Abstract
Beneficial plant-microbe interactions have utmost importance for enhancing plant growth, improving soil structure, and managing plant diseases. Not surprisingly, such mutual interactions, where plants provide nourishment to rhizospheric microbes and in return microbes help in facilitating plant growth and stress amelioration, actually lay the foundation of sustainable agriculture. To cope with the major challenge of pathogen attack, beneficial rhizospheric microbes have proven their efficacy by induced systemic resistance (ISR). Therefore, such microbes are increasingly used in the form of biofertilizers and biopesticides. Moreover, such plant-microbe interactions elicit a range of defense-responsive activities in order to combat the pathogen challenge. The main microbes-mediated defense strategies adopted by plants include activation of antioxidant status of the plant by reprogramming defense-related enzymes, modulation of quorum sensing phenomenon, and activation of phenylpropanoid pathway leading to phenolics production, lignin deposition, and transgenerational defense response. In this chapter, we highlight the relevance of beneficial interactions between plant and microbes in enhancing plants’ innate immune system against pathogen attack. This review provides a better understanding of the recent advances and major outcome of positive plant-microbe interactions and linking their relevance to plant defense response.
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References
Alvarez MV, Moreira MR, Ponce A (2012) Antiquorum sensing and antimicrobial activity of natural agents with potential use in food. J Food Saf 32:379–387
Asada K (1999) The water–water cycle in chloroplasts: scavenging of active oxygen and dissipation of excess photons. Annu Rev Plant Physiol Plant Mol Biol 50:601–639
Asada K, Takahashi M (1987) Production and scavenging of active oxygen in photosynthesis. In: Kyle DJ, Osmond CB, Arntzen CJ (eds) Photoinhibition. Elsevier, Amsterdam, pp 227–287
Bais HP, Weir TL, Perry LG, Gilroy S, Vivanco JM (2006) The role of root exudates in rhizosphere interactions with plants and other organisms. Annu Rev Plant Biol 57:234–266
Bakker PAHM, Pieterse CMJ, Van Loon LC (2007) Induced systemic resistance by fluorescent pseudomonas spp. Phytopathology 97:239–243
Balakrishnan N, Subramanian KS (2012) Mycorrhizal symbiosis and bioavailability of micronutrients in maize grain. Maydica 57:129–138
Banerjee M, Yesmin L (2002) Sulfur-oxidizing plant growth promoting rhizobacteria for enhanced canola performance. US Patent 07491535
Bashan Y, Holguin G (1998) Proposal for the division of plant growth-promoting rhizobacteria into two classifications: biocontrol-PGPB (plant growth-promoting bacteria) and PGPB. Soil Biol Biochem 30:1225–1228
Berendsen RL, Pieterse CMJ, Bakker PAHM (2012) The rhizosphere microbiome and plant health. Trends Plant Sci 17:478–486
Berg G (2009) Plant–microbe interactions promoting plant growth and health: perspectives for controlled use of microorganisms in agriculture. Appl Microbiol Biotechnol 84:11–18
Berg G, Smalla K (2009) Plant species and soil type cooperatively shape the structure and function of microbial communities in the rhizosphere. FEMS Microbiol Ecol 68:1–13
Bernards MA, Lewis NG (1998) The macromolecular aromatic domain in suberized tissue: a changing paradigm. Phytochemistry 47:915–933
Boerjan W, Ralph J, Baucher M (2003) Lignin biosynthesis. Annu Rev Plant Biol 54:519–546
Bowler C, Slooten L, Vandenbranden S, De Rycke R, Botterman J, Sybesma C, Van Montagu M, Inzé D (1991) Manganese superoxide dismutase can reduce cellular damage mediated by oxygen radicals in transgenic plants. EMBO J 10:1723–1732
