虫害诱导的植物挥发物:基本特性、生态学功能及释放机制

植物在遭受植食性昆虫攻击时,能通过释放挥发物调节植物、植食性昆虫及其天敌三者间的相互关系,并由此而防御植食性昆虫。主要就虫害诱导的植物挥发物的基本特性、生态学功能及其释放机制进行了系统性综述,并提出了今后的研究方向。     

虫害诱导的植物挥发物代谢调控机制研究进展

长期受自然界的非生物/生物侵害,植物逐步形成了复杂的防御机制,为防御植食性昆虫的为害,植物释放虫害诱导产生的挥发性化合物(herbivore-induced plant volatiles,HIPVs)。HIPVs是植物-植食性昆虫-天敌三级营养关系之间协同进化的结果。HIPVs的化学组分因植物、植食性昆虫种类的不同而有差异。生态系统中,HIPVs可在植物与节肢动物、植物与微生物、虫害植物与邻近的健康植物、或同一植株的受害和未受害部位间起作用,介导防御性反应。HIPVs作为寄主定位信号,在吸引捕食性、寄生性天敌过程中起着重要作用。HIPVs还可以作为植物间信息交流的工具,启动植株的防御反应而增强抗虫性。不论从生态学还是经济学角度来看,HIPVs对于农林生态系中害虫综合治理策略的完善具有重要意义。前期的研究在虫害诱导植物防御的化学生态学方面奠定了良好基础,目前更多的研究转向阐述虫害诱导植物抗性的分子机制。为了深入了解HIPVs的代谢调控机制,主要从以下几个方面进行了综述。因为植食性昆虫取食造成的植物损伤是与昆虫口腔分泌物共同作用的结果,所以首先阐述口腔分泌物在防御反应中的作用。挥发物诱导素volicitin和β-葡萄糖苷酶作为口腔分泌物的组分,是产生HIPVs的激发子,通过调节伤信号诱发HIPVs的释放。接着阐述了信号转导途径对HIPVs释放的调节作用,并讨论了不同信号途径之间的交互作用。就HIPVs的代谢过程而言,其过程受信号转导途径(包括茉莉酸、水杨酸、乙烯、过氧化氢信号途径)的调控,其中茉莉酸信号途径是诱发HIPVs释放的重要途径。基于前人的研究,综述了HIPVs的主要代谢过程及其过程中关键酶类的调控作用。文中的HIPVs主要包括萜烯类化合物、绿叶挥发物和莽草酸途径产生的芳香族化合物,如水杨酸甲酯和吲哚等。作为化学信号分子,这些化合物中的一部分还能激活邻近植物防御基因的表达。萜烯合酶是各种萜烯类化合物合成的关键酶类,脂氧合酶、过氧化氢裂解酶也是绿叶挥发物代谢途径中的研究热点,而苯丙氨酸裂解酶和水杨酸羧基甲基转移酶分别是合成水杨酸及其衍生物水杨酸甲酯的关键酶类。这些酶类的基因在转录水平上调控着HIPVs代谢途径。最后展望了HIPVs的研究前景。     

植物的诱导抗虫性

植物对植食性昆虫的抗性可包括两个方面,即植物的组成抗性（constitutiveresistance）和诱导抗性（inducedresistance）。组成抗性是指植物在遭受植食性昆虫进攻前就已存在的抗虫特性;而诱导抗性是指植物在遭受植食性昆虫进攻后所表现出来的一种抗虫特性[1,2]。根据作用世代的不同,诱导抗性又分为迅速的诱导抗性（rapidlyinducedre－sistance,RIR）和滞后的诱导抗性（delayedinducedresistance,DIR）。前者是指对当前世代的植食性昆虫的影响,而后者是指对后续的1～几个世代的植食性昆虫的影响[2]。研究植物的诱导抗虫性,不仅能在…     

虫害诱导的植物挥发物:植物与植食性昆虫及其天敌相互作用的进化产物

综述了植物、植食性昆虫及其天敌相互作用的进化过程。虫害诱导的植物挥发物的特征和功能是植物-植食性昆虫-天敌之间长期进化的结果。在植物、植食性昆虫与天敌相互作用的进化过程中,3个不同营养级,包括植物、植食性昆虫和天敌有着各自的调节和利用虫害诱导的植物挥发物的策略。但有一些问题,如通过实验研究得出的诱导防御在田间是否真正能起到保护作用等需进一步研究、阐明。     

