虫害诱导植物间接防御反应的激发与信号转导途径

植物通过产生和释放挥发性物质增加植食性昆虫的天敌对其寄主或猎物的定位,减少植食性昆虫对植物的取食,从而达到间接防御的目的。植物对植食性昆虫所做出间接防御反应激发因子和信号转导途径的研究,对应用虫害诱导植物挥发物引诱害虫天敌,并进一步从植物、植食性昆虫及其天敌间三级营养关系,研究动植物协同进化机理和病虫害防治具有深远意义。本文根据国内外最新研究进展,对虫害诱导植物间接防御反应的激发因子,昆虫取食信号的转导途径及对植物间接防御相关基因的激活等方面进行了系统地综述。     

植物对植食性哺乳动物的化学防卫

综述植物次生合物防卫植食性哺乳动物食的研究进展,植物组织的次生化合物主要为酚类、萜类及含N类化合物,植物对动物觅食的化学防卫对策以次生化合物的各类而有差异,次生化合物通过对动物的食物摄入、消化、代谢,以及敏殖活动的效应,以抵御动物的觅食。将植物化学防卫与动物适应对策相结合,探讨动物－植物协同进化模式,是该研究领域的主要发展趋势。     

Nitrogen and water affect direct and indirect plant systemic induced defense in cotton

Plants have direct and indirect constitutively produced and inducible defenses against herbivores and pathogens, which can substantially aid in their ability to defend themselves. However, very little is known about the influence of agronomic factors on such defenses. Here, we tested the effects of nitrogen levels and water availability on the ability of cotton plants to deter feeding by Spodoptera exigua through induction of anti-feedants, and to attract Microplitis croceipes through systemic induction of volatile emission. Cotton plants were grown with various nitrogen levels and were either exposed to water stress or normal water before being exposed to S. exigua for 48 h for induction of defenses. Dual choices of various nitrogen and water treatments were provided to M. croceipes in flight tunnel bioassays. Dual choices of leaf tissue from the various nitrogen and water treatments were provided to S. exigua larvae. Both water stress and nitrogen levels under and over the recommended levels increased leaf tissue consumption and decreased attraction of M. croceipes to the plants. Analyses of induced volatiles released from herbivore damaged plants indicate that their concentrations differ among the nitrogen levels tested with plants receiving no nitrogen or twice the recommended dose having amounts much lower than plants receiving the recommended dose. Because both direct and indirect plant defense mechanisms are negatively affected by improper nitrogen and insufficient water, we argue that these factors should be considered for a better natural control of pests in cotton and most probably in other crops.     

High herbivore pressure favors constitutive over induced defense

Theoretical and empirical studies show that, when past or current herbivory is a reliable cue of future attack and defenses are costly, defenses can be induced only when needed and thereby permit investment in other functions such as growth or reproduction. Theory also states that, in environments where herbivory is constantly high, constitutive defenses should be favored. Here, we present data to support the second aspect of the induced resistance hypothesis. We examined herbivore‐induced responses for four species of Inga (Fabaceae), a common canopy tree in Neotropical forests. We quantified chemical defenses of expanding leaves, including phenolic, saponin and toxic amino acids, in experimental field treatments with and without caterpillars. Because young leaves lack fiber and are higher in protein than mature leaves, they typically lose >25% of their leaf area during the few weeks of expansion. We predicted that the high rates of attack would select for investment in constitutive defenses over induction. Our data show that chemical defenses were quite unresponsive to herbivory. We demonstrated that expanding leaves showed no or only small increases in investment in secondary metabolites, and no qualitative changes in the phenolic compound profile in response to herbivory. The proteinogenic amino acid tyrosine, which can be toxic at high concentrations, showed the greatest levels of induction. Synthesis: These results provide some of the first support for theoretical predictions that the evolution of induced vs. constitutive defenses depends on the risk of herbivory. In habitats with constant and high potential losses to herbivores, such as tropical rainforests, high investments in constitutive defenses are favored over induction.     

