Chromatin modification acts as a memory for systemic acquired resistance in the plant stress response

Priming of defence genes for amplified response to secondary stress can be induced by application of the plant hormone salicylic acid or its synthetic analogue acibenzolar S‐methyl. In this study, we show that treatment with acibenzolar S‐methyl or pathogen infection of distal leaves induce chromatin modifications on defence gene promoters that are normally found on active genes, although the genes remain inactive. This is associated with an amplified gene response on challenge exposure to stress. Mutant analyses reveal a tight correlation between histone modification patterns and gene priming. The data suggest a histone memory for information storage in the plant stress response.     

Abscisic acid modulates salicylic acid biosynthesis for systemic acquired resistance in tomato

Among the regulatory mechanisms of systemic acquired resistance (SAR) in tomato, antagonistic interaction between salicylic acid (SA) and abscisic acid (ABA) signaling pathways was investigated. Treatment with 1,2-benzisothiazol-3(2H)-one1,1-dioxide (BIT) induced SAR in tomato thorough SA biosynthesis. Pretreatment of ABA suppressed BIT-induced SAR including SA accumulation, suggesting that ABA suppressed SAR by inhibiting SA biosynthesis.     

Nitric oxide and reactive oxygen species are required for systemic acquired resistance in plants

Systemic acquired resistance (SAR) is a form of broad-spectrum disease resistance that is induced in response to primary infection and that protects uninfected portions of the plant against secondary infections by related or unrelated pathogens. SAR is associated with an increase in chemical signals that operate in a collective manner to confer protection against secondary infections. These include, the phytohormone salicylic acid (SA), glycerol-3-phosphate (G3P), azelaic acid (AzA) and more recently identified signals nitric oxide (NO) and reactive oxygen species (ROS). NO, ROS, AzA and G3P function in the same branch of the SAR pathway, and in parallel to the SA-regulated branch. NO and ROS function upstream of AzA/G3P and different reactive oxygen species functions in an additive manner to mediate chemical cleavage of the C9 double bond on C18 unsaturated fatty acids to generate AzA. The parallel and additive functioning of various chemical signals provides important new insights in the overlapping pathways leading to SAR.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demon-strated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the second-ary metabolites accumulation and is referred to as a copper-inducible elicitor (CIE). Finally, we discuss how copper-inducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

Plastid omega3-fatty acid desaturase-dependent accumulation of a systemic acquired resistance inducing activity in petiole exudates of Arabidopsis thaliana is independent of jasmonic acid

Systemic acquired resistance (SAR) is an inducible defense mechanism that is activated throughout the plant, subsequent to localized inoculation with a pathogen. The establishment of SAR requires translocation of an unknown signal from the pathogen-inoculated leaf to the distal organs, where salicylic acid-dependent defenses are activated. We demonstrate here that petiole exudates (PeXs) collected from Arabidopsis leaves inoculated with an avirulent (Avr) Pseudomonas syringae strain promote resistance when applied to Arabidopsis, tomato ( Lycopersicum esculentum ) and wheat ( Triticum aestivum ). Arabidopsis FATTY ACID DESATURASE7 ( FAD7 ), SUPPRESSOR OF FATTY ACID DESATURASE DEFICIENCY1 ( SFD1 ) and SFD2 genes are required for accumulation of the SAR-inducing activity. In contrast to Avr PeX from wild-type plants, Avr PeXs from fad7 , sfd1 and sfd2 mutants were unable to activate SAR when applied to wild-type plants. However, the SAR-inducing activity was reconstituted by mixing Avr PeXs collected from fad7 and sfd1 with Avr PeX from the SAR-deficient dir1 mutant. Since FAD7 , SFD1 and SFD2 are involved in plastid glycerolipid biosynthesis and SAR is also compromised in the Arabidopsis monogalactosyldiacylglycerol synthase1 mutant we suggest that a plastid glycerolipid-dependent factor is required in Avr PeX along with the DIR1- encoded lipid transfer protein for long-distance signaling in SAR. FAD7 -synthesized lipids provide fatty acids for synthesis of jasmonic acid (JA). However, co-infiltration of JA and methylJA with Avr PeX from fad7 and sfd1 did not reconstitute the SAR-inducing activity. In addition, JA did not co-purify with the SAR-inducing activity confirming that JA is not the mobile signal in SAR.     

