Rhizobacteria-mediated Induced Systemic Resistance: Triggering, Signalling and Expression

Selected strains of rhizosphere bacteria reduce disease by activating a resistance mechanism in the plant named rhizobacteria-mediated induced systemic resistance (ISR). Rhizobacteria-mediated ISR resembles pathogen-induced systemic acquired resistance (SAR) in that both types of induced resistance render uninfected plant parts more resistant towards a broad spectrum of plant pathogens. Some rhizobacteria trigger the salicylic acid (SA)-dependent SAR pathway by producing SA at the root surface. In other cases, rhizobacteria trigger a different signalling pathway that does not require SA. The existence of a SA-independent ISR pathway has been demonstrated in Arabidopsis thaliana. In contrast to pathogen-induced SAR, ISR induced by Pseudomonas fluorescens WCS417r is independent of SA accumulation and pathogenesis-related (PR) gene activation but, instead, requires responsiveness to the plant hormones jasmonic acid (JA) and ethylene. Mutant analyses showed that ISR follows a novel signalling pathway in which components from the JA and ethylene response are successively engaged to trigger a defensive state that, like SAR, is controlled by the regulatory factor NPR1. Interestingly, simultaneous activation of both the JA/ethylene-dependent ISR pathway and the SA-dependent SAR pathway results in an enhanced level of protection. Thus combining both types of induced resistance provides an attractive tool for the improvement of disease control. This review focuses on the current status of our research on triggering, signalling, and expression of rhizobacteria-mediated ISR in Arabidopsis.     

The Relationship Between Pathogen-induced Systemic Resistance (ISR) and Multigenic (horizontal) Resistance in Plants

Plants have developed mechanisms to successfully co-exist in the presence of pathogenic organisms. Some interactions between plants and pathogens are based on recognition of specific elicitor molecules from avirulent pathogen races (avr gene products), which is described in the gene-for-gene resistance theory. Another type of resistance, multigenic (horizontal) resistance, is a less well-studied phenomenon that depends upon multiple genes in the plant host. All plants possess resistance mechamisms which can be induced upon pre-treatment of plants with a variety of organisms or compounds. This general phenomenon is known as induced systemic resistance (ISR). At least in some plant species, ISR depends on the timely accumulation of multiple gene products, such as hydrolytic enzymes, peroxidases or other gene products related to plant defences. The pre-treatment of plants with an inducing organism or compound appears to incite the plant to mount an effective defense response upon subsequent encounters with pathogens, converting what would have been a compatible interaction to an incompatible one. Our studies in three plant–pathogen systems clearly document that multigenic-resistant plants constitutively express specific isozymes of hydrolytic enzymes that release cell wall elicitors, which in turn may activate other defense mechanisms. ISR induces constitutive accumulation of these and other gene products prior to challenge. ISR is known to function against multiple organisms, and there is no specificity observed in the accumulation patterns of defense-related gene products when ISR is induced. It is therefore hypothesized that the constitutive accumulation of specific isozymes of hydrolytic enzymes, or other defense related gene products, is an integral part of both multigenic resistance and the phenomenon of ISR. Further, plants in which ISR has been activated appear to move from a latent resistance state to one in which a multigenic, non-specific form of resistance is active.     

Costs and trade-offs associated with induced resistance

Plants resist attack by pathogens and herbivorous insects through constitutive and inducible defences. Based on differences in signalling pathways and spectra of effectiveness, different types of induced resistance have been defined. Systemic acquired resistance (SAR) occurs in distal plant parts following localized infection by a necrotizing pathogen. It is controlled by a signalling pathway that depends upon the accumulation of salicylic acid (SA) and the regulatory protein NPR1. In contrast, induced systemic resistance (ISR) is induced by selected strains of non-pathogenic plant growth promoting bacteria (PGPR). ISR functions independently of SA, but requires NPR1 and is regulated by jasmonic acid (JA) and ethylene (ET). It is generally believed that induced resistance evolved to save energy under pathogen or insect-free conditions, although costs still arise when defences are activated following attack. Costs can arise from the allocation of resources to defence and away from plant growth and development, and there are also ecological costs which result from trade-offs between induced resistance and the plant's interaction with beneficial organisms e.g. mycorrhizal fungi. To date, few studies have examined the costs and trade-offs associated with induced resistance to pathogens. There is a clear need for long-term studies of costs and trade-offs associated with induced resistance in crops under commercial conditions. Without such information, the potential offered by induced resistance is unlikely to be realized.     

