎发育相比, 红树植物胎生是一个遗传的程序, 在进化过程中形成了一些特殊的结构。植物激素对胎生发育起关键的调控作用, 繁殖体发育过程中, 其盐离子的种类与浓度的动态变化则是对海岸潮间带生境的重要适应特征。这种胎生繁殖体依靠在母体上完善的一系列功能性特征能更有效地适应落地后的滩涂环境。然而, 红树植物胎生发育过程的分子机理及调控机制还有待研究。理解胎生这一特殊适应性现象的本质及其进化过程将为红树林保护繁育、适应气候变化提供理论依据。

关键词: 红树植物, 胎生现象, 生态适应性, 海岸潮间带

Abstract:

Vivipary in plants refers to a phenomenon that sexually reproduced offsprings germinate while still attached to the maternal bodies. This is mostly manifested in mangrove plants, which occur in tropical and subtropical intertidal zones and encounter harsh environmental conditions such as high salinity, high temperatures, waterlogging, hypoxia and tidal waves. Vivipary has long been recognized as one of the most important adaptive features under such a complex environment. Here we discuss four aspects of vivipary: morphological anatomy, physiology and biochemistry, molecular biology and ecological adaptation. We also discuss shortcomings in current studies and prospect of future directions. Differing from regular seed development, viviparous seeds in mangroves are evolved with many special structures, indicating a genetically based process. Hormones play an important role in regulating the process, whilst the dynamics of salt ion concentration during embryo and propagule development seems to be an adaptive feature. The ecological significance of vivipary is fully exhibited in the propagules that can effectively establish themselves on muddy tidal zones. Such a success heavily relies on sound functional features developed on the mother plants. However, the molecular mechanism and the regulation of viviparous seed development in mangroves remain elusive. Systematic studies of vivipary in mangroves not only help to understand the nature and evolutionary process of this distinct adaptive phenomenon, but also provide the foundation for mangrove forest restoration and protection in many parts of the world.

Key words: mangrove plants, vivipary, ecological adaption, coastal intertidal zone

周晓旋, 蔡玲玲, 傅梅萍, 洪礼伟, 沈英嘉, 李庆顺. 红树植物胎生现象研究进展. 植物生态学报, 2016, 40(12): 1328-1343. DOI: 10.17521/cjpe.2016.0087

Xiao-Xuan ZHOU, Ling-Ling CAI, Mei-Ping FU, Li-Wei HONG, Ying-Jia SHEN, Qingshun Quinn LI. Progress in the studies of vivipary in mangrove plants. Chinese Journal of Plant Ecology, 2016, 40(12): 1328-1343. DOI: 10.17521/cjpe.2016.0087