Cassán FD, GarcĂa de Salamone I (2008) Azospirillum sp.: cell physiology, plant interactions and agronomic research in Argentina. AsociaciĂłn Argentina de MicrobiologĂa, Argentina, p 266
Cazale AC, Droillard MJ, Wilson C, Heberle-Bors E, Barbier-Brygoo H, Laurière C (1999) MAP kinase activation by hypo-osmotic stress of tobacco cell suspensions: towards the oxidative burst response? Plant J 19:297–307
Clark RB, Zobel RW, Zeto SK (1999) Effects of mycorrhizal fungus isolates on mineral acquisition by Panicum virgatum acidic soils. Mycorrhiza 9:167–176
Conrath U (2011) Molecular aspects of defence priming. Trends Plant Sci 16(10):524–531
Contreras-Cornejo HA, Macias-Rodriguez L, Cortes-Penagos C, Lopez-Bucio J (2009) Trichoderma virens, a plant beneficial fungus, enhances biomass production and promotes lateral root growth through an auxin-dependent mechanism in Arabidopsis. Plant Physiol 149:1579–1592
Crépin A, Barbey C, Beury-Cirou A, Hélias V, Taupin L, Reverchon S, Nasser W, Faure D, Dufour A, Orange N, Feuilloley M, Heurlier K, Burini JF, Latour X (2012a) Quorum sensing signaling molecules produced by reference and emerging soft-rot bacteria (Dickeya and Pectobacterium spp.). PLoS One 7(4):e35176
Crépin A, Barbey C, Cirou A, Tanniéres M, Orange N, Orange N, Feuilloley M, Dessaux Y, Burini JF, Faure D, Latour X (2012b) Biological control of pathogen communication in the rhizosphere: a novel approach applied to potato soft rot due to Pectobacterium atrosepticum. Plant Soil 358:27–37
Dat J, Vandenabeele S, Vranová E, Van Montagu M, Inzé D, Van Breusegem F (2000) Dual action of the active oxygen species during plant stress responses. Cell Mol Life Sci 57:779–795
Davin LB, Lewis NG (2000) Dirigent proteins and dirigent sites explain the mystery of specificity of radical precursor coupling in lignan and lignin biosynthesis. Plant Physiol 123:453–462
De Vleeschauwer D, Höfte M (2007) Using Serratia plymuthica to control fungal pathogens of plant. CAB Rev 2:46
De Werra P, Péchy-Tarr M, Keel C, Maurhofer M (2009) Role of gluconic acid production in the regulation of biocontrol traits of Pseudomonas fluorescens CHA0. Appl Environ Microbiol 75:4162–4174
Desikan R, A-HMackerness S, Hancock JT, Neill SJ (2001) Regulation of the Arabidopsis transcriptosome by oxidative stress. Plant Physiol 127:159–172
Diallo S, Crépin A, Barbey C, Orange N, Burini JF, Latour X (2011) Mechanisms and recent advances in biological control mediated through the potato rhizosphere. FEMS Microbiol Ecol 75:351–364
Dicke M, Hilker M (2003) Induced plant defences: from molecular biology to evolutionary ecology. Basic Appl Ecol 4:3–14
Dixon RA, Paiva N (1995) Stress induced phenylpropanoid metabolism. Plant Cell 7:1085–1097
Djonovic S, Pozo MJ, Dangott LJ, Howell CR, Kenerley CM (2006) Sm1, a proteinaceous elicitor secreted by the biocontrol fungus Trichoderma virens induces plant defense responses and systemic resistance. Mol Plant-Microbe Interact 8:838–853
Djonovic S, Vargas WA, Kolomiets MV, Horndeski M, Wiest A, Kenerley CM (2007) A proteinaceous elicitor Sm1 from the beneficial fungus Trichoderma virens is required for induced systemic resistance in maize. Plant Physiol 145:875–889
Dobbelare S, Vanderleydern J, Okon Y (2003) Plant-growth promoting effects of diazotrophs in the rhizosphere. Crit Rev Plant Sci 22:107–149
Dong YH, Wang LH, Xu JL, Zhang HB, Zhang XF, Zhang LH (2001) Quenching quorum-sensing-dependent bacterial infection by an N-acyl homoserine lactonase. Nature 411:813–817
Durrant WE, Dong X (2004) Systemic acquired resistance. Annu Rev Phytopathol 42:185–209