内生细菌EBS05对烟草诱导抗性的信号转导途径研究

 樟树内生枯草芽胞杆菌EBS05是一株对多种植物病原菌具有较强拮抗活性,并能诱导烟草系统抗性的生防菌株。本文以缺失Surfactin A合成相关基因的突变菌株EBS05^T为材料,研究了内生细菌EBS05对烟草诱导系统抗性的激发子及其信号转导途径。结果表明,菌株EBS05产生的Surfactin A是诱导烟草对TMV系统抗性的有效激发子;Surfactin A诱导处理后,SA信号转导途径下游的关键调节基因NPR1首先被激活,并持续超量表达,进而触发PR1b和PR1a基因持续超量表达,表明Surfactin A诱导烟草对TMV的系统抗性是通过激活SA信号转导途径实现的。同时,Surfactin A诱导处理后24~72 h,JA/ET信号转导途径调节基因PDF1.2被激活,且超量表达,表明在Surfactin A诱导烟草对TMV系统抗性的信号转导过程中,可能存在SA信号途径和JA/ET信号途径的交叉协同作用。     

Dimethyl disulfide is an induced systemic resistance elicitor produced by Bacillus cereus C1L

BACKGROUND: Bacillus cereus C1L is a plant growth‐promoting rhizobacterium and can elicit induced systemic resistance (ISR) in plants against necrotrophic pathogens. However, little is known about ISR elicitors produced by B. cereus C1L, and no ISR elicitor has been identified and characterised. Therefore, the objective of this study is to identify volatile ISR elicitor(s) produced by B. cereus C1L. RESULTS: The volatile metabolites produced by B. cereus C1L were extracted, separated and identified by solid‐phase microextraction, gas chromatography and mass spectrometry. Dimethyl disulfide (DMDS) was the only separated metabolite being determined. Afterwards, application of DMDS by means of soil drench significantly protected tobacco and corn plants against Botrytis cinerea and Cochliobolus heterostrophus, respectively, under greenhouse conditions. The results reveal that DMDS could play an important role in ISR by B. cereus C1L. CONCLUSION: This is the first report of DMDS as an elicitor produced by an ISR‐eliciting B. cereus strain and its ability to suppress plant fungal diseases under greenhouse conditions. It is suggested that DMDS has potential for practical use in controlling plant foliar diseases besides soil fumigation. Copyright © 2012 Society of Chemical Industry     

Concepts and Direction of Induced Systemic Resistance in Plants and its Application

Resistance to plant disease is often specific and metabolites and receptors contributing to this specificity may have specific structures. However, simple, structurally-unrelated compounds induce systemic resistance in unrelated plants to diverse pathogens including fungi, bacteria and viruses. Both resistance and induced systemic resistance (ISR) are associated with the rapid accumulation of the same structurally unrelated putative defense compounds that have diverse functions. It has been suggested that cultivar (race)-specific resistance is initiated by the specific interaction of a pathogen product (or pathogen induced product) and a plant receptor. However, restricted infection by pathogens can result in ISR and many different compounds can cause ISR. It is thus evident that there are both specific and non-specific routes to the master switch for ISR and there may be more than one master switch. Are reactive oxygen species and free radicals regulating the master switch(es) via both routes? It is also evident there are many switches, other than the master switch. Adding to the complexity of resistance and ISR are the observations that different compounds and pathways may mediate different biochemical resistances. Activation of one of the pathways may antagonize or enhance the activation or effectiveness of another. The review will address these complexities and questions and propose directions of research which require high priority. Factors which encourage and suppress the application of ISR in agriculture will also be addressed.     