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2016.0087

https://www.plant-ecology.com/CN/Y2016/V40/I12/1328

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科名 Family	种名 Species	生境 Habitat	参考文献 Reference
天南星科 Araceae	Dieffenbachia longispatha	湿润林 Wet forest	Janzen, 1983
	隐棒花 Cryptocoryne sp. (1)	海岸 Coast	Pijl, 1972
	Aglaodorum griffithii	海岸 Coast	Pijl, 1972
棕榈科 Arecaceae	椰子 Cocos nucifera	海岸 Coast	Sankaran et al., 2012
棕榈科 Arecaceae	水椰 Nypa fruticans*	海岸 Coast	Tomlinson, 1986
菊科 Asteraceae	Abrotanella linearis	湿润林 Wet forest	Simpson, 1979
菊科 Asteraceae	Pachystegia insignis	湿润林 Wet forest	Simpson, 1979
海榄雌科 Avicenniaceae	Avicennia spp. (8)*	海岸 Coast	Marilyn & Graham, 1984
木棉科 Bombacaceae	Montezuma speciosissima	湿润林 Wet forest	Marrero, 1942
石竹科 Caryophyllaceae	Scheidea diffusa	山地森林 Montane	Wagner et al., 1999
牛栓藤科 Connaraceae	Connarus grandis	湿润林 Wet forest	Corner, 1976
山茱萸科 Cornaceae	Corokia macrocarpa	湿润林 Wet forest	Fountain & Outred, 1991
葫芦科 Cucurbitaceae	佛手瓜 Sechium edule	农田 Farmland	Singh & Mathur, 2004
丝粉藻科 Cymodoceaceae	Amphibolus spp. (2)	海岸 Coast	Ducker & Knox, 1976
	Thalassodendron spp. (2)	海岸 Coast	Tomlinson, 1986
大戟科 Euphorbiaceae	麻疯树 Jatropha curcas	农田 Farmland	Deore & Johnson, 2008
豆科 Fabaceae	Inga spp. (2)	沼泽林 Swamp forest	Leck et al., 2008
	Pithecellobium racemosum	湿润林 Wet forest	Leite & Rankin, 1981
玉蕊科 Lecythidaceae	玉蕊 Barringtonia racemosa	海岸 Coast	Guppy, 1912
百合科 Liliaceae	Crinum capense	河岸 Riverine	Guppy, 1912
	Hymenocallis spp. (2)	河岸 Riverine	Dahlgren et al., 1985
	Nerine sp. (1)	草地 Grassland	Dahlgren et al., 1985
	Ripogonum scandens	湿润林 Wet forest	Burrows, 1997
桑科 Moraceae	Morus latifolia	湿润林 Wet forest	Ellis et al., 1985
	Artocarpus incisa	湿润林 Wet forest	Guppy, 1912
紫金牛科 Myrsinaceae	Aegiceras spp. (2)*	海岸 Coast	Tomlinson, 1986
紫茉莉科 Nyctaginaceae	Pisonia longirostris	沼泽林 Swamp forest	Corner, 1976
Pellicierieaceae	Pelliciera rhizophora*	海岸 Coast	Tomlinson, 1986
白花丹科 Plumbaginaceae	Aegialitis spp. (2)	海岸 Coast	Tomlinson, 1986
蓼科 Polygonaceae	荞麦 Fagopyrum esculentum	农田 Farmland	Joshi & Paroda, 1991
禾本科 Poaceae	Dinochloa sp. (1)	湿润林 Wet forest	Dransfield & Bulletin, 1981
	Melocanna baccifera	湿润林 Wet forest	Ramanayake & Weerawardene, 2003
红树科 Rhizophoraceae	Bruguiera spp. (6)*	海岸 Coast	Tomlinson, 1986
	Ceriops spp. (3)*	海岸 Coast	Tomlinson, 1986
	Kandelia spp. (2)*	海岸 Coast	Sheue et al., 2003
	Rhizophora spp. (8)*	海岸 Coast	Tomlinson, 1986
茜草科 Rubiaceae	Coprosma robusta	湿润林 Wet forest	Burrows, 1995
	Ophiorrhiza tomentosa	湿润林 Wet forest	Tan & Rao, 1981
	Ophiorrhiza mungos	湿润林 Wet forest	Dintu et al., 2015
山榄科 Sapotaceae	Euphrasia disperma	湿润林 Wet forest	Simpson, 1977
仙人掌科 Cactaceae	Coryphantha vivipara	干旱 Drought	Cota-Sánchez, 2004
	Disocactus martianus	热带 Tropics	Cota-Sánchez, 2004
	Epiphyllum X Fern la Borde	栽培 Cultivation	Cota-Sánchez, 2004
	Epiphyllum phyllanthus	热带 Tropics	Conde, 1975
	Lepismium ianthothele	热带 Tropics	Cota-Sánchez, 2004
表1 (续) Table 1 (continued)
科名 Family	种名 Species	生境 Habitat	参考文献 Reference
仙人掌科 Cactaceae	Lepismium monacanthum	热带 Tropics	Cota-Sánchez, 2004
	Rhipsalis pilocarpa	热带 Tropics	Lombardi JA, 1993
	Rhipsalis baccifera	干旱 Drought	Cota-Sánchez, 2004
	Cleistocactus smaragdiflorus	干旱 Drought	Cota-Sánchez, 2004
	Harrisia martinii	干旱 Drought	Cota-Sánchez, 2004