The pathway for systemic electrical signal conduction in the wounded tomato plant

The pathway of a systemic electrical signal possibly linking wounding and the systemic synthesis of proteinase inhibitor was investigated in tomato (Lycopersicon esculentumMill. cv. Moneymaker) plants. Heat, causing wounding to a cotyledon, was used to induce both a travelling electrical signal and systemic proteinase inhibitor activity. Intracellular recordings of changes in the membrane potential of different cell types were measured in the petiole of leaf 1, the first true leaf, and impaled cells were identified by injection of fluorescent dye (Lucifer Yellow CH). No difference was found between the membrane potentials of the different cell types; the mean membrane potential of all the cell types was -148 ± 3 mV. Only sieve-tube elements and companion cells produced large (79 ± 3.3 mV) action-potential-like depolarisations following wounding, although smaller (23 ± 1.6 mV) depolarisations were observed in other cell types. It was concluded that the electrical signal possibly linking a wound stimulus in a cotyledon with the induction of systemic proteinase inhibitor synthesis was propagated in the sieve-tube element/companion cell complex.Abbreviations LYCH Lucifer Yellow CH - PI proteinase inhibitor This work was supported by the Biotechnology and Biological Sciences Research Council (UK).     

Host plant defense signaling in response to a coevolved herbivore combats introduced herbivore attack

Defense‐free space resulting from coevolutionarily naïve host plants recently has been implicated as a factor facilitating invasion success of some insect species. Host plants, however, may not be entirely defenseless against novel herbivore threats. Volatile chemical‐mediated defense signaling, which allows plants to mount specific, rapid, and intense responses, may play a role in systems experiencing novel threats. Here we investigate defense responses of host plants to a native and exotic herbivore and show that (1) host plants defend more effectively against the coevolved herbivore, (2) plants can be induced to defend against a newly‐associated herbivore when in proximity to plants actively defending against the coevolved species, and (3) these defenses affect larval performance. These findings highlight the importance of coevolved herbivore‐specific defenses and suggest that naïveté or defense limitations can be overcome via defense signaling. Determining how these findings apply across various host–herbivore systems is critical to understand mechanisms of successful herbivore invasion.     

Control of systemically induced herbivore resistance by plant vascular architecture

Patterns of systemically induced resistance (SIR) in Eastern Cottonwood, Populus deltoides, measured by reduced feeding of the leaf-chewing beetle, Plagiodera versicolora, were shown to be directly related to the distribution of the plant vasculature. Mechanical damage to single leaves resulted in SIR in non-adjacent, orthostichous leaves (vertically aligned on the stem) with direct vascular connections, both up and down the shoot; but no SIR in adjacent, non-orthostichous leaves with less direct vascular connections. The control that the plant vasculature exerts over signal distribution following wounding can therefore be used to predict SIR patterns, explain variation in the distribution of SIR, and relate this ecologically important phenomenon to biochemical processes of systemic gene expression and biochemical resistance mechanisms.     

Herbivore-induced ethylene suppresses a direct defense but not a putative indirect defense against an adapted herbivore

Herbivory induces both direct and indirect defenses in plants; however, some combinations of these defenses may not be compatible. The jasmonate signal cascade activated both direct (nicotine accumulations) and indirect (mono- and sesquiterpene emissions) whole-plant defense responses in the native tobacco Nicotiana attenuata Torr. Ex Wats. Nicotine accumulations were proportional to the amount of leaf wounding and the resulting increases in jasmonic acid (JA) concentrations. However, when larvae of the nicotine-tolerant herbivore, Manduca sexta, fed on plants or their oral secretions were applied to leaf punctures, the normal wound response was dramatically altered, as evidenced by large (4- to 10-fold) increases in the release of (i) volatile terpenoids and (ii) ethylene, (iii) increased (4- to 30-fold) accumulations of endogenous JA pools, but (iv) decreased or unchanged nicotine accumulations. The ethylene release, which was insensitive to inhibitors of induced JA accumulation, was sufficient to account for the attenuated nicotine response. Applications of ethylene and ethephon suppressed the induced nicotine response and pre-treatment of plants with a competitive inhibitor of ethylene receptors, 1-methylcyclopropene, restored the full nicotine response. This ethylene burst, however, did not inhibit the release of volatile terpenoids. Because parasitoids of Manduca larvae are sensitive to the dietary intake of nicotine by their hosts, this ethylene-mediated switching from direct to a putative indirect defense may represent an adaptive tailoring of a plant's defense response. Received: 13 June 1999 / Accepted: 21 August 1999     

Bacterial phytopathogen infection disrupts belowground plant indirect defense mediated by tritrophic cascade