The Ecological Concept of Costs of Induced Systemic Resistance (ISR)

Plant defence is thought to provide benefits for the defended plants. Theoretical concepts must, therefore, explain why there is variation in defensive traits, which naively might be assumed to be present constitutively in fixed high amounts. Explanations are mainly based on the assumption of fitness costs. Investment in defence is thought to reduce the fitness of plants in enemy-free environments. Fitness costs often result from allocation costs, i.e. allocation of limited resources to defence, which then cannot be used for growth or other fitness-relevant processes. This theoretical concept can provide a useful tool for the interpretation of induced plant responses against pathogens, named induced systemic (or systemic acquired) resistance (ISR or SAR). Phenotypic plasticity, leading to induced responses, might have evolved mainly to reduce costs, since investment in defence is restricted to situations actually requiring defence. ISR can incur allocation costs and other, indirect costs, which ultimately may lead to fitness costs. Evolution of any defensive trait depends on both what a plant ideally 'should do and what it actually 'is able to do. Costs of defence constrain its expression. This might have important influences on the evolution of plant defensive traits, as well as on the exploitation of natural defences in agricultural crop protection.     

Induced Systemic Resistance (ISR) by Pseudomonas spp. Impairs Pre- and Post-Infection Development of Pythium aphanidermatum on Cucumber Roots

The effect of induced systemic resistance (ISR) by Pseudomonas rhizobacteria on the pre- and post-infection development of Pythium aphanidermatum on cucumber roots was investigated. Cucumber plants (cv. Corona) were grown in vermiculite, roots were split with one side bacterized with Pseudomonas corrugata strain 13 or P. aureofaciens strain 63-28 (bacterized roots) and the other distant side was treated with water (distant, induced roots). For the non-induced control, roots on the bacterized side were treated with buffer instead of the bacterial treatment. Intact, non-split roots were also treated with the bacteria or buffer as a control. Cucumber root tissue from these treatments were harvested and incubated with a zoospore suspension of P. aphanidermatum for three hours. Most of the zoospores in the suspension were stimulated to encyst or germinate. The numbers of germinated zoospores were significantly decreased on distant induced cucumber roots in comparison to non-induced controls. Germination was also reduced on intact bacterized roots, compared to controls. There was less attachment, germ tube production and penetration on roots bacterized or induced by the rhizobacteria compared to non-induced roots. Effects were significantly greater on bacterized roots (roots colonized by bacteria) compared to distant induced roots (roots with the opposite side bacterized). Systemic resistance induced by the two Pseudomonas spp. also reduced pathogen spread on split cucumber roots in planta. Crown infection from induced or bacterized roots was delayed for four to six days in comparison to the non-induced control. Results indicated that Pseudomonas spp. can exert both an indirect influence on P. aphanidermatum zoospore behaviour and infection via induced systemic resistance (ISR) and a local influence via antibiosis or local induced resistance.     

Assessment of resistance pathways induced in Arabidopsis thaliana by hypovirulent Rhizoctonia spp. isolates