Friend J (1976) Lignification in infected tissue. In: Friend J, Threfall DR (eds) Biochemical aspects of plantparasite relationships. Academic, London, pp 291–303
Glick BR (2005) Modulation of plant ethylene levels by the bacterial enzyme ACC deaminase. FEMS Microbiol Lett 252:1–7
Gottlieb S, Pelczar MJ (1951) Microbiological aspects of lignin degradation. Bacteriol Rev 15:55–76
Gram L, Grossart H, Schlingloff A, Kiørboe T (2002) Possible quorum sensing in marine snow bacteria: production of acylated homoserine lactones by roseobacter strains isolated from marine snow. Appl Environ Microbiol 8(68):4111–4116
Grant JJ, Loake GJ (2000) Role of reactive oxygen intermediates and cognate redox signaling in disease resistance. Plant Physiol 124:21–29
Harman G, Shoresh M (2007) The mechanisms and applications of opportunistic plant symbionts. In: Vurro M, Gressel J (eds) Novel biotechnologies for biocontrol agent enhancement and management. Springer, Amsterdam, pp 131–155
Harman GE, Howell CR, Viterbo A, Chet I, Lorito M (2004) Trichoderma species – opportunistic, avirulent plant symbionts. Nat Rev Microbiol 2:43–56
Harrison MJ (1999) Molecular and cellular aspects of the arbuscular mycorrhizal symbiosis. Annu Rev Plant Physiol 50:361–389
Harrison MJ (2005) Signaling in the arbuscular mycorrhizal symbiosis. Annu Rev Microbiol 59:19–42
Hatfield R, Vermerris W (2001) Lignin formation in plants. The dilemma of linkage specificity. Plant Physiol 126:1351–1357
Hayatsu M, Tago K, Saito M (2008) Various players in the nitrogen cycle: diversity and functions of the microorganisms involved in nitrification and denitrification. Soil Sci Plant Nutr 54:33–45
Howell CR, Hanson LE, Stipanovic RD, Puckhaber LS (2000) Induction of terpenoid synthesis in cotton roots and control of Rhizoctonia solani by seed treatment with Trichoderma virens. Phytopathology 90:248–252
Hummerschmidt R (1999) Phytoalexins: what have we learned after 60 years? Annu Rev Phytopathol 37:285–306
Jain A, Singh S, Sarma BK, Singh HB (2012) Microbial consortium mediated reprogramming of defence network in pea to enhance tolerance against Sclerotinia sclerotiorum. J Appl Microbiol 112:537–550
Jetiyanon K (2007) Defensive-related enzyme response in plants treated with a mixture of Bacillus strains (IN937a and IN937b) against different pathogens. Biol Control 42:178–185
Jeun YC, Park KS, Kim CH, Fowler WD, Kloepper JW (2004) Cytological observations of cucumber plants during induced resistance elicited by rhizobacteria. Biol Control 29:34–42
Jones JDG, Dangl JL (2006) The plant immune system. Nature 444:323–329
Kamilova F, Validov S, Azarova T, Mulders I, Lugtenberg B (2005) Enrichment for enhanced competitive plant root tip colonizers selects for a new class of biocontrol bacteria. Environ Microbiol 7:1809–1817
Kamilova F, Kravchenko LV, Shaposhnikov AI, Makarova N, Lugtenberg BJJ (2006) Effects of the tomato pathogen Fusarium oxysporum f. sp. radicis-lycopersici and of the biocontrol bacterium Pseudomonas fluorescens WCS365on the composition of organic acids and sugars in tomato root exudate. Mol Plant-Microbe Interact 19:1121–1126
Keswani C, Mishra S, Sarma BK, Singh SP, Singh HB (2014) Unraveling the efficient applications of secondary metabolites of various Trichoderma spp. Appl Microbiol Biotechnol 98:533–544
Kim JM, To TK, Seki M (2012) An epigenetic integrator: new insights into genome regulation, environmental stress responses and developmental controls by histone deacetylase 6. Plant Cell Physiol 53(5):794–800