Plants can defend themselves against herbivores through activation of defensive pathways and attraction of third‐trophic‐level predators and parasites. Trophic cascades that mediate interactions in the phytobiome are part of a larger dynamic including the pathogens of the plant itself, which are known to greatly influence plant defenses. As such, we investigated the impact of a phloem‐limited bacterial pathogen, Candidatus Liberibacter asiaticus (CLas), in cultivated citrus rootstock on a well‐studied belowground tritrophic interaction involving the attraction of an entomopathogenic nematode (EPN), Steinernema diaprepesi, to their root‐feeding insect hosts, Diaprepes abbreviatus larvae. Using belowground olfactometers, we show how CLas infection interferes with this belowground interaction by similarly inducing the release of a C12 terpene, pregeijerene, and disconnecting the association of the terpene with insect presence. D. abbreviatus larvae that were not feeding but in the presence of a CLas‐infected plant were more likely to be infected by EPN than those near uninfected plants. Furthermore, nonfeeding larvae associated with CLas‐infected plants were just as likely to be infected by EPN as those near noninfected plants with D. abbreviatus larval damage. Larvae of two weevil species, D. abbreviatus and Pachnaeus litus, were also more attracted to plants with infection than to uninfected plants. D. abbreviatus larvae were most active when exposed to pregeijerene at a concentration of 0.1 μg/μl. We attribute this attraction to CLas‐infected plants to the same signal previously thought to be a herbivore‐induced plant volatile specifically induced by root‐feeding insects, pregeijerene, by assessing volatiles collected from the roots of infected plants and uninfected plants with and without feeding D. abbreviatus. Synthesis. Phytopathogens can influence the structuring of soil communities extending to the third trophic level. Field populations of EPN may be less effective at host‐finding using pregeijerene as a cue in citrus grove agroecosystems with high presence of CLas infection.     

植物挥发性信号物质介导抗性的生态功能

植物产生的挥发性化合物能够作为媒介参与植物与周围环境之间的信息交流及相互作用。植物挥发性物质在吸引传粉者、促进种子传播、抑制其它植物种子萌发等方面具有重要的作用。近年来,关于植物挥发性物质在生态系统中的信号作用研究已经成为国内外的研究热点,受到广泛关注。总结了植物挥发性物质作为信号物质在提高植物抗性方面的国内外研究成果,阐述了植物挥发性物质不仅能够直接提高植物的抗性,而且可以作为信号物质在同株、同种异株和不同种植物之间进行传递,进而间接提高目标植物的抗性。最后,还对植物挥发性物质的研究方法和潜在的生态功能进行了探讨。     

The oxidative burst in plant defense: Function and signal transduction

The rapid production and accumulation of active oxygen species (AOS), the oxidative burst, has been shown to occur in a variety of plant/pathogen systems. In particular, two species, hydrogen peroxide (H₂O₂) and the superoxide radical anion O₂^? have received considerable attention. H₂O₂ and O₂^?, while acting directly as antimicrobial agents, may also serve as second messengers or catalysts in plants to activate a more diverse set of defense responses. Some of the better studied downstream responses promoted by AOS are (1) the cross-linking of cell wall proteins, (2) the induction of defense-related genes, (3) the stimulation of phytoalexin biosynthesis and (4) promotion of the hypersensitive response (HR). A useful model for studying the oxidative burst in plants is the neutrophil NADPH ox-idase complex, the primary source of AOS production in mammals. Several of the subunits of the neutrophil NADPH oxidase complex have been immunologically identified in plants. Furthermore, many of the components known to be involved in the signal transduction pathway in neutrophils have also been found to play a role in the oxidative burst in plants. Just as various ligands activate the oxidase complex in neutrophils, several ligands (elicitors or pathogens) also lead to induction of the oxidative burst in plant cells. The similarities between the neutrophil and plant oxidative bursts will be elaborated in this review. Following stimulation with elicitors, different signal transduction pathways are activated in plants, depending on the source of elicitor used. While the identities and chronologies of the major intermediates in these pathways remain largely unknown, there is strong evidence at least for participation of phospholipases, H⁺/K⁺ exchange, Ca²⁺ influxes, protein kinases and phosphatases, and GTP binding proteins. In an effort to integrate these various signaling events into a single scheme, we have constructed a hypothetical model that proposes how different elicitors might induce the oxidative burst in the same cell by different pathways.     

The jasmonate pathway alters herbivore feeding behaviour: consequences for plant defences