Certain hypovirulent Rhizoctonia isolates effectively protect plants against well-known important pathogens among Rhizoctonia isolates as well as against other pathogens. The modes of action involved in this protection include resistance induced in plants by colonization with hypovirulent Rhizoctonia isolates. The qualifications of hypovirulent isolates (efficient protection, rapid growth, effective colonization of the plants, and easy application in the field) provide a significant potential for the development of a commercial microbial preparation for application as biological control agents. Understanding of the modes of action involved in protection is important for improving the various aspects of development and application of such preparations. The hypothesis of the present study is that resistance pathways such as systemic acquired resistance (SAR), induced systemic resistance (ISR), and phytoalexins are induced in plants colonized by the protective hypovirulent Rhizoctonia isolates and are involved in the protection of these plants against pathogenic Rhizoctonia. Changes in protection levels of Arabidopsis thaliana mutants defective in defense-related genes (npr1-1, npr1-2, ndr1-1, npr1-2/ndr1-1, cim6, wrky70.1, snc1, and pbs3-1) and colonized with the hypovirulent Rhizoctonia isolates compared with that of the wild type (wt) plants colonized with the same isolates confirmed the involvement of induced resistance in the protection of the plants against pathogenic Rhizoctonia spp., although protection levels of mutants constantly expressing SAR genes (snc1 and cim6) were lower than that of wt plants. Plant colonization by hypovirulent Rhizoctonia isolates induced elevated expression levels of the following genes: PR5 (SAR), PDF1.2, LOX2, LOX1, CORI3 (ISR), and PAD3 (phytoalexin production), which indicated that all of these pathways were induced in the hypovirulent-colonized plants. When SAR or ISR were induced separately in plants after application of the chemical inducers Bion and methyl jasmonate, respectively, only ISR activation resulted in a higher protection level against the pathogen, although the protection was minor. In conclusion, plant colonization with the protective hypovirulent Rhizoctonia isolates significantly induced genes involved in the SAR, ISR, and phytoalexin production pathways. In the studied system, SAR probably did not play a major role in the mode of protection against pathogenic Rhizoctonia spp.; however, it may play a more significant role in protection against other pathogens.     

Plant immunisation: from myth to SAR

The idea that plants might be able to develop a form of acquired immunity to infection following exposure to a pathogen has been current ever since discovery of the animal immune system in the later years of the nineteenth century. Early attempts to demonstrate a comparable system in plants focused on the detection of precipitating antibodies and hence were doomed to failure. Nevertheless, largely anecdotal evidence for plant immunisation continued to accumulate, culminating in the discovery of phytoalexins in the 1940s. Convincing evidence for systemic changes in plant resistance following an inducer inoculation was not available until 20 years later, when pioneering work on tobacco infected with blue mould (Peronospora tabacina) or tobacco mosaic virus (TMV) showed that tissues remote from the inoculation site were altered in disease reaction type. Increased resistance was expressed as a reduction in lesion numbers and size, and a reduced rate of pathogen reproduction. Systemic acquired resistance (SAR) has now been demonstrated in at least 20 plant species in at least six plant families, although detailed genetic or molecular analysis has mainly been confined to a few models, such as tobacco, cucumber and Arabidopsis. SAR is associated with the coordinate induction of genes encoding defence proteins which can be used as molecular markers of the response. The availability of Arabidopsis mutants altered in the induction and expression of SAR is now providing new insights into the signal transduction pathway(s) involved, and will enable comparison with the molecular mechanisms operating in other plant taxa. Important unresolved questions concern the nature of the translocated signal, the mechanism of defence ‘priming’, efficacy of the response against different pathogens, and practical exploitation of SAR in crop protection. The first generation of chemical plant defence activators is now commercially available and optimal use of these SAR inducers in integrated disease control requires further evaluation. The prospects for engineering transgenic crops altered in the regulation or expression of SAR is also a subject for further investigation. © 1999 Society of Chemical Industry     

Systemic resistance induced in Arabidopsis thaliana by Trichoderma asperellum SKT-1, a microbial pesticide of seedborne diseases of rice