科名 Family	种名 Species	生境 Habitat	参考文献 Reference
天南星科 Araceae	Dieffenbachia longispatha	湿润林 Wet forest	Janzen, 1983
	隐棒花 Cryptocoryne sp. (1)	海岸 Coast	Pijl, 1972
	Aglaodorum griffithii	海岸 Coast	Pijl, 1972
棕榈科 Arecaceae	椰子 Cocos nucifera	海岸 Coast	Sankaran et al., 2012
棕榈科 Arecaceae	水椰 Nypa fruticans*	海岸 Coast	Tomlinson, 1986
菊科 Asteraceae	Abrotanella linearis	湿润林 Wet forest	Simpson, 1979
菊科 Asteraceae	Pachystegia insignis	湿润林 Wet forest	Simpson, 1979
海榄雌科 Avicenniaceae	Avicennia spp. (8)*	海岸 Coast	Marilyn & Graham, 1984
木棉科 Bombacaceae	Montezuma speciosissima	湿润林 Wet forest	Marrero, 1942
石竹科 Caryophyllaceae	Scheidea diffusa	山地森林 Montane	Wagner et al., 1999
牛栓藤科 Connaraceae	Connarus grandis	湿润林 Wet forest	Corner, 1976
山茱萸科 Cornaceae	Corokia macrocarpa	湿润林 Wet forest	Fountain & Outred, 1991
葫芦科 Cucurbitaceae	佛手瓜 Sechium edule	农田 Farmland	Singh & Mathur, 2004
丝粉藻科 Cymodoceaceae	Amphibolus spp. (2)	海岸 Coast	Ducker & Knox, 1976
	Thalassodendron spp. (2)	海岸 Coast	Tomlinson, 1986
大戟科 Euphorbiaceae	麻疯树 Jatropha curcas	农田 Farmland	Deore & Johnson, 2008
豆科 Fabaceae	Inga spp. (2)	沼泽林 Swamp forest	Leck et al., 2008
	Pithecellobium racemosum	湿润林 Wet forest	Leite & Rankin, 1981
玉蕊科 Lecythidaceae	玉蕊 Barringtonia racemosa	海岸 Coast	Guppy, 1912
百合科 Liliaceae	Crinum capense	河岸 Riverine	Guppy, 1912
	Hymenocallis spp. (2)	河岸 Riverine	Dahlgren et al., 1985
	Nerine sp. (1)	草地 Grassland	Dahlgren et al., 1985
	Ripogonum scandens	湿润林 Wet forest	Burrows, 1997
桑科 Moraceae	Morus latifolia	湿润林 Wet forest	Ellis et al., 1985
	Artocarpus incisa	湿润林 Wet forest	Guppy, 1912
紫金牛科 Myrsinaceae	Aegiceras spp. (2)*	海岸 Coast	Tomlinson, 1986
紫茉莉科 Nyctaginaceae	Pisonia longirostris	沼泽林 Swamp forest	Corner, 1976
Pellicierieaceae	Pelliciera rhizophora*	海岸 Coast	Tomlinson, 1986
白花丹科 Plumbaginaceae	Aegialitis spp. (2)	海岸 Coast	Tomlinson, 1986
蓼科 Polygonaceae	荞麦 Fagopyrum esculentum	农田 Farmland	Joshi & Paroda, 1991
禾本科 Poaceae	Dinochloa sp. (1)	湿润林 Wet forest	Dransfield & Bulletin, 1981
	Melocanna baccifera	湿润林 Wet forest	Ramanayake & Weerawardene, 2003
红树科 Rhizophoraceae	Bruguiera spp. (6)*	海岸 Coast	Tomlinson, 1986
	Ceriops spp. (3)*	海岸 Coast	Tomlinson, 1986
	Kandelia spp. (2)*	海岸 Coast	Sheue et al., 2003
	Rhizophora spp. (8)*	海岸 Coast	Tomlinson, 1986
茜草科 Rubiaceae	Coprosma robusta	湿润林 Wet forest	Burrows, 1995
	Ophiorrhiza tomentosa	湿润林 Wet forest	Tan & Rao, 1981
	Ophiorrhiza mungos	湿润林 Wet forest	Dintu et al., 2015
山榄科 Sapotaceae	Euphrasia disperma	湿润林 Wet forest	Simpson, 1977
仙人掌科 Cactaceae	Coryphantha vivipara	干旱 Drought	Cota-Sánchez, 2004
	Disocactus martianus	热带 Tropics	Cota-Sánchez, 2004
	Epiphyllum X Fern la Borde	栽培 Cultivation	Cota-Sánchez, 2004
	Epiphyllum phyllanthus	热带 Tropics	Conde, 1975
	Lepismium ianthothele	热带 Tropics	Cota-Sánchez, 2004
表1 (续) Table 1 (continued)
科名 Family	种名 Species	生境 Habitat	参考文献 Reference
仙人掌科 Cactaceae	Lepismium monacanthum	热带 Tropics	Cota-Sánchez, 2004
	Rhipsalis pilocarpa	热带 Tropics	Lombardi JA, 1993
	Rhipsalis baccifera	干旱 Drought	Cota-Sánchez, 2004
	Cleistocactus smaragdiflorus	干旱 Drought	Cota-Sánchez, 2004
	Harrisia martinii	干旱 Drought	Cota-Sánchez, 2004