Kloepper JW, Leong J, Teintze M, Schroth MN (1980) Enhanced plant growth by siderophores produced by plant growth-promoting rhizobacteria. Nature 286:885–886
Kloepper JW, Ryu CM, Zhang SA (2004) Induced systemic resistance and promotion of plant growth by Bacillus spp. Phytopathology 94:1259–1266
Lambert DH, Cole H, Baker DE (1980) The role of boron in plant response to mycorrhizal infection. Plant Soil 57:431–438
Lavania M, Chauhan PS, Chauhan SVS, Singh HB, Nautiyal CS (2006) Induction of plant defense enzymes and phenolics by treatments with plant growth promoting rhizobacteria Serratia marcescens NBRI 1213. Curr Microbiol 52:363–368
Lewis NG, Yamamoto E (1990) Lignin: occurrence, biogenesis and biodegradation. Annu Rev Plant Physiol Plant Mol Biol 41:455–496
Liu A, Hamel C, Hamilton RI, Ma BL, Smith DL (2000) Acquisition of Cu, Zn, Mn and Fe by mycorrhizal maize (Zea mays L.) growth in soil at different P and micronutrient levels. Mycorrhiza 9:331–336
Lugtenberg BJJ, Chin-A-Woeng TFC, Bloemberg GV (2002) Microbe– plant interactions: principles and mechanisms. Antonie Van Leeuwenhoek 81:373–383
Luna E, Ton J (2012) The epigenetic machinery controlling transgenerational systemic acquired resistance. Plant Signal Behav 7:615–618
Luna E, Bruce TJA, Roberts MR, Flors V, Ton J (2012) Next-generation systemic acquired resistance. Plant Physiol 158:844–853
Mäe A, Montesano M, Koiv V, Palva ET (2001) Transgenic plants producing the bacterial pheromone N-acyl-homoserine lactone exhibit enhanced resistance to the bacterial phytopathogen Erwinia carotovora. Mol Plant-Microbe Interact 14:1035–1042
Mandal S, Mitra A (2007) Reinforcement of cell wall in roots of Lycopersicon esculentum through induction of phenolic compounds and lignin by elicitors. Physiol Mol Plant Pathol 71:201–209
Meziane H, Van der Sluis I, Van Loon LC, Ho¨fte M, Bakker PAHM (2005) Determinants of Pseudomonas putida WCS358 involved in inducing systemic resistance in plants. Mol Plant Pathol 6:177–185
Mittler R (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci 7:405–410
Molina L, Constantinescu F, Michel L, Reimmann C, Duffy B, Defago G (2003) Degradation of pathogen quorum-sensing molecules by soil bacteria: a preventive and curative biological control mechanism. FEMS Microbiol Ecol 45:71–81
Morrissey JP, Dow JM, Mark L, O’Gara F (2004) Are microbes at the root of a solution to world food production? EMBO Rep 5:922–926
Nicholson RL, Hammerschmidt R (1992) Phenolic compounds and their role in disease resistance. Annu Rev Phytopathol 30:369–389
Ongena M, Jourdan E, Scha¨ fer M, Kech C, Budzikiewicz H, Luxen A, Thonart P (2005) Isolation of an N-alkylated benzylamine derivative from Pseudomonas putida BTP1 as elicitor of induced systemic resistance in bean. Mol Plant Microbe Interact 18:562–569
Ongena M, Jourdan E, Adam A, Paquot M, Brans A, Joris B, Arpigny JL, Thonart P (2007) Surfactin and fengycin lipopeptides of Bacillus subtilis as elicitors of induced systemic resistance in plants. Environ Microbiol 9:1084–1090
Park KS, Kloepper JW (2000) Activation of PR-1a promoter by rhizobacteria which induce systemic resistance in tobacco against Pseudomonas syringae pv. tabaci. Biol Control 18:2–9
Pei ZM, Murata Y, Benning G, Thomine S, Klüsener B, Allen GJ, Grill E, Schroeder JI (2000) Calcium channels activated by hydrogen peroxide mediate abscisic acid signaling in guard cells. Nature 406:731–734
Pierson EA, Wood D, Cannon JAW, Blachere FM, Pierson LS (1998a) Interpopulation signaling via N-acyl-homoserine lactones among bacteria in the wheat rhizosphere. Mol Plant-Microbe Interact 11:1078–1084
Pierson LS, Wood DW, Pierson EA (1998b) Homoserine lactone-mediated gene regulation in plant-associated bacteria. Annu Rev Phytopathol 36:207–225
