ELICITORS ACTIVATE DEFENSE MECHANISM

Soil is the key determinant for soil microbial population and their activities. Plant residue or manure, effect soil microorganisms and soil suppressiveness by promoting beneficial soil microorganisms. Several strategies have been developed to control plant disease such as soil management,  use of resistant varieties,  crop rotation,  removal of infected plants, use of plant protection product and besides all these one method is to activate the defense mechanism that is inherent to plants when plants come into contact with pathogenic or non-pathogenic microorganisms. Defense reaction occur so that the infection does not spread in plant. Presence of pathogen is detected through the microbial signal.

Extracellular signal compound involved in activation of defense reaction is known as elicitors. Elicitors are of microbial origin or are from other biotic sources. Elicitor can induce defense responses even at a very low concentration (nM) (Shibuya and Minami 2001).  Fungi are rich source of elicitors (Eder and Cosio 1994). The elicitors in fungi are glucans, chitin and chitosan oligosaccharides as well as glycopeptides, glycoproteins, and ergosterol (Boller 1995). Plants can recognize elicitors such as:

  1. Flagellin (Felix et al., 1999) and harpin from bacteria (Dong et al., 1999)
  2. Chitin (Egusa et al., 2015; Felix et al., 1993), ergosterol (Granado et al., 1995) and cell wall glucans (Sharp et al., 1984; Yamaguchi et al., 2000) from fungi
  3. Laminarins from algae (Aziz et al., 2003)

Three categories of elicitors have been determined so far:

  • Pathogen-associated molecular patterns (PAMPs): Released specifically from pathogen.
  • Microbe-associated molecular patterns (MAMPs): Released by beneficial/non-pathogenic microorganisms such as yeasts, and plant growth-promoting rhizobacteria or plant growth-promoting fungi.
  • Damage- or danger-associated molecular patterns (DAMPs): Emitted from the plant itself (Henry et al., 2012). Endogenous molecular patterns is related to injured plant tissue.

Elicitors act by activating pattern recognition receptors (PRRs). Resistance inducing PAMPs may be provided to plants by beneficial microorganisms. Elicitors of induced systemic resistance (ISR) produced by plant growth promoting rhizobacteria (PGPR) include siderophores such as pseudobactin (Meziane et al., 2005).

Elicitor molecule bind with the specific plant receptor located on plant cell membranes. These receptors are able to recognize the molecular pattern of elicitors activating the signal transduction pathway leading to production of phytoalexin, PR protein, reactive oxygen species (ROS) and strengthening cell wall which provides resistance against pathogen or environmental stress. Many plant genes that respond to environmental and developmental changes are regulated by jasmonic acid derived from linolenic acid via octadecanoid pathway (Reinbothe et al., 1994). Jasmonic acid (JA) and its derivatives are part of a general signal transduction system. Methyl jasmonate (MeJA) being volatile is released in atmosphere and can induce defense response in neighbouring plants (Reinbothe et al., 1994). Jasmonates induce the synthesis of proteinase inhibitor proteins in response to wounding and pathogen attack (Farmer and Ryan 1990, 1992) and trigger the formation of phytoalexins and their biosynthetic enzymes (Gundlach et al., 1992) and thionins, polypeptides with antifungal activity, in barley (Andresen et al., 1992).

Elicitors of plant defense responses are exogenous and endogenous:

  • Exogenous elicitors: Elicitors which are the primary signal in pathogen interaction. Proteins, oligosaccharides, glycoprotein, fatty acids and their derivatives can function as exogenous elicitors.
  • Endogenous elicitors: Elicitors which are component of intercellular signal transduction system of plants (Eder and Cosio 1994).

Plant defense systems are activated in response to chitin in fungal (pathogen) cell walls, which is perceived as a microbe- or pathogen-associated molecular pattern (MAMP/PAMP) (Egusa et al., 2015). Chitin elicitor triggers disease resistance in both dicot and monocot plants (Shibuya and Minami 2001).

In soil chitin comes from insect (exoskeleton) and fungi (constituent of the cell wall). Chitin is a major component of fungal cell walls and serves as a microbe-associated molecular pattern (MAMP) for the detection of various potential pathogens of plants. Plant possess chitin degrading enzyme to digest fungal cell wall during infection as well as plant cell membranes contain chitin-specific receptors, which are known to activate induced defense mechanisms.  Chitin recognition results in activation of defense signaling pathway.

A range of “chitin elicitor binding proteins” (CEBiP) have been isolated from a number of crops (Miya et al., 2007) and all these glycoproteins possess a highly conserved extracellular lysine motif (LysM) that binds chitin directly when in contact with the plasma membrane in which it is embedded. Chitin and chitosan induce host defense responses in both monocot and dicot plants. Application of chitin, chitosan and glucosamine on crop plants show a range of beneficial responses (Sharp 2013) such as improve crop yield, toxic to plant pest and pathogen, enhance beneficial microorganisms and induce plant defense response. Elicitor substances have been widely used in plant disease control which has least impact on the environment.

