根寄生植物种子萌发刺激物研究进展

根寄生植物是被子植物中一类寄生在寄主根部, 以摄取寄主水分和营养物质为生的特殊植物类群, 其种子萌发需要寄主萌发刺激物的诱导。该文主要阐述根寄生植物种子萌发的特异性, 以及目前已发现的刺激种子萌发的信号物质及其调节机制和生物合成途径, 并就萌发刺激物的识别机制及其在根寄生植物或丛枝菌根真菌与寄主建立寄生关系过程中所起的作用进行综述, 提出根寄生植物种子萌发研究中存在的问题, 并对其研究前景进行了展望。     

寄生植物对寄主植物的化学识别

植物间寄生关系的研究近年来受到了广泛的重视。大量的研究表明,寄主释放的次生物质对植物间寄生关系的建立和维持起了重要的调节作用。寄主植物的次生物质对寄生植物的化学防御和昆虫授粉等生态功能起重要的作用,寄主植物次生物质对寄生植物生理与生态的调节作用是受寄生植物基因调节的。更为重要的是寄主植物释放的次生物质成为寄生植物的种子萌发和吸器发生的异源识别物质。能够刺激寄生植物种子萌发的次生物质主要是倍半萜和氢醌类物质,而诱导吸器发生的物质则是酚酸、醌和黄酮类化合物,诱导吸器发生的核心结构是对苯醌。这些异源识别物质大多是寄主植物释放的化感抑制物质,显示寄生植物在化学防御方面要比寄主植物高级。异源识别化合物的活性与其氧化潜力显著相关。由于寄生植物中存在一抑制异源识别物质诱导吸器发生的调节过程,因此吸器的产生与寄生植物根部接触异源识别物质的浓度与时间呈正相关关系,这一调节过程对寄生植物准确识别寄主并寄生其上是十分重要的。对寄生植物和寄主植物间的化学识别关系的揭示有助于人们防治有害寄生植物和开发利用有价值的寄生植物资源。     

寄生植物概述

对寄生植物的概念、寄生类型、寄生生物学特性和经济价值与危害进行了总结与概括,并对世界和我国广泛分布的重点寄生植物进行了着重介绍。对国内外目前关于寄生植物种子萌发刺激物质、吸器形成诱导物质以及寄生植物与寄主植物间形成寄生关系的作用机理进行了探讨。     

根寄生植物吸器发育及其调控的研究进展

很多根寄生植物是重要的杂草,严重影响农业生产。吸器是寄生植物与寄主连接的特异性器官,是寄生植物与寄主间信号和物质交流的"桥梁",而吸器诱导因子在诱导吸器发育过程中扮演着重要角色。该文针对寄生植物吸器的发育过程,吸器诱导因子的种类、信号转导和基因调控,寄生植物如何避免寄主的防御这三方面的进展进行了综述,以期为进一步揭示寄生植物识别寄主的信号物质的分子机制提供参考。     

寄生植物和寄主间的分子交流研究进展

寄生植物和寄主植物间存在广泛的物质、能量和信息交流事件。寄生植物通过吸器从寄主吸收所需的多种矿物质元素,以及氨基酸、糖类、中间代谢产物等营养物质。此外,寄生植物与其寄主还进行着诸如蛋白质、RNA等大分子物质,及病毒、类病毒、植原体等微生物的交流。研究表明,mRNA在寄生植物和寄主间进行着大量的转移,其结构和功能的特异性可能对转移起着决定性的作用。对寄生植物来说,部分mRNA可能在寄生过程和调节自身生长发育过程中发挥着重要作用。本文对近年来寄生植物和寄主间的分子交流事件的相关报道进行综述,其中着重讨论mRNA转移的研究进展,以期对转移mRNA的结构和转移机理有进一步了解,同时对mRNA转移在寄生植物-寄主系统中的角色进行定位,为研究不同寄生系统中的mRNA转移奠定基础。     