BACKGROUND: Trichoderma asperellum SKT-1 is a microbial pesticide of seedborne diseases of rice. To investigate the mechanisms of disease suppression in SKT-1, the ability to induce systemic resistance by SKT-1, or its cell-free culture filtrate (CF), was tested using Arabidopsis thaliana Col-0 plants. RESULTS: Both SKT-1 and its CF elicit an induced systemic resistance against the bacterial leaf speck pathogen Pseudomonas syringae pv. tomato DC3000 in Col-0 plants. Involvement of plant hormones in the induced resistance by SKT-1 and CF was assessed using Arabidopsis genotypes such as the jasmonic acid (JA)-resistant mutant jar1, the ethylene (ET)-resistant mutant etr1, the plant impaired in salicylic acid (SA) signalling transgenic NahG and the mutant npr1 impaired in NPR1 activity. In soil experiments using SKT-1, no significant disease suppression effect was observed in NahG transgenic plants or npr1 mutant plants. Expression levels of SA-inducible genes such as PR-1, PR-2 and PR-5 increased substantially in the leaves of Col-0 plants. Expression levels of JA/ET-induced genes such as PDF1.2a, PR-3, PR-4 and AtVsp1 were also induced, but the levels were not as high as for SA-inducible genes. In a hydroponic experiment using CF from SKT-1, all Arabidopsis genotypes showed an induced systemic resistance by CF and increased expression levels of JA/ET- and SA-inducible genes in leaves of CF-treated plants. CONCLUSION: The SA signalling pathway is important in inducing systemic resistance to colonisation by SKT-1, and both SA and JA/ET signalling pathways combine in the signalling of induced resistance by CF. These results indicate that the response of A. thaliana is different from that found in root treatments with barley grain inoculum and CF from SKT-1. Copyright © 2011 Society of Chemical Industry     

Study of Bacterial Determinants Involved in the Induction of Systemic Resistance in Bean by Pseudomonas putida BTP1

The ability of Pseudomonas putida BTP1 to induce resistance in bean to Botrytis cinerea was demonstrated in soil experiments on plants pre-inoculated at the root level with the bacteria before challenge with the leaf pathogen. As a first step to characterize the molecules from BTP1 responsible for induction of systemic resistance in bean, heat-killed cells and supernatant from culture in an iron-limited medium were tested for their protective effect. Most of the resistance-eliciting activity of the strain was retained in the crude cell-free culture fluid. In vivo assays with samples from successive fractionation steps of the BTP1 supernatant led, (i) to the conclusion that salicylic acid, pyochelin and pyoverdin, previously identified as Pseudomonas determinants for induced systemic resistance (ISR), were not involved in systemic resistance triggered by BTP1, and (ii) to the isolation of fractions containing one main metabolite that retained most of the resistance-inducing activity in bean. Although this molecule remains to be structurally characterized, its isolation is an addition to the range of determinants from plant growth-promoting rhizobacteria (PGPR) known to stimulate plant defences.     

Colonization of <Emphasis Type="Italic">Arabidopsis</Emphasis> roots by <Emphasis Type="Italic">Trichoderma atroviride</Emphasis> promotes growth and enhances systemic disease resistance through jasmonic acid/ethylene and salicylic acid pathways

Trichoderma spp. are common soil fungi used as biocontrol agents due to their capacity to produce antibiotics, induce systemic resistance in plants and parasitize phytopathogenic fungi of major agricultural importance. The present study investigated whether colonization of Arabidopsis thaliana seedlings by Trichoderma atroviride affected plant growth and development. Here it is shown that T. atroviride promotes growth in Arabidopsis. Moreover, T. atroviride produced indole compounds in liquid cultures. These results suggest that indoleacetic acid-related indoles (IAA-related indoles) produced by T. atroviride may have a stimulatory effect on plant growth. In addition, whether colonization of Arabidopsis roots by T. atroviride can induce systemic protection against foliar pathogens was tested. Arabidopsis roots inoculation with T. atroviride provided systemic protection to the leaves inoculated with bacterial and fungal pathogens. To investigate the possible pathway involved in the systemic resistance induced by T. atroviride, the expression profile of salicylic acid, jasmonic acid/ethylene, oxidative burst and camalexin related genes was assessed in Arabidopsis. T. atroviride induced an overlapped expression of defence-related genes of SA and JA/ET pathways, and of the gene involved in the synthesis of the antimicrobial phytoalexin, camalexin, both locally and systemically. This is the first report where colonization of Arabidopsis roots by T. atroviride induces the expression of SA and JA/ET pathways simultaneously to confer resistance against hemibiotrophic and necrotrophic phytopathogens. The beneficial effects induced by the inoculation of Arabidopsis roots with T. atroviride and the induction of the plant defence system suggest a molecular dialogue between these organisms.     