Advances in research of induced resistance to insects in cotton

Any change in a plant that occurs following herbivory or environmental factors is an induced response. These changes include phytochemical induction, increases in physical defenses, emission of volatiles that attract predators and parasitoids of herbivores, and reduction in plant nutritional quality for herbivores, which is termed induced resistance. Induced resistance has been demonstrated ubiquitously in plants. It is one of our goals to review what is known about the induced resistance to herbivorous insects in cotton, including three resistance secondary metabolites (terpenoid, tannin, and flavonoids) that are contained at any significant levels of resistance to herbivorous insects in cotton cultivates. In many cases, the quantities or quality of secondary metabolites in plant are changed after attacked by insects. This review focuses on induced plant resistance as quantitative or qualitative enhancement of defense mechanism against insect pests, especially on the abiotic-elicitors-induced resistance in cotton plants. The abiotic-elicitor of cupric chloride, an exogenous inorganic compound, may induce the secondary metabolites accumulation and is referred to as a copperinducible elicitor (CIE). Finally, we discuss how copperinducible elicitor may be used in the Integrated Pest Management (IPM) system for cotton resistance control.     

3-Acetonyl-3-hydroxyoxindole: a new inducer of systemic acquired resistance in plants

Systemic acquired resistance (SAR) is an inducible defence mechanism which plays a central role in protecting plants from microbial pathogen attack. Guided by bioassays, a new chemical inducer of SAR was isolated from the extracts of Strobilanthes cusia and identified to be 3-acetonyl-3-hydroxyoxindole (AHO), a derivative of isatin. Tobacco plants treated with AHO exhibited enhanced resistance to tobacco mosaic virus (TMV) and to the fungal pathogen Erysiphe cichoracearum (powdery mildew), accompanied by increased levels of pathogenesis-related gene 1 ( PR-1 ) expression, salicylic acid (SA) accumulation and phenylalanine ammonia-lyase activity. To study the mode of action of AHO, its ability to induce PR-1 expression and TMV resistance in nahG transgenic plants expressing salicylate hydroxylase, which prevents the accumulation of SA, was analysed. AHO treatment did not induce TMV resistance or PR-1 expression in nahG transgenic plants, suggesting that AHO acts upstream of SA in the SAR signalling pathway. In addition, using two-dimensional gel electrophoresis combined with mass spectrometry, five AHO-induced plant proteins were identified which were homologous to the effector proteins with which SA interacts. Our data suggest that AHO may represent a novel class of inducer that stimulates SA-mediated defence responses.     

Induction of systemic resistance in rice by leaf extracts of Datura metel against sheath blight disease

The leaf extracts of Datura metel [both aqueous leaf extract (ALE) and ethanolic leaf extract (ELE)] were observed here to find if they can induce systemic resistance in the rice commonly found in Eastern India. The results showed that after the treatment, the enzyme activities of all the defence-related enzymes increased to a certain level even without pathogenic infection in comparison with non-treated seedlings and then, maintain at constant level throughout the study period. When treated seedlings were infected with Rhizoctonia solani, the enzyme activities were increased more than in uninfected seedlings. The elevated enzyme activities gave the indication of an induced systemic resistance in rice. The ELE of D. metel showed better induction effect than ALE.     