是谁唤醒了寄生植物种子

众所周知，植物种子发芽的环境条件是水分、温度和空气。然而寄生植物的种子，却因其发芽后特殊的生活方式，而有着与众不同的萌发习性。有人做过试验，试图用种种常规方法和特殊手段促使寄生植物肉众蓉的种子萌发，但都没有成功。然而当这些种子被埋在寄主植物梭梭极的近旁时，很快就萌发了。这种现象在列当、独脚金、蛇板等众多根部寄生植物中，也普遍存在。列当、独脚金都是声名狼藉的农田恶性杂草，前者被称为“吸血鬼”，后者则有“女巫草”的绰号。它们都有十分巨大的生殖能力，每一株可产生十几万粒十分微小的种子。这些种子散布在农…     

MicroRNA在种子发育、休眠与萌发过程中的作用

植物microRNA(miRNA)是一类小分子非编码RNA,对植物的生长发育发挥着重要调控作用。种子发育、休眠与萌发是植物生命进程中的重要阶段。在这一阶段内,种子受各种环境因子及内源激素调控,并且不同植物种子具有不同发育及休眠特性。随着人们对种子发育、休眠及萌发机理的探究,越来越多miRNA被鉴定,它们能够基于植物激素信号传导、抗氧化作用、关键转录因子调控等途径参与种子形态建成、物质代谢及各种胁迫响应。本文主要综述了近年来植物miRNA的形成及调控机理,以及在种子发育、休眠及萌发过程中发挥的调控作用,旨在为今后的研究方向提供参考。     

不同列当抗性的向日葵品种根系分泌物对向日葵列当种子萌发的影响

以对向日葵列当抗性不同的向日葵幼苗为材料,利用水培法收集其根系分泌物,经二氯甲烷萃取后结合硅胶柱色谱分离法,研究不同列当抗性的向日葵品种根系分泌物对向日葵列当种子萌发的影响。结果表明:(1)在一定浓度范围内,向日葵根系分泌物对向日葵列当种子萌发具有低浓度促进、高浓度抑制的作用;二氯甲烷萃取的向日葵根系分泌物浓度为0.1mg/mL时,其向日葵列当萌发率高于其他浓度的作用。(2)‘星火大白边’和‘白葵杂9号’根系分泌物提取物作用下向日葵列当萌发率最高分别为46.92%和43.88%,显著高于免疫品种‘MGS’根系分泌物提取物作用下列当的萌发率(27.81%)。(3)根系分泌物提取物正己烷-乙酸乙酯1∶1洗脱组分的萌发刺激活性显著高于其他组分,‘星火大白边’、‘白葵杂9号’和‘MGS’的该组分作用下向日葵列当最高萌发率分别为49.90%、45.66%和30.00%。(4)‘MGS’的根系分泌物提取物与GR24或脱氢木香内酯共同作用时种子萌发率显著低于GR24或脱氢木香内酯的单独作用,说明‘MGS’根系分泌物中可能含有抑制GR24或脱氢木香内酯刺激向日葵列当种子萌发的物质,使其对向日葵列当表现出免疫特性。     

寄生植物对寄主的选择和影响

寄生植物对寄主的选择性与寄主营养物质的含量、合成的次生物质和硬度有关.多寄主存在的条件下,有利于寄生植物的生长发育.通过改变寄主光合产物的流向和影响寄主叶片的气孔调节,寄生植物对寄主生理和形态产生显著的影响.寄生植物的存在使植物群落的生物量、组成多样性和动态发生改变.大量研究显示,寄生植物对寄主的选择和影响与植食性动物对取食植物的选择和影响有很多相似之处.气候变化对寄生植物与寄主关系影响的研究还刚刚起步,寄生植物对寄主选择和影响的研究对有害寄生植物的防除和有益寄生植物的利用有重要的价值,应该重视和加强.     