植物系统性获得抗病性的产生机理和途径

坏死型病原物侵染或某些生化制剂诱导处理后,植株未受侵染或处理部位产生对随后病原物侵染的抗性,称为植物系统性获得抗性,ＳＡＲ具有抗性表现系统、持久、抗病对象广谱三大特点。坏死型病原物侵染或某些生化制剂处理后,植株受处理部位迅速产生系统性信号,经韧皮部传导到未侵染或处理部位,诱发ＳＡＲ基因表达。水杨酸是诱发ＳＡＲ的系统性信号之一。此外,上部非处理部位处于敏化状态,能更迅速有效地产生针对挑战接种病原物的     

Tobacco cultivars vary in induction of systemic resistance against <Emphasis Type="Italic">Cucumber mosaic virus</Emphasis> and growth promotion by <Emphasis Type="Italic">Pseudomonas chlororaphis</Emphasis> O6 and its <Emphasis Type="Italic">gacS</Emphasis> mutant

The colonization of plant roots with certain rhizosphere bacteria promotes plant growth and induces long lasting systemic protection against a broad spectrum of plant pathogens. The role of the global regulator, GacS, in the rhizosphere colonist Pseudomonas chlororaphis O6 in stimulating growth promotion and induced resistance against Cucumber mosaic virus was examined in tobacco. Responses were compared in tobacco cvs Samsun and GX3. Root colonization of Samsun with wild-type O6 and the gacS complemented mutant-elicited reduced viral symptoms and viral titre. On GX3, there was little affect on symptoms when roots were colonized by the wild-type, gacS mutant or complemented mutant but colonization by both the wild-type and the gacS mutant lowered viral titre. Wild-type O6 and the gacS mutant caused plant growth to be maintained in both tobacco cultivars after viral infection, although the affect was stronger with GX3 than Samsun. In contrast, although a chemical inducer, benzothiadiazole, reduced symptoms and viral titre in both cultivars, plant growth was suppressed. Our results indicate rhizobacteria-elicited induced viral resistance without a negative impact on growth but there was a differential response between cultivars. Detailed knowledge regarding the mechanisms inherent to these differences between cultivars requires further investigation.     

Application of Rhizobacteria for Induced Resistance

This article provides a review of experiments conducted over a six-year period to develop a biological control system for insect-transmitted diseases in vegetables based on induced systemic resistance (ISR) mediated by plant growth-promoting rhizobacteria (PGPR). Initial experiments investigated the factors involved in treatment with PGPR led to ISR to bacterial wilt disease in cucumber caused by Erwinia tracheiphila. Results demonstrated that PGPR-ISR against bacterial wilt and feeding by the cucumber beetle vectors of E. trachiphiela were associated with reduced concentrations of cucurbitacin, a secondary plant metabolite and powerful beetle feeding stimulant. In other experiments, PGPR induced resistance against bacterial wilt in the absence of the beetle vectors, suggesting that PGPR-ISR protects cucumber against bacterial wilt not only by reducing beetle feeding and transmission of the pathogen, but also through the induction of other plant defense mechanisms after the pathogen has been introduced into the plant. Additional greenhouse and field experiments are described in which PGPR strains were selected for ISR against cucumber mosaic virus (CMV) and tomato mottle virus (ToMoV). Although results varied from year to year, field-grown tomatoes treated with PGPR demonstrated a reduction in the development of disease symptoms, and often a reduction in the incidence of viral infection and an increase in tomato yield. Recent efforts on commercial development of PGPR are described in which biological preparations containing industrial formulated spores of PGPR plus chitosan were formulated and evaluated for use in a transplant soil mix system for developing plants that can withstand disease attack after transplanting in the field.     