Pieterse CMJ (2012) Prime time for transgenerational defense. Plant Physiol 158:545
Pieterse CM, Dicke M (2007) Plant interactions with microbes and insects: from molecular mechanisms to ecology. Trends Plant Sci 12:564–569
Pieterse CMJ, van der Does D, Zamioudis C, Leon-Reyes A, van Wees SCM (2012) Hormonal modulation of plant immunity. Annu Rev Cell Dev Biol 28:489–521
Pineda A, Zheng SJ, van Loon JJA, Pieterse CMJ, Dicke M (2010) Helping plants to deal with insects: the role of beneficial soil-borne microbes. Trends Plant Sci 15:507–514
Pozo MJ, Azcon-Aguilar C (2007) Unraveling mycorrhiza-induced resistance. Curr Opin Plant Biol 10:393–398
Ran LX, Li ZN, Wu GJ, Van Loon LC, Bakker PAHM (2005) Induction of systemic resistance against bacterial wilt in Eucalyptus urophylla by fluorescent Pseudomonas spp. Eur J Plant Pathol 113:59–70
Ride JP (1978) The role of cell wall alterations in resistance to fungi. Ann Appl Biol 89:302–306
Ryan RP, Monchy S, Cardinale M, Taghavi S, Crossman L, Avison MB, Berg G, van der Lelie D, Dow JM (2009) Versatility and adaptation of bacteria from the genus Stenotrophomonas. Nat Microbiol Rev 7:514–525
Ryu CM, Farag MA, Hu CH, Reddy MS, Wie HX, Paré PW, Kloepper JW (2003) Bacterial volatiles promote growth of Arabidopsis. Proc Natl Acad Sci 100:4927–4932
Ryu CM, Farag MA, Hu CH, Reddy MS, Kloepper JW, Pare’ PW (2004) Bacterial volatiles induce systemic resistance in Arabidopsis. Plant Physiol 134:1017–1026
Saleem M, Arshad M, Hussain S, Bhatti AS (2007) Perspective of plant growth promoting rhizobacteria (PGPR) containing ACC deaminase in stress agriculture. J Ind Microbiol Biotechnol 34:635–648
Schrey SD, Tarkka MT (2008) Friends and foes: Streptomycetes as modulators of plant disease and symbiosis. Antonie Van Leeuwenhoek 94:11–19
Schuhegger R, Ihring A, Gantner S, Bahnweg G, Knappe C, Hartmann A, Langebartels C (2006) Induction of systemic resistance in tomato by N-acyl-l-homoserine lactone-producing rhizosphere bacteria. Plant Cell Environ 29:909–918
Sederoff RR, MacKay JJ, Ralph J, Hatfield RD (1999) Unexpected variation in lignin. Curr Opin Plant Biol 2:145–152
Serfling A, Wirsel SGR, Lind V, Deising HB (2007) Performance of the biocontrol fungus Piriformospora indica on wheat under greenhouse and field conditions. Phytopathology 97:523–531
Shoresh M, Harman GE, Mastouri F (2010) Induced systemic resistance and plant responses to fungal biocontrol agents. Annu Rev Phytopathol 48:21–43
Silva HSA, Romeiro RDS, Macagnan D, Halfeld-Vieira BDA, Pereira MCB, Mounteer A (2004) Rhizobacterial induction of systemic resistance in tomato plants non-specific protection and increase in enzyme activities. Biol Control 29:288–295
Singh A, Sarma BK, Upadhyay RS, Singh HB (2013) Compatible rhizosphere microbes mediated alleviation of biotic stress in chickpea through enhanced antioxidant and phenylpropanoid activities. Microbiol Res 168:33–40
Singhai PK, Sarma BK, Srivastava JS (2011) Biological management of common scab of potato through Pseudomonas species and vermicompost. Biol Control 57:150–157
Slaughter A, Daniel X, Flors V, Luna E, Hohn B, Mauch-Mani B (2012) Descendants of primed Arabidopsis plants exhibit resistance to biotic stress. Plant Physiol 158:835–843
Smith SE, Jakobsen I, Gronlund M, Smith FA (2011) Roles of arbuscular mycorrhizas in plant phosphorus nutrition: Interactions between pathways of phosphorus uptake in arbuscular mycorrhizal roots have important implications for understanding and manipulating plant phosphorus acquisition. Plant Physiol 156:1050–1057