Induction of systemic acquired resistance in Zea mays L. by Aspergillus flavus and A. parasiticus derived elicitors

Host defence mechanisms can be elicited by using different elicitors produced from the pathogen/host. In this study, an effort has been made to study the effect of two fungal elicitors derived from Aspergillus flavus and A. parasiticus on induction of various defence-related enzymes in maize (Zea mays L.). Foliar application was done on 20-days-old maize plant with 10% A. flavus fungal culture filtrate (AFFCF) and A. parasiticus fungal culture filtrate (APFCF) as elicitors to trigger systemic acquired resistance (SAR). As a response of SAR, an increase in activities of phenylalanine ammonia lyase (PAL), peroxidase (POX), β-1,3-glucanase, nitrate reductase (NR) and nitrite reductase (NiR), total proteins were found highest on 4th day after treatment (DAT), whereas total carbohydrate and total chlorophyll on 2nd and 6th DAT, respectively, in comparison with the control plants. The SDS PAGE analysis revealed the induction of PR proteins, namely Chitinase (25, 29?kDa) and β-1,3-glucanase (33?kDa), in treated plants in comparison with untreated control plants. The treated plants showed enhanced growth and development as well as increase in yield. About 100% survival rate was found in maize seeds treated with AFFCF and APFCF and grown on respective fungal infested soil than control. The enhanced activities of defence enzymes and elevated protein, carbohydrate, chlorophyll content in treated maize plants suggest the induction of SAR against A. flavus and A. parasiticus by using the same fungal elicitors.     

Induced systemic resistance (ISR) against pathogens – a promising field for ecological research

Putative fitness costs provide an explanation for why ISR is induced instead of constitutive, and they might constrain the use of ISR as preventative protection of cultivated plants. Though ISR is mainly elicited by and effective against pathogens, further biotic agents such as leaf-chewing herbivores, leaf miners, aphids and even non-pathogenic root-colonising bacteria can induce systemic pathogen resistance, while some ISR traits can have a defensive effect against herbivores. ‘Cross-resistance’ elicited by and effective against non-microbial plant enemies thus might add significantly to the function of ISR. On the other hand, ‘trade-offs” have been reported, i.e. increased susceptibility to herbivores in ISR-expressing plants. Finally, ISR is a rather unspecific response, being active against different microbes. It thus might have effects on mutualistic bacteria and fungi, too. The question of how expression of ISR affects the large variety of mutualistic and antagonistic plant-microbe and plant-insect interactions cannot yet be answered. This knowledge is, however, needed to obtain a risk assessment for the use of chemically induced or genetically engineered ISR in crop protection. This review aims to provide an overview and to highlight some of the many open questions which require intensive ecological research.     

Induction of systemic resistance in tobacco against Tobacco mosaic virus by Bacillus spp.

Bacillus pumilus strain EN16 and Bacillus subtilis strain SW1 were tested for their systemic resistance and protection abilities against tobacco mosaic virus disease under greenhouse conditions. The results showed that strain EN16 and SW1 treatment significantly reduced mosaic symptoms and disease severity, resulting in 52 and 71% protection at 14 days of inoculation, respectively. A decreased amount of virus was detected in EN16- or SW1-treated tobacco plants by ELISA. Moreover, 5- and 7-day intervals between inducer treatment and pathogen inoculations were respectively required for strain EN16 and SW1 to induce optimal resistance. Further analysis on phenylalanine ammonia-lyase, peroxidase, polyphenol oxidase and pathogenesis-related (PR) proteins in tobacco showed that the amounts of defense enzymes and PR proteins significantly increased in Bacillus-treated plants challenged with pathogen when compared to control.     

茉莉酸对棉花单宁含量和抗虫相关酶活性的诱导效应

以植物生长调节物茉莉酸(Jasmonic acid,JA)为诱导子,以常规棉为研究对象,探讨了外源茉莉酸对棉花幼苗单宁和蛋白酶抑制素以及其它抗虫相关酶活性诱导的浓度依赖性和持久性,讨论了棉花抗虫相关物质的抗虫效果.结果表明,0.01、0.1和1.0 mmol/L茉莉酸都能在2周内诱导棉花单宁和胰蛋白酶抑制素(Proteinase inhibitors,PIs)含量增加,诱导多酚氧化酶(Polyphenol oxidase,PPO)、苯丙氨酸解氨酶(Phenylalanine ammonia-lyase,PAL)、过氧化物酶(Peroxidase,POD)和过氧化氢酶(Catalase,CAT)活性升高.对3种浓度茉莉酸的诱导效应进行分析表明,0.1 mmol/L茉莉酸对于诱导PIs、PPO、POD和CAT最有效,0.1和1.0 mmol/L茉莉酸对于诱导棉花单宁和苯丙氨酸解氨酶等效,二者的诱导效应均高于0.01 mmol/L.对茉莉酸诱导抗性的持久性进行分析表明,最佳诱导效应发生的时间各不相同:POD活性在JA处理后第1天最高,随后呈下降趋势,PIs和单宁含量分别在JA处理后第7天和第14天达最大值;JA处理后第1天和第7天的PPO活性无明显差异,但明显高于第14天;JA处理后第7天和第14天的PAL活性无明显差异,但明显高于第1天;JA处理后第1、7和14天棉花叶片的CAT活性均无明显差异.以上结果表明,茉莉酸可通过增加棉叶单宁和PIs含量、提高棉叶PAL、PPO、POD和CAT活性等增强棉花幼苗的抗虫性.     