Increased rhizosphere populations of Trichoderma asperellum strain T34 caused by secretion pattern of root exudates in tomato plants inoculated with Botrytis cinerea

Root exudates secreted from plants can modify rhizosphere microbiota by enhancing or inhibiting the growth of biological control agents (BCAs) and/or pathogens. Similarly, microorganisms can modify the secretion of plant root exudates. The aim of this study was to analyse the effect of a Botrytis cinerea leaf infection on the secretion of tomato root exudates and on the populations of the BCA Trichoderma asperellum strain T34 (T34). This study found that the secretion pattern of root exudates in tomato plants was influenced by B. cinerea infection in plant leaves. An increase in the levels of gluconic acid was observed, while levels of sucrose and inositol decreased. A decrease in the severity of B. cinerea by the induction of systemic resistance triggered by T34 was also observed. Tomato plants infected with B. cinerea maintained the populations of T34 in the roots, while populations of T34 decreased in plants not inoculated with the pathogen. Samples exposed to media containing gluconic acid (as the only carbon source or at the same concentration found in roots exudates) saw an increase in the in vitro growth of T34 compared to media without gluconic acid. In conclusion, a change in the secretion pattern of root exudates caused by B. cinerea, together with the enhanced growth of T34 in the presence of gluconic acid, indicates the existence of leaf to root communication. The result of this is enhanced populations of T34, and in turn induced disease resistance and a consequential reduction in disease severity.     

Induction of transient ethylene and reduction in severity of tomato bacterial wilt by Pythium oligandrum

Pythium oligandrum (PO) is a mycoparasite on a wide range of fungi and suppresses diseases caused by fungal pathogens when colonizing the rhizosphere. PO and its cell wall proteins (CWPs) have elicitor activity that induces defence responses in plants. The potential of a mycelial homogenate of PO to suppress bacterial diseases was studied in roots of tomato ( Lycopersicon esculentum cv. Micro-Tom) inoculated with Ralstonia solanacearum . PO-treated plants showed enhanced resistance to R. solanacearum and reduction in severity of wilt symptoms. As ethylene often acts as one of the signal molecules for induced resistance, its production following treatment of tomato roots with the mycelial homogenate or CWP of PO was measured. The level of ethylene in PO- and CWP-treated plants was transiently elevated six- to 11-fold at 4–8 h after treatment, followed by high expression of three basic ethylene-inducible defence-related genes ( PR-2b , PR-3b and PR-5b ). Analysis of PR-5b gene expression in the leaves of PO- and CWP-treated plants suggested that PR gene expression was induced systemically. The expression of LeERF2 and LeETR4 , which confer an ethylene-dependent signalling pathway, was also significantly accelerated by such treatments. These results indicate that PO has the potential to control bacterial wilt disease and that CWP may play an important role in the induction of resistance to R. solanacearum accompanying the activation of the ethylene-dependent signalling pathway.     

Induction of systemic disease resistance and pathogen defence responses in Asparagus officinalis inoculated with nonpathogenic strains of Fusarium oxysporum

The ability of nonpathogenic isolates of Fusarium oxysporum (np Fo ) to induce systemic resistance and defence responses against subsequent challenge with a pathogenic strain of F. oxysporum f. sp. asparagi ( Foa ) was examined in Asparagus officinalis . In a split-root experiment, roots inoculated with np Fo exhibited a hypersensitive response and those subsequently inoculated with Foa displayed resistance. Induction of systemic resistance in np Fo -treated plants led to significantly fewer necrotic lesions ( P  = 0·05) and reduced Foa disease severity compared with plants not treated with np Fo . In hyphal-sandwich root inoculation experiments, activities of peroxidase and phenylalanine ammonia-lyase and lignin content were higher in np Fo -treated plants and increased more rapidly than in np Fo -untreated plants after Foa inoculation. Antifungal activity (inhibition of fungal spore germination and germ-tube growth) from exudates of roots inoculated with Foa were observed for np Fo -treated plants but not for np Fo -untreated plants. Thus, isolates of np Fo may function as inducers of systemic acquired resistance (SAR) and defence responses against Foa invasion in A. officinalis .