植物间挥发物信号的研究进展

植物VOCs信号是植物间进行信息交流的"语言",可由多种生物和非生物因素诱导产生。非寄生植物释放的VOCs信号可影响其所在群落中其它植物的种子萌发与幼苗生长;而寄主植物释放的VOCs信号却是诱导寄生植物种子萌发和幼苗生长的信号物质。VOCs作为植物间的伤害信息可以诱导临近的同种或异种植物做好防御准备,从而通过主动或间接防御以减少外界的伤害。植物间通过VOCs信号进行信息交流,从而实现其繁衍与防御。该文通过对VOCs信号的种类、诱导产生因素、传递及作用进行综述,以期对VOCs信号的研究有所帮助。     

IAA production during germination of Orobanche spp. seeds

Broomrapes (Orobanche spp.) are parasitic plants, whose growth and development fully depend on the nutritional connection established between the parasite and the roots of the respective host plant. Phytohormones are known to play a role in establishing the specific Orobanche-host plant interaction. The first step in the interaction is seed germination triggered by a germination stimulant secreted by the host-plant roots. We quantified indole-3-acetic acid (IAA) and abscisic acid (ABA) during the seed germination of tobacco broomrape (Orobanche ramosa) and sunflower broomrape (O. cumana). IAA was mainly released from Orobanche seeds in host-parasite interactions as compared to non-host-parasite interactions. Moreover, germinating seeds of O. ramosa released IAA as early as 24 h after the seeds were exposed to the germination stimulant, even before development of the germ tube. ABA levels remained unchanged during the germination of the parasites' seeds. The results presented here show that IAA production is probably part of a mechanism triggering germination upon the induction by the host factor, thus resulting in seed germination.     

Thermoinhibition uncovers a role for strigolactones in Arabidopsis seed germination

Strigolactones are host factors that stimulate seed germination of parasitic plant species such as Striga and Orobanche. This hormone is also important in shoot branching architecture and photomorphogenic development. Strigolactone biosynthetic and signaling mutants in model systems, unlike parasitic plants, only show seed germination phenotypes under limited growth condition. To understand the roles of strigolactones in seed germination, it is necessary to develop a tractable experimental system using model plants such as Arabidopsis. Here, we report that thermoinhibition, which involves exposing seeds to high temperatures, uncovers a clear role for strigolactones in promoting Arabidopsis seed germination. Both strigolactone biosynthetic and signaling mutants showed increased sensitivity to seed thermoinhibition. The synthetic strigolactone GR24 rescued germination of thermoinbibited biosynthetic mutant seeds but not a signaling mutant. Hormone analysis revealed that strigolactones alleviate thermoinhibition by modulating levels of the two plant hormones, GA and ABA. We also showed that GR24 was able to counteract secondary dormancy in Arabidopsis ecotype Columbia (Col) and Cape Verde island (Cvi). Systematic hormone analysis of germinating Striga helmonthica seeds suggested a common mechanism between the parasitic and non-parasitic seeds with respect to how hormones regulate germination. Thus, our simple assay system using Arabidopsis thermoinhibition allows comparisons to determine similarities and differences between parasitic plants and model experimental systems for the use of strigolactones.     

Secondary metabolite signalling in host-parasitic plant interactions

The parasitic weeds Orobanche and Striga spp. are a serious threat to agriculture in large parts of the world. The lifecycle of the parasitic weeds is closely regulated by the presence of their hosts, and secondary metabolites that are produced by host plants play an important role in this interaction. Model plants, such as Arabidopsis and maize mutant collections, have been increasingly used to study these chemical signals, especially those host-produced stimulants that induce the germination of parasite seeds.     