虫害诱导植物挥发物(HIPVs)对植食性昆虫的行为调控

虫害诱导植物挥发物(herbivore induced plant volatiles,HIPVs)具有植物种类、品种、生育期和部位的特异性,也具有植食性昆虫种类、虫龄、为害程度、为害方式和其他一些环境因子的特异性。由于其释放量明显大于健康植株,因此更易被天敌、害虫以及邻近的植物等所利用,从而调节植物、植食性昆虫与天敌三者之间的相互作用关系,增强植物在自然界的生存竞争能力。本文对HIPVs在植食性昆虫寄主定位行为中的作用、HIPVs对植食性昆虫的种群调控功能及其应用现状2个方面加以综述,并在展望中对目前研究中存在的一些问题进行了探讨。     

昆虫对植物抗虫性的诱导

植物的抗虫性主要表现为受遗传因素控制的遗传抗性和受环境因素控制的生态抗性两个方面。植物的抗性主要由遗传因素决定,但其抗性表达和抗虫幅度又受到环境因子的影响。环境因子可改变植物的生理状态和理化性质,使植物不合适作为某种昆虫的寄主,从而提高了植物的抗性水平。植食性昆虫对植物抗虫性的诱导是环境影响抗性表达的一个重要方面,它是植物对取食者的一种重要的防御策略,同时也是植物与植食者协同进化的结果,进一步了解和充分发挥诱导抗性的作用,对抗虫有种工作和害虫的综合治理都有指导意义。三植食性昆虫对植物抗虫性的诱导…     

外用不同浓度茉莉酸甲酯诱导的茶树挥发物的种类和时序变化

【目的】明确MeJA对茶树挥发物的诱导作用。【方法】采用顶空活体取样法对不同浓度MeJA处理后的茶苗挥发物进行抽提,并利用GC-MS对挥发物进行鉴定。【结果】不同剂量MeJA显著地影响茶树挥发物的种类组成和释放量,50μL MeJA处理可显著诱导茶树释放香叶烯、萜品油烯、罗勒烯等10种单萜类化合物,法呢烯、橙花叔醇和红没药烯等7种倍半萜类化合物,苯甲醇、苯乙腈和吲哚等5种氨基酸衍生物,以及3种未知化合物;而100μL MeJA处理仅能诱导茶树释放7种化合物。不同挥发物对MeJA处理的响应时间不同,但其释放量都具有昼高夜低的趋势。并且,释放量的大小明显受到光照强度的影响。【结论】外用MeJA喷雾处理可诱导茶树挥发物的产生和释放。     

虫害诱导蔬菜作物挥发物的研究进展

蔬菜作物释放的虫害诱导植物挥发物(Herbivore-induced plant volatiles,HIPVs)是蔬菜作物受害虫胁迫后产生的一类启动防御反应的化合物,可以作为蔬菜作物的对外重要信息交流媒介和防御措施实施前体.蔬菜作物通过释放HIPVs来增强自身防御反应、调节昆虫行为和向邻近作物发出"预警信号",从而直...     

虫害诱导的水稻挥发物对褐飞虱的驱避作用

水稻叶片在遭受害虫的危害时,释放一些特异性的挥发性物质。这些挥发性信息物质在调节植物、植食性昆虫及其天敌的相互关系中有重要作用。利用“Y”型嗅觉仪研究了褐飞虱对虫害诱导的11种水稻挥发物（浓度为2 Μl 化合物溶解在100 Μl丙酮溶液中）的行为反应。结果表明,(E)-2-己烯醛、(E)-2-己烯-1-醇、-2-庚醇和水杨酸甲酯等4种化合物对褐飞虱成虫有显著的驱避作用,而(Z)-3-己烯-1-醇、-2-庚酮、柠檬烯、罗勒烯、芳樟醇、-β-子丁香烯和橙花叔醇等7种物质对褐飞虱成虫的选择行为无显著影响。浓度梯度的实验表明,浓度(不同体积的待测化合物用100 Μl丙酮稀释)较低的(E)-2-己烯醛（0.5 Μl、1 Μl）和芳樟醇（05 Μl、1 Μl、5 Μl）对褐飞虱成虫的选择行为无显著影响,而较高浓度的(E)-2-己烯醛（5 Μl、10 Μl）和芳樟醇（10 Μl）对褐飞虱成虫均有明显的驱避作用。     

虫害诱导植物挥发物（HIPVs）：从诱导到生态功能

植物被植食性昆虫危害后,其地上和地下部分均能释放虫害诱导植物挥发物(HIPVs).HIPVs不仅能招引植食性昆虫的天敌,而且还可对植食性昆虫和临近植物产生影响,从而调节植物、植食性昆虫、天敌三者之间的相互关系.根据最近的研究结果,主要就HIPVs的诱导物类型、虫害后植物体内的早期信号和信号传导、HIPVs的合成、释放、组成,以及其生态功能等进行了系统性的综述,并提出了今后该领域的研究方向.     