The jasmonate pathway is a highly conserved defensive cascade in plants that regulates the induction of resistance against herbivores; however, its role in herbivore feeding behaviour remains unknown. We used a mutant tomato plant (def‐1) deficient in the production of jasmonate‐related defensive proteins to test the hypothesis that genotypes with a reduced ability to induce resistance have a higher and more concentrated pattern of herbivore damage. Wild‐type and def‐1 plants received either damage by Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) caterpillars or no damage. After treatment, we tested for systemic responses by allowing a free roaming S. exigua caterpillar to feed on the undamaged portions of plants. Weight‐gain and leaf consumption of S. exigua were highest on def‐1 plants, regardless of prior herbivore damage. Def‐1 plants also had fewer numbers of leaves and leaflets eaten, and fewer feeding holes, which was indicative of a more concentrated distribution of damage on mutant compared to wild‐type plants. Following these results, we mimicked the amount and distribution of feeding damage that wild‐type or jasmonate‐deficient plants would receive on wild‐type plants to test whether changes in feeding behaviour may feedback to influence the expression of induced resistance. We mimicked the distribution of damage in wild‐type and jasmonate‐deficient plants by allowing caterpillars to feed on either one (leaf 1 or 2) or two leaves (leaf 1 and 2). Increased herbivore damage resulted in higher proteinase inhibitor (PI) activity, a jasmonate‐regulated defensive protein, and lower S. exigua performance on wild‐type but not jasmonate‐deficient plants. Compared to undamaged plants, a concentrated pattern of herbivore damage increased systemic resistance; these induced responses were greater on leaflets with stronger vascular connections to the damaged leaf. A more dispersed pattern of caterpillar damage altered the expression of induced responses, but the outcome depended on the specific pattern of damage. When leaf 1 was damaged and then leaf 2, the undamaged (third) leaf (which is more strongly connected to leaf 1 than 2) expressed reduced the PI activity compared to plants receiving concentrated damage to leaf 1; whereas in plants where leaf 2 was first damaged and then leaf 1, there were no differences in PI activity in leaf 3 compared to plants receiving concentrated damage to leaf 2. Thus, induction of the jasmonate pathway may not only determine the amount and distribution of feeding damage by herbivores, but this may feedback to affect the subsequent expression of plant defence.     

Comparing the consequences of induced and constitutive plant resistance for herbivore population dynamics

Although it has been suggested that induced and constitutive plant resistance should have different effects on insect herbivore population dynamics, there is little experimental evidence that plant resistance can influence herbivore populations longer than one season. We used a density-manipulation experiment and model fitting to examine the effects of constitutive and induced resistance on herbivore dynamics over both the short and long term. We used likelihood methods to fit population dynamic models to recruitment data for populations of Mexican bean beetles on soybean varieties with no resistance, constitutive resistance, or induced resistance. We compared model configurations that fit parameters for resistance types separately to models that did not account for resistance type. Models representing the hypothesis that the three resistance types differed in their effects on beetle dynamics received the most support. Induced resistance resulted in lower population growth rates and stronger density dependence than no resistance. Constitutive resistance resulted in lower population growth rates and stronger density dependence than induced resistance. Constitutive resistance had a stronger effect on both short-term beetle recruitment and predicted beetle population dynamics than induced resistance. The results of this study suggest that induced and constitutive resistance can differ in their effects on herbivore populations even in a relatively complex system.     

Parameters affecting plant defense pathway mediated recruitment of entomopathogenic nematodes

Entomopathogenic nematodes are natural enemies and effective biological control agents of subterranean insect herbivores. Interactions between herbivores, plants, and entomopathogenic nematodes are mediated by plant defense pathways. These pathways can induce release of volatiles and recruit entomopathogenic nematodes. Stimulation of these plant defense pathways for induced defense against belowground herbivory may enhance biological control in the field. Knowledge of the factors affecting entomopathogenic nematode behaviour belowground is needed to effectively implement such strategies. To that end, we explore the effect of elicitor, elicitor dose, mechanical damage, and entomopathogenic nematode release distance on recruitment of entomopathogenic nematode infective juveniles to corn seedlings. Increasing doses of methyl jasmonate and methyl salicylate elicitors recruited more entomopathogenic nematodes as did mechanical damage. Recruitment of entomopathogenic nematodes was higher at greater release distances. These results suggest entomopathogenic nematodes are highly tuned to plant status and present a strategy for enhancing biological control using elicitor-stimulated recruitment of entomopathogenic nematodes.     

The circadian clock contributes to diurnal patterns of plant indirect defense in nature

The plant circadian clock regulates the rhythms of plant metabolism. Many herbivore-induced plant volatiles(HIPVs) fluctuate, diurnally, but the role of the circadian clock in the emission of HIPVs and their ecological consequences remains largely unknown.Here, we show that the timing of herbivore attack can alter the outcome of tri-trophic interactions, and this is mediated by the circadian clock, under both field and glasshouse conditions. Although most HIPV emissions did not have a circadian rhythm, the circadian clock modulated HIPV emissions in a time-dependent manner. HIPVs mediate tri-trophic interactions, and the circadian clock may affect these interactions by modulating HIPV emission in nature.