An improved axenic system for studying pre-infection development of the parasitic plant Orobanche ramosa

BACKGROUND AND AIMS: Broomrapes (Orobanche spp.) are holoparasitic weeds that cause devastating losses in many economically important crops. The molecular mechanisms that control early stages of host infection in Orobanche are poorly understood, partly due to the lack of experimentally tractable in vitro systems that allow the efficient application of molecular tools. Here an improved axenic system for the analysis of pre-infection stages in O. ramosa in the absence of the host plant is described. METHODS: An optimized protocol for seed disinfection, based on formaldehyde, was developed. Orobanche ramosa seeds were conditioned in Petri dishes with filter paper, stimulated by addition of the synthetic strigol analogue GR24, and the percentage of germination as well as attachment-organ formation was determined. KEY RESULTS: Treatment of O. ramosa seeds with tobacco-root exudate or with GR24 resulted in highly reproducible germination rates around 70 %. A conditioning period of 8 d was both necessary and sufficient to allow optimal germination in response to GR24. Conditioned seeds that were dehydrated for several months remained fully responsive to GR24 without the need of a new conditioning period. Treatments as short as 5 min with GR24 were sufficient to fully and irreversibly induce the seed germination response. Approximately half of the germinated seeds initiated attachment-organ development. Similar rates of attachment organ induction were also detected in the rare cases of seeds that had germinated spontaneously on water. CONCLUSIONS: The results suggest that the conditioning period produces persistent changes in the seeds required for responsiveness to external stimulants. The rapid action of GR24 suggests that it may act via a receptor-mediated signalling mechanism. While germination in O. ramosa is induced by exogenous stimuli, attachment organ differentiation appears to be triggered by unknown endogenous signals. The new in vitro culture system will have useful applications for the molecular analysis of early stages of parasitic development in Orobanche.     

Induction of seed germination in <Emphasis Type="Italic">Orobanche</Emphasis> spp. by extracts of traditional Chinese medicinal herbs

The co-evolution of Orobanche spp. and their hosts within the same environment has resulted in a high degree of adaptation and effective parasitism whereby the host releases parasite germination stimulants, which are likely to be unstable in the soil. Our objective was to investigate whether extracts from non-host plants, specifically, Chinese medicinal plants, could stimulate germination of Orobanche spp. Samples of 606 Chinese medicinal herb species were extracted with deionized water and methanol. The extracts were used to induce germination of three Orobanche species; Orobanche minor, Orobanche cumana, and Orobanche aegyptiaca. O. minor exhibited a wide range of germination responses to the various herbal extracts. O. cumana and O. aegyptiaca exhibited an intermediate germination response to the herbal extracts. O. minor, which has a narrow host spectrum, showed higher germination rates in response to different herbal extracts compared with those of O. cumana and O. aegyptiaca, which have a broader host spectrum. Methanolic extracts of many Chinese herbal species effectively stimulated seed germination among the Orobanche spp., even though they were not the typical hosts. The effective herbs represent interesting examples of potential trap crops. Different countries can also screen extracts from indigenous herbaceous plants for their ability to induce germination of Orobanche spp. seeds. The use of such species as trap plants could diminish the global soil seed bank of Orobanche.     

The strigolactone germination stimulants of the plant-parasitic Striga and Orobanche spp. are derived from the carotenoid pathway

The seeds of parasitic plants of the genera Striga and Orobanche will only germinate after induction by a chemical signal exuded from the roots of their host. Up to now, several of these germination stimulants have been isolated and identified in the root exudates of a series of host plants of both Orobanche and Striga spp. In most cases, the compounds were shown to be isoprenoid and belong to one chemical class, collectively called the strigolactones, and suggested by many authors to be sesquiterpene lactones. However, this classification was never proven; hence, the biosynthetic pathways of the germination stimulants are unknown. We have used carotenoid mutants of maize (Zea mays) and inhibitors of isoprenoid pathways on maize, cowpea (Vigna unguiculata), and sorghum (Sorghum bicolor) and assessed the effects on the root exudate-induced germination of Striga hermonthica and Orobanche crenata. Here, we show that for these three host and two parasitic plant species, the strigolactone germination stimulants are derived from the carotenoid pathway. Furthermore, we hypothesize how the germination stimulants are formed. We also discuss this finding as an explanation for some phenomena that have been observed for the host-parasitic plant interaction, such as the effect of mycorrhiza on S. hermonthica infestation.     