Isolation of a psychrotolerant Debaryomyces hansenii strain from fermented tea plant (Camellia sinensis) leaves

Abstract

A psychrotolerant, halotolerant and alkalophilic yeast was isolated from fermented leaves of Camellia sinensis Kuntze, the tea plant. The yeast strain, named Tea-Y1, was both phenotypically and genotypically identified as belonging to the species Debaryomyces hansenii. This assignment was confirmed by scanning and transmission electron microscopy. The analysis of growth curves demonstrated the ability this yeast strain to grow in a temperature range between 4°C and 28°C, with an optimum of 23°C. The ecology of this yeast in the C. sinensis phyllosphere, as well as its possible role in tea fermentation and storage, with particular reference to iced tea, are discussed.     

Potentiality of actinobacteria to combat against biotic and abiotic stresses in tea [Camellia sinensis (L) O. Kuntze]

Tea (Camellia sinensis (L) O. Kuntze) is a long-duration monoculture crop prone to several biotic (fungal diseases and insect pest) and abiotic (nutrient deficiency, drought and salinity) stress that eventually result in extensive annual crop loss. The specific climatic conditions and the perennial nature of the tea crop favour growth limiting abiotic factors, numerous plant pathogenic fungi (PPF) and insect pests. The review focuses on the susceptibility of tea crops to PPF/pests, drought, salinity and nutrient constraints and the potential role of beneficial actinobacteria in promoting tea crop health. The review also focuses on some of the major PPF associated with tea, such as Exobasidium vexans, Pestalotiopsis theae, Colletotrichum acutatum, and pests (Helopeltis theivora). The phylum actinobacteria own a remarkable place in agriculture due to the biosynthesis of bioactive metabolites that assist plant growth by direct nutrient assimilation, phytohormone production, and by indirect aid in plant defence against PPF and pests. The chemical diversity and bioactive significance of actinobacterial metabolites (antibiotics, siderophore, volatile organic compounds, phytohormones) are valuable in the agro-economy. This review explores the recent history of investigations in the role of actinobacteria and its secondary metabolites as a biocontrol agent and proposes a commercial application in tea cultivation.     

Theanine transporters identified in tea plants (Camellia sinensis L.)

Theanine, a unique non‐proteinogenic amino acid, is an important component of tea, as it confers the umami taste and relaxation effect of tea as a beverage. Theanine is primarily synthesized in tea roots and is subsequently transported to young shoots, which are harvested for tea production. Currently, the mechanism for theanine transport in the tea plant remains unknown. Here, by screening a yeast mutant library, followed by functional analyses, we identified the glutamine permease, GNP1 as a specific transporter for theanine in yeast. Although there is no GNP1 homolog in the tea plant, we assessed the theanine transport ability of nine tea plant amino acid permease (AAP) family members, with six exhibiting transport activity. We further determined that CsAAP1, CsAAP2, CsAAP4, CsAAP5, CsAAP6, and CsAAP8 exhibited moderate theanine affinities and transport was H⁺‐dependent. The tissue‐specific expression of these six CsAAPs in leaves, vascular tissues, and the root suggested their broad roles in theanine loading and unloading from the vascular system, and in targeting to sink tissues. Furthermore, expression of these CsAAPs was shown to be seasonally regulated, coincident with theanine transport within the tea plant. Finally, CsAAP1 expression in the root was highly correlated with root‐to‐bud transport of theanine, in seven tea plant cultivars. Taken together, these findings support the hypothesis that members of the CsAAP family transport theanine and participate in its root‐to‐shoot delivery in the tea plant.     

Isolation and characterization of highly polymorphic microsatellites in tea (Camellia sinensis)

Relatively little is known about the diversity and origins of tea. The highest value tea products are sold on the basis of their region of origin but there are currently no methods available to verify the claims made on packages. We have developed 15 microsatellite loci for tea. These have been evaluated for polymorphism in a set of tea clones to determine their usefulness for authentication purposes. The majority of the microsatellites developed proved to be highly polymorphic both between and within different geographical origins and offer the potential to investigate the population genetics and genetic origins of tea.     