Signaling of systemic acquired resistance in tobacco depends on ethylene perception

The hypersensitive interaction between Tobacco mosaic virus (TMV) and tobacco results in accumulation of salicylic acid (SA), defense gene expression, and development of systemic acquired resistance (SAR) in uninfected leaves. The plant hormones SA and ethylene have been implicated in SAR. From a study with ethylene-insensitive (Tetr) tobacco, we concluded that ethylene perception is required to generate the systemic signal molecules in TMV-infected leaves that trigger SA accumulation, defense gene expression, and SAR development in uninfected leaves. Ethylene perception was not required for the responses of the plant to the systemic signal that leads to SAR development.     

Insect eggs induce a systemic acquired resistance in Arabidopsis

Although they constitute an inert stage of the insect's life, eggs trigger plant defences that lead to egg mortality or attraction of egg parasitoids. We recently found that salicylic acid (SA) accumulates in response to oviposition by the Large White butterfly Pieris brassicae, both in local and systemic leaves, and that plants activate a response that is similar to the recognition of pathogen‐associated molecular patterns (PAMPs), which are involved in PAMP‐triggered immunity (PTI). Here we discovered that natural oviposition by P. brassicae or treatment with egg extract inhibit growth of different Pseudomonas syringae strains in Arabidopsis through the activation of a systemic acquired resistance (SAR). This egg‐induced SAR involves the metabolic SAR signal pipecolic acid, depends on ALD1 and FMO1, and is accompanied by a stronger induction of defence genes upon secondary infection. Although P. brassicae larvae showed a reduced performance when feeding on Pseudomonas syringae‐infected plants, this effect was less pronounced when infected plants had been previously oviposited. Altogether, our results indicate that egg‐induced SAR might have evolved as a strategy to prevent the detrimental effect of bacterial pathogens on feeding larvae.     

Endophytic Bacillus subtilis enriched with chitin offer induced systemic resistance in cotton against aphid infestation

Endophytic bacterial strains EPCO 102 and EPCO 16 were effectively reduced the aphid population under greenhouse conditions. Biochemical characterisation and 16S rRNA sequence analysis confirmed as Bacillus subtilis. Talc-based powder formulation for Bacillus subtilis strains EPCO 102, EPCO 16 and Pseudomonas fluorescens Pf1 (with and without chitin amendments) were tried and its effect in inducing systemic resistance were tested under greenhouse conditions. The combined application of bioformulation as seed, soil and foliar spray significantly reduced the aphid infestation. Chitin addition in the formulation showed additional reduction in the infestation by the aphids. Application of Pf1 culture along with chitin showed less aphid infestation and this efficacy was on par with the chemical insecticide treatment, followed by EPCO 16 + Chitin. In addition, these endophytic bacterial strains along with chitin induced the activity of peroxidase, polyphenol oxidase, phenylalanine ammonia-lyase, chitinase, β-1.3-glucanase and phenol accumulation in cotton, which favours reduction in aphid infestation.     