Spaink HP (2000) Root nodulation and infection factors produced by rhizobial bacteria. Annu Rev Microbiol 54:257–288
Stein E, Molitor A, Kogel KH, Waller F (2008) Systemic resistance in Arabidopsis conferred by the mycorrhizal fungus Piriformospora indica requires jasmonic acid signaling and the cytoplasmic function of NPR1. Plant Cell Physiol 49:1747–1751
Tran H, Ficke A, Asiimwe T, Ho¨fte M, Raaijmakers JM (2007) Role of the cyclic lipopeptide massetolide A in biological control of Phytophthora infestans and in colonization of tomato plants by Pseudomonas fluorescens. New Phytol 175:731–742
Truchado P, Tomás-Barberán F, Larrosa M, Allende A (2012) Food phytochemicals act as quorum sensing inhibitors reducing production and/or degrading autoinducers of Yersinia enterocolĂtica and Erwinia carotovora. Food Control 24:78–85
Unno Y, Okubo K, Wasaki J, Shinano T, Osaki M (2005) Plant growth promotion abilities and microscale bacterial dynamics in the rhizosphere of lupin analysed by phytate utilization ability. Environ Microbiol 7:396–404
Van der Putten WH, Vet LM, Harvey JA, Wäckers FL (2001) Linking above- and belowground multitrophic interactions of plants, herbivores, pathogens, and their antagonists. Trends Ecol Evol 16:547–554
Van Loon LC, Bakker PAHM, Pieterse CMJ (1998) Systemic resistance induced by rhizosphere bacteria. Annu Rev Phytopathol 36:453–483
Van Oosten VR, Bodenhausen N, Reymond P, Van Pelt JA, Van Loon LC, Dicke M, Pieterse CMJ (2008) Differential effectiveness of microbially induced resistance against herbivorous insects in Arabidopsis. Mol Plant-Microbe Interact 21:919–930
van Rhijn P, Vanderleyden J (1995) The Rhizobium-plant symbiosis. Microbiol Rev 59:124–142
van Wees SCM, de Swart EAM, van Pelt JA, van Loon LC, Pieterse CMJ (2000) Enhancement of induced disease resistance by simultaneous activation of salicylate – and jasmonate-dependent defense pathways in Arabidopsis thaliana. Proc Natl Acad Sci U S A 97:8711–8716
Vance CP, Anderson JO, Sherwood RT (1976) Soluble and cell wall peroxidases in reed canary grass in relation to disease resistance and localized lignin formation. Plant Physiol 57:920–922
Vinale F, Sivasithamparam K, Ghisalberti EL, Marra R, Barbetti MJ, Li H, Woo SL, Lorito M (2008) A novel role for Trichoderma secondary metabolites in the interactions with plants. Physiol Mol Plant Pathol 72:80–86
Whitmore FW (1978) Lignin-carbohydrate complex formed in isolated cell walls of callus. Phytochemistry 17:421–425
Woo SL, Scala F, Ruocco M, Lorito M (2006) The molecular biology of the interactions between Trichoderma spp., phytopathogenic fungi and plants. Phytopathology 96:181–185
Yates IE, Bacon CW, Hinton DM (1997) Effects of endophytic infection by Fusarium moniliforme on corn growth and cellular morphology. Plant Dis 81:723–728
Zhang H, Xie X, Kim MS, Kornyeyev DA, Holaday S, Par’e PW (2008) Soil bacteria augment Arabidopsis photosynthesis by decreasing glucose sensing and abscisic acid levels in planta. Plant J 56:264–273
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HBS and BKS are grateful to the Department of Biotechnology, Govt. of India, for providing financial support (BT/PR5990/AGR/5/587/2012). SM is thankful to UGC for awarding Dr. D.S. Kothari Postdoctoral Fellowship.
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Mishra, S., Singh, A., Keswani, C., Saxena, A., Sarma, B.K., Singh, H.B. (2015). Harnessing Plant-Microbe Interactions for Enhanced Protection Against Phytopathogens. In: Arora, N. (eds) Plant Microbes Symbiosis: Applied Facets. Springer, New Delhi. https://doi.org/10.1007/978-81-322-2068-8_5
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