 References:

Andresen, I., Becher, W., Schluter, K., Parthier, B. and Apel, K. 1992 The Identification of Leaf Thionin as One of the Main Jasmonate-Induced Proteins of Barley (Hordeum vulgare) Plant Mol. Biol. 19: 193-204

Aziz, A., Poinssot, B., Daire, X., Adrian, M., Bézier, A., Lambert, B., Joubert, J. M. and Pugin, A. 2003 Laminarin Elicits Defense Responses in Grapevine and Induces Protection Against Botrytis cinerea and Plasmopara viticola. Mol. Plant Microbe Interact. 16:  1118-1128

Boller, T. 1995 Chemoperception of Microbial Signals in Plant Cells. Annu. Rev. Plant Physiol. Plant Mol. Biol. 46: 189 214

Dong, H., Delaney, T.P., Bauer, D.W. and Beer, S.V. 1999 Harpin Induces Disease Resistance in Arabidopsis Through the Systemic Acquired Resistance Pathway Mediated by Salicylic Acid and the NIM1 Gene. Plant J. 20:  207-215

Eder, J. and Cosio, E. G. 1994 Elicitors of Plant Defense Responses. International Review of Cytology 148:  1-36

doi.org/10.1016/S0074-7696 (08)62404-3

Egusa, M., Matsui, H., Urakami, T., Okuda, S., Ifuku, S., Nakagami, H. and Kaminaka, H. 2015 Chitin Nanofiber Elucidates the Elicitor Activity of Polymeric Chitin in Plants. Front Plant Sci. 6: 1098

doi:  10.3389/fpls.2015.01098

Farmer, E. E. and Ryan, C. A. 1992 Octadecanoid Precursors of Jasmonic Acid Activate the Synthesis of Wound-Inducible Proteinase Inhibitors.  Plant Cell 4(2): 129-134

Farmer, E. E. and Ryan, C. A.  1990 Interplant Communication: Airborne Methyl Jasmonate Induces Synthesis of Proteinase Inhibitors in Plant Leaves.  Proc. Natl. Acad. Sci. USA 87:  7713-7716

Felix, G., Duran, J.D., Volko, S. and Boller, T. 1999 Plants Have a Sensitive Perception System for the Most Conserved Domain of Bacterial Flagellin. The Plant J. 18(3): 265–276

doi: 10.1046/j.1365-313X.1999.00265.x

Felix, G., Regenass, M. and Boller, T. 1993 Specific Perception of Subnanomolar Concentrations of Chitin Fragments by Tomato Cells: Induction of Extracellular Alkalinization, Changes in Protein Phosphorylation, and Establishment of a Refractory State. The Plant J. 4(2):  307-316

doi: 10.1046/j.1365-313X.1993.04020307.x

Granado, J., Felix, G. and Boller, T. 1995 Perception of Fungal Sterols in Plants. Plant Physiol. 107:  485-490

Gundlach, H., Miller, M. J., Kutchan, T. M. and  Zenk, M. H. 1992 Jasmonic Acid is a Signal Transducer in Elicitor-Induced Plant Cell Cultures Proc. Natl. Acad. Sci. USA 89(6):  2389-2393

Henry, G., Thonart, P. and Ongena, M. 2012 PAMPs, MAMPs, DAMPs and Others: an Update on the Diversity of Plant Immunity Elicitors Base 16: 1-9

Meziane, H., Van der Sluis I., Van Loon, L. C., Hofte, M. and Bakker P. A. H. M. 2005 Determinants of Pseudomonas putida WCS358 Involved in Inducing Systemic Resistance in Plants. Mol. Plant Pathol. 6: 177 – 185

Miya, A., Albert, P., Shinya, T., Desaki, Y., Ichimura, K., Shirasu, K., Narusaka, Y., Kawakami, N., Kaku, H. and  Shibuya, N. 2007 CERK1, a LysM Receptor Kinase, is Essential for Chitin Elicitor Signaling in Arabidopsis. Proc. Natl. Acad. Sci. USA 104(49): 19613–19618

Reinbothe, S., Mollenhauer, B. and Reinbothe, C. 1994 JIPs and RIPs: The Regulation of Plant Gene Expression by Jasmonates in Response to Environmental Cues and Pathogens.  Plant Cell 6(9):  1197−1209

doi: 10.1105/tpc.6.9.1197

Sharp, R. G.  2013 A Review of the Applications of Chitin and Its Derivatives in Agriculture to Modify Plant-Microbial Interactions and Improve Crop Yields.  Agronomy 3(4): 757-793

doi:10.3390/agronomy3040757

Sharp, J. K., Valent, B. and Albersheim, P. 1984 Purification and Partial Characterization of a β-Glucan Fragment That Elicits Phytoalexin Accumulation in Soybean. J. Biol. Chem. 259(18): 11312-11320

Shibuya, N. and Minami, E. 2001 Oligosaccharide Signaling for Defense Responses in Plant. Physiol. Mol. Plant Pathol. 59 223–233

10.1006/pmpp.2001.0364

Yamaguchi, T., Yamada, A., Hong, N., Ogawa, T., Ishii, T. and Shibuya, N.  2000 Differences in the Recognition of Glucan Elicitor Signals Between Rice and Soybean: β-Glucan Fragments from the Rice Blast Disease Fungus Pyricularia Oryzae That Elicit Phytoalexin Biosynthesis in Suspension-Cultured Rice Cells. Plant Cell 12: 817-826

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