Induced systemic resistance (ISR) against pathogens in the context of induced plant defences

Induced systemic resistance (ISR) of plants against pathogens is a widespread phenomenon that has been intensively investigated with respect to the underlying signalling pathways as well as to its potential use in plant protection. Elicited by a local infection, plants respond with a salicylic-dependent signalling cascade that leads to the systemic expression of a broad spectrum and long-lasting disease resistance that is efficient against fungi, bacteria and viruses. Changes in cell wall composition, de novo production of pathogenesis-related-proteins such as chitinases and glucanases, and synthesis of phytoalexins are associated with resistance, although further defensive compounds are likely to exist but remain to be identified. In this Botanical Briefing we focus on interactions between ISR and induced resistance against herbivores that is mediated by jasmonic acid as a central signalling molecule. While many studies report cross-resistance, others have found trade-offs, i.e. inhibition of one resistance pathway by the other. Here we propose a framework that explains many of the thus far contradictory results. We regard elicitation separately from signalling and from production, i.e. the synthesis of defensive compounds. Interactions on all three levels can act independently from each other.     

Induction of systemic resistance in tomato against broomrape (Phelipanche aegyptiaca)

Rhizosphere dwelling bacteria can increase plant resistance to biotic and abiotic stresses, and they promote plant growth through various mechanisms. In this study, three bioassays were conducted including the following: (a) screening for effective bacterial isolates in the suppression of broomrape, (b) evaluating induced systemic resistance against broomrape and (c) comparing the selected bacterium isolate with plant chemical inducers. Fifteen plant growth‐promoting rhizobacteria (PGPR) were examined to assess their biocontrol potential against Egyptian broomrape (Phelipanche aegyptiaca). Ten isolates significantly reduced the broomrape biomass compared to the control. The Lysinibacillus boronitolerans B124 reduced the dry weight of broomrape plants from 2.15 g in control to 0.45 g. Bacillus megaterium B6 was the best isolate in reducing the number of broomrape tubercles. In addition, the activity of three selected bacterial isolates was investigated in induced systemic resistance to broomrape by split‐root method. The Bacillus pumilus INR7 reduced the number of visible broomrape tubercles by 90%, and B. megaterium B71 and L. boronitolerans B124 were the next two in rank. Compared with the control, L. boronitolerans B124 reduced the dry weight of broomrape from 1.49 g in control to 0.39 g. In a subsequent experiment, L. boronitolerans B124 was evaluated along with some resistance‐inducing volatile compounds. Lysinibacillus boronitolerans B124 decreased the number of broomrapes by 87% on average, while the lowest dry weight of broomrape was observed in methyl jasmonate treatment. In conclusion, PGPR have considerable potential to be used in the integrated management of broomrape. It is also possible to use a mixture of rhizobacteria and defence inducers, such as biogenic volatiles as a promising approach in the management of this noxious parasitic weed.     

Variation in the response of tomato (Solanum lycopersicum) breeding lines to the effects of benzo (1,2,3) thiadiazole‐7‐carbothioic acid S‐methyl ester (BTH) on systemic acquired resistance and seed germination

Genetic variation may play a major role in how plants respond to activators of systemic acquired resistance. To examine this, the defence activator benzo(1,2,3)thiadiazole‐7‐carbothioic acid S‐methyl ester (BTH) was applied to seed of different breeding lines of tomato (Solanum lycopersicum) with diverse pedigrees, and the levels of induced resistance against Pseudomonas syringae pv. tomato, changes in defence gene expression and detrimental effects on seed germination and seedling emergence were measured. Two breeding lines, 7007 and 7024, were selected as non‐responsive and responsive to BTH. The SAR‐associated genes, SlPR1a and SlPR3b, were induced earlier or more strongly over the control prior to inoculation for line 7024 but not for line 7007. This was not observed for the ISR‐related genes, SlPin2 and SlPR2b. BTH inhibition of seed germination and seedling emergence was more delayed in line 7024 than 7007. However, applying BTH as a seed or soil drip reduced the delay. Thus, greater levels of BTH response have both positive (i.e., induced resistance and expression of SAR‐related gene expression) and negative (i.e., inhibition of seed germination and seedling emergence) effects and can differ significantly between genotypes. Thus, recommendations for use of induced resistance activators should include plant genotype recommendations and consider possible negative impacts of greater responsiveness.