物种 Species	基因名称 Gene name	生物功能 Biological functions	脱落酸 Abscisic acid (ABA)	文献来源 Reference
玉米 Zea mays	VP1	编码与ABA信号相关的转录激活因子, 在胚胎与胚乳中特异表达, 能控制种子成熟中的发育过程 Encoding a transcription activator related to ABA signaling, expressed specifically in embryo and endosperm tissues, and controls multiple developmental responses associated with the maturation phase of seed formation	不敏感 Insensitive	McCarty et al., 1989, 1991
	VP2	编码或调控4-羟苯丙酮酸二加氧酶, 可催化4-羟苯丙酮酸生成高龙胆酸, 高龙胆酸可影响质体醌的合成, 进而影响淀粉体中胡萝卜素的去饱和过程 Encoding or regulating the expression of 4-hydroxyphenylpyruvate dioxygenase, an enzyme catalyzing the synthesis of homogentisic acid from 4-hydroxyphenylpyruvate. Homogentisic acid affects the biosynthesis of plastoquinone, then blocks the process of carotene desaturation in amyloplasts	合成缺陷 Synthesis deficit	Matthews et al., 2003
	VP5	编码茄红素脱氢酶, 进而影响ABA的积累和种子的发育 Encoding a phytoene desaturase which is essential for ABA accumulation and seed development	合成缺陷 Synthesis deficit	Hable et al., 1998
	VP7	参与ABA合成通路中番茄红素转化为胡萝卜素的环化反应 Playing a role in cyclization reaction of lycopene to carotene in the ABA synthesis pathway	合成缺陷 Synthesis deficit	Singh et al., 2003
	VP10	编码Cnx1直系同源蛋白, 该酶能催化钼蛋白辅助因子的合成 Encoding an orthologous protein of Cnx1 which is required for molybdenum cofactor biosynthesis	正常 Normal	Porch et al., 2006
	VP14	编码一种类囊体膜蛋白, 催化ABA合成的关键步骤 Encoding a thylakoid membrane protein that catalyzes the committed step in ABA synthesis	合成缺陷 Synthesis deficit	Bertoni, 2010
	VP15	编码植物钼喋呤合成酶的小亚基, 影响ABA的合成 Encoding a plant molybdopterin small subunit that affects ABA synthesis	合成缺陷 Synthesis deficit	Suzuki et al., 2006
水稻 Oryza sativa	ENL1	编码SNF2解旋酶家族蛋白, 在水稻合胞体胚乳的发育过程中起重要作用 Encoding a SNF2 helicase family protein that is essential for syncytial endosperm development	正常 Normal	Hara et al., 2015
	PHS1	编码茄红素脱氢酶, 参与类胡萝卜素合成途径 Encoding a phytoene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS2	编码ζ胡萝卜素去饱和酶, 参与类胡萝卜素合成途径 Encoding a ζ-carotene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS3	编码类胡萝卜素异构酶, 参与类胡萝卜素合成途径 Encoding a carotenoid isomerase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS4	编码番茄红素β-环化酶, 参与类胡萝卜素合成途径 Encoding a lycopene β-cyclase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
向日葵 Helianthus annuus	ND-1	编码ζ胡萝卜素去饱和酶, 参与类胡萝卜素合成途径 Encoding a ζ-carotene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Conti et al., 2004
拟南芥 Arabidopsis thaliana	ABI3	ABA信号识别的主要调节因子, 编码与玉米Vp-1基因同源的转录因子, 调控胚胎成熟与幼苗的早期发育过程 A main regulator of ABA signal recognition, encoding a transcription factor that is homologous to VP1 in maize, functions in regulation of embryo maturation and early development of plant	不敏感 Insensitive	Meinke et al., 1994
	LEC1	编码与胚胎成熟和细胞分化相关的转录激活因子, 正调控脂肪酸的合成 Encoding a transcription activator related to embryo maturation and cell differentiation, acts as a positive regulator to fatty acid synthesis	正常 Normal	Meinke et al., 1994