Micropropagation of tea (Camellia sinensis (L.) O. Kuntze) using Thidiazuron

The effect of thidiazuron (TDZ) on the micropropagation of Camellia sinensis (China hybrid) was compared with that of benzylaminopurine (BAP) using nodal segments from in vitro raised seedlings. Extremely low concentrations of TDZ (1pM–100nM) alone were effective in inducing shoot bud proliferation and maintaining high rates of shoot multiplication on hormone-free media. On the other hand, higher concentrations of BAP (1–10M) and its continued presence were required to initiate and sustain shoot proliferation. While wider ranges of BAP combined favourably with auxins like NAA or IBA, only specific combinations of TDZ and NAA were effective for shoot proliferation. TDZ treated explants yielded healthy shoots, with sturdy leaves, even during the initial stages of growth, whereas, the effect of BAP was cumulative over subcultures in attaining a high proliferative rate.     

茶(Camellia sinensis L.)对铜的吸收与累积

对广东省5个大型茶场中不同土壤背景值的8个茶园进行了土壤及茶(CamelliasinensisL.)树各部位的铜含量分析调查。地处我国南方的广东省茶园,土壤中铜含量在2.39～53.05mg kg之间。有效态铜含量相对较低,仅在0.45～3.17mg kg之间。茶树中各部位铜含量大小依次为枝条>幼叶>根>成熟叶。茶树枝条和幼叶铜含量较高,平均分别达14.5和14.3mg kg,远高于成熟叶铜含量水平(9.7mg kg)。而同时期相应茶场的茶叶制成品铜含量则较幼叶(一芽两叶)铜的含量高。土壤中的总钾、有效磷和总铜等都对茶树各部位铜的累积产生较大影响,但只有土壤有效磷含量对铜在茶树体内向枝条部分的转移和累积影响达显著程度,枝条铜含量与土壤有效磷之间的相关系数r=-0.74,呈显著负相关(P<0.05)。     

依赖于叶黄素循环的热耗散对茶树叶片防御强光破坏的相关试验研究

经叶黄素循环抑制剂——二硫苏糖醇(DIT)处理的茶树叶片,以850μmol．m^-2．s^-1的PFD照射120min后,福鼎大白茶的叶黄素循环组分中的环氧玉米黄素(A)和玉米黄素(Z)含量之和降低了76．5％,结果导致非光化学猝灭(NPQ)、光系统Ⅱ(PSⅡ)的光化学效率(Fv／Fm)、光化学猝灭系数(qP)、PSⅡ实际光化学量子效率(ψPSⅡR)和光合电子传递速率(ETR)明显下降,而F0显著上升,暗恢复后Fv／Fm恢复程度小于未经DIT处理的叶片。自然光强下,NPQ与与叶黄素循环的脱环氧化程度(A Z)/(V A Z)比值呈明显的正线性关系(R=0．9488^***)。这些结果充分证明依赖与叶黄素循环的热耗散是茶树叶片光合器官防御强光破坏的主要途径。     

Effect of nitrogen form and root-zone pH on growth and nitrogen uptake of tea (Camellia sinensis) plants

BACKGROUND AND AIMS: Tea (Camellia sinensis) is considered to be acid tolerant and prefers ammonium nutrition, but the interaction between root zone acidity and N form is not properly understood. The present study was performed to characterize their interaction with respect to growth and mineral nutrition. METHODS: Tea plants were hydroponically cultured with NH4+, NO3- and NH(4+) + NO3-, at pH 4.0, 5.0 and 6.0, which were maintained by pH stat systems. KEY RESULTS: Plants supplied with NO3- showed yellowish leaves resembling nitrogen deficiency and grew much slower than those receiving NH4+ or NH(4+) + NO3- irrespective of root-zone pH. Absorption of NH4+ was 2- to 3.4-fold faster than NO3- when supplied separately, and 6- to 16-fold faster when supplied simultaneously. Nitrate-grown plants had significantly reduced glutamine synthetase activity, and lower concentrations of total N, free amino acids and glucose in the roots, but higher concentrations of cations and carboxylates (mainly oxalate) than those grown with NH4+ or NH(4+) + NO3-. Biomass production was largest at pH 5.0 regardless of N form, and was drastically reduced by a combination of high root-zone pH and NO3-. Low root-zone pH reduced root growth only in NO(3-)-fed plants. Absorption of N followed a similar pattern as root-zone pH changed, showing highest uptake rates at pH 5.0. The concentrations of total N, free amino acids, sugars and the activity of GS were generally not influenced by pH, whereas the concentrations of cations and carboxylates were generally increased with increasing root-zone pH. CONCLUSIONS: Tea plants are well-adapted to NH(4+)-rich environments by exhibiting a high capacity for NH4+ assimilation in their roots, reflected in strongly increased key enzyme activities and improved carbohydrate status. The poor plant growth with NO3- was largely associated with inefficient absorption of this N source. Decreased growth caused by inappropriate external pH corresponded well with the declining absorption of nitrogen.