Adaptation of the parasitic plant lifecycle: germination is controlled by essential host signaling molecules

Parasitic plants are plants that connect with a haustorium to the vasculature of another, host, plant from which they absorb water, assimilates, and nutrients. Because of this parasitic lifestyle, parasitic plants need to coordinate their lifecycle with that of their host. Parasitic plants have evolved a number of host detection/host response mechanisms of which the germination in response to chemical host signals in one of the major families of parasitic plants, the Orobanchaceae, is a striking example. In this update review, we discuss these germination stimulants. We review the different compound classes that function as germination stimulants, how they are produced, and in which host plants. We discuss why they are reliable signals, how parasitic plants have evolved mechanisms that detect and respond to them, and whether they play a role in host specificity. The advances in the knowledge underlying this signaling relationship between host and parasitic plant have greatly improved our understanding of the evolution of plant parasitism and are facilitating the development of more effective control measures in cases where these parasitic plants have developed into weeds.

Root parasitic plants grow on the roots of other plants and germinate only in the presence of that host, on which they completely depend, through the perception of host presence signaling molecules called germination stimulants.

Outstanding questions

• Have we overlooked the role of germination stimulants in facultative parasites?
• What is the biological relevance of the observation that many plant species produce and secrete a range of different strigolactones?
• Have parasitic plants evolved mechanisms to compensate for low phosphorus availability, a condition that stimulates their germination?
• What is the contribution of the HTL strigolactone receptors to host specificity in parasitic plants or does downstream signaling play a role?
• What other, nonstrigolactone, germination stimulants can parasitic plants respond to and does this require adaptation in the HTL receptors?
• What is the role of germination and underlying mechanism in the rapid adaptation of (orobanchaceous) parasitic plants to a new host?

     

Phosphorus deficiency in red clover promotes exudation of orobanchol,the signal for mycorrhizal symbionts and germination stimulant for root parasites

Plant derived sesquiterpene strigolactones, which have previously been characterized as germination stimulants for root parasitic plants, have recently been identified as the branching factors which induce hyphal branching morphogenesis, a critical step in host recognition by arbuscular mycorrhizal (AM) fungi. We show here that, in red clover plants (Trifolium pratense L.), which is known as a host for both AM fungi and the root holoparasitic plant Orobanche minor Sm., reduced supply of phosphorus (P) but not of other elements examined (N, K, Mg, Ca) in the culture medium significantly promotes the release of a strigolactone, orobanchol, by the roots of this plant. In red clover plants, the level of orobanchol exudation appeared to be regulated by P availability and was in good agreement with germination stimulation activity of the root exudates. This implies that under P deficiency, plant roots attract not only symbiotic fungi but also root parasitic plants through the release of strigolactones. This is the first report demonstrating that nutrient availability influences both symbiotic and parasitic interactions in the rhizosphere.     

Signaling Organogenesis in Parasitic Angiosperms: Xenognosin Generation, Perception, and Response

Abstract Parasitic strategies within the angiosperms generally succeed by tightly coupling developmental transitions with host recognition signals in a process referred to as xenognosis. Within the Scrophulariaceae, Striga asiatica is among the most studied and best understood parasitic member with respect to the processes of host recognition. Specific xenognosins regulate seed germination, the development of the host attachment organ, the haustorium, and several later stages of host-parasite integration. Here we discuss the signals regulating the development of the haustorium, the critical vegetative/parasitic transition in the life cycle of this obligate parasite. We provide evidence for the localized production of H₂O₂ at the Striga root tip and suggest how this oxidant is used to exploit host peroxidases and cell wall pectins to generate a simple benzoquinone signal. This benzoquinone xenognosin proves to be both necessary and sufficient for haustorial induction in cultured seedlings. Furthermore, evidence is provided that benzoquinone binding to a redox active site completes a ``redox circuit' to mediate signal perception. This redox reaction regulates the time-dependent expression of specific marker genes critical for the development of the mature host attachment organ. These studies extend the emerging series of events necessary for the molecular regulation of organogenesis within the parasitic plants and suggest novel signaling features and molecular mechanisms that may be common across higher plants.