物种 Species	基因名称 Gene name	生物功能 Biological functions	脱落酸 Abscisic acid (ABA)	文献来源 Reference
玉米 Zea mays	VP1	编码与ABA信号相关的转录激活因子, 在胚胎与胚乳中特异表达, 能控制种子成熟中的发育过程 Encoding a transcription activator related to ABA signaling, expressed specifically in embryo and endosperm tissues, and controls multiple developmental responses associated with the maturation phase of seed formation	不敏感 Insensitive	McCarty et al., 1989, 1991
	VP2	编码或调控4-羟苯丙酮酸二加氧酶, 可催化4-羟苯丙酮酸生成高龙胆酸, 高龙胆酸可影响质体醌的合成, 进而影响淀粉体中胡萝卜素的去饱和过程 Encoding or regulating the expression of 4-hydroxyphenylpyruvate dioxygenase, an enzyme catalyzing the synthesis of homogentisic acid from 4-hydroxyphenylpyruvate. Homogentisic acid affects the biosynthesis of plastoquinone, then blocks the process of carotene desaturation in amyloplasts	合成缺陷 Synthesis deficit	Matthews et al., 2003
	VP5	编码茄红素脱氢酶, 进而影响ABA的积累和种子的发育 Encoding a phytoene desaturase which is essential for ABA accumulation and seed development	合成缺陷 Synthesis deficit	Hable et al., 1998
	VP7	参与ABA合成通路中番茄红素转化为胡萝卜素的环化反应 Playing a role in cyclization reaction of lycopene to carotene in the ABA synthesis pathway	合成缺陷 Synthesis deficit	Singh et al., 2003
	VP10	编码Cnx1直系同源蛋白, 该酶能催化钼蛋白辅助因子的合成 Encoding an orthologous protein of Cnx1 which is required for molybdenum cofactor biosynthesis	正常 Normal	Porch et al., 2006
	VP14	编码一种类囊体膜蛋白, 催化ABA合成的关键步骤 Encoding a thylakoid membrane protein that catalyzes the committed step in ABA synthesis	合成缺陷 Synthesis deficit	Bertoni, 2010
	VP15	编码植物钼喋呤合成酶的小亚基, 影响ABA的合成 Encoding a plant molybdopterin small subunit that affects ABA synthesis	合成缺陷 Synthesis deficit	Suzuki et al., 2006
水稻 Oryza sativa	ENL1	编码SNF2解旋酶家族蛋白, 在水稻合胞体胚乳的发育过程中起重要作用 Encoding a SNF2 helicase family protein that is essential for syncytial endosperm development	正常 Normal	Hara et al., 2015
	PHS1	编码茄红素脱氢酶, 参与类胡萝卜素合成途径 Encoding a phytoene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS2	编码ζ胡萝卜素去饱和酶, 参与类胡萝卜素合成途径 Encoding a ζ-carotene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS3	编码类胡萝卜素异构酶, 参与类胡萝卜素合成途径 Encoding a carotenoid isomerase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
	PHS4	编码番茄红素β-环化酶, 参与类胡萝卜素合成途径 Encoding a lycopene β-cyclase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Fang et al., 2008
向日葵 Helianthus annuus	ND-1	编码ζ胡萝卜素去饱和酶, 参与类胡萝卜素合成途径 Encoding a ζ-carotene desaturase which is involved in carotenoid synthesis pathway	合成缺陷 Synthesis deficit	Conti et al., 2004
拟南芥 Arabidopsis thaliana	ABI3	ABA信号识别的主要调节因子, 编码与玉米Vp-1基因同源的转录因子, 调控胚胎成熟与幼苗的早期发育过程 A main regulator of ABA signal recognition, encoding a transcription factor that is homologous to VP1 in maize, functions in regulation of embryo maturation and early development of plant	不敏感 Insensitive	Meinke et al., 1994
	LEC1	编码与胚胎成熟和细胞分化相关的转录激活因子, 正调控脂肪酸的合成 Encoding a transcription activator related to embryo maturation and cell differentiation, acts as a positive regulator to fatty acid synthesis	正常 Normal	Meinke et al., 1994

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