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R-Avr Genes Interactions and Plant Defense SystemBY: Ruchi V. Trivedi | Category: Agriculture | Submitted: 2016-10-17 00:29:32
Article Summary: "Host plant having Resistance gene and corresponding gene in the pathogen is Avirulence gene, their interaction play curcial role in plant defense system. .."
R-Avr Genes Interactions and Plant Defense System
Authors: Ruchi V. Trivedi and Bhupendra Singh Panwar
The resistance of the host plant is controlled by single R (resistance) gene (usually dominant) .The corresponding gene in the pathogen is called an Avr (avirulence) gene (also usually dominant). It is important to note that the actions of the R genes and the Avr genes can only be detected by test inoculations of pathogens on plants (i.e. using infection assays for resistance or susceptibility).
Plant resistance proteins (R proteins) recognize corresponding pathogen avirulence (Avr) proteins either indirectly through detection of changes in their host protein targets or through direct R-Avr protein interactions. Although indirect recognition imposes selection against Avr effector function, pathogen effector molecules recognized through direct interaction may overcome resistance through sequence diversification rather than loss of function.
There are four classes of R predicted protein .
⋆ NBS/LRR class (Nulceotide binding site /leucine rich repeat)
⋆ LRR/TM class (Leucine rich repeat/ transmembrane receptor)
⋆ Kinase class
⋆ LRR/TM/Kinase class.
Based on the predicted protein structures the cloned resistance genes can be grouped in to five classes.
• Hm1 (This gene is distinct from the interactions which involve in R genes that may couple the recognition of specific pathogen races to expression of defence related genes.)
• Pto (Pseudomonas tomato resistance)
• RPM1(Resistance to Pseudomonas syringae ssp. maculicola 1)
• Cf9 (Resistance to Cladosporium fulvum-9)
• Xa 21 (Xanthomonas campestris resistance 21)
Mechanism of R-Avr interaction
Direct recognition ( ‘Gene-for-Gene’ hypothesis ).
The ‘Gene-for-Gene’ hypothesis proposed by Flor (1971) suggests that for each avirulence gene product synthesized by the pathogen, the resistant host carries a complementary, single, dominant R gene whose product recognizes the Avr product. During infection, an interaction between these two components induces a defence response.
1) Flax resistance genes - flax rust avirulence genes (Peter N. Dodds et al., 2006)
The flax rust fungus Avr L567 genes, whose products are recognised by the L5, L6, and L7 R proteins of flax, are highly diverse, with twelve sequence variants identified from six rust strains. Out of twelve seven show necrotic response within corresponding R gene. Yeast Two- Hybrid assay indicates that recognition is based on direct R- Avr protein interaction.
2) Pseudomonas tomato resistance (Pto)– AvrPto. (Mucyn, T. S. et al., 2006)
Pto is a tomato serinethreonine protein kinase. Pto is polymorphic and hence satisfies the genetic criteria for the definition of a disease resistance protein. Pto activity requires the NB-LRR protein Prf, and the proteins form a molecular complex. Prf is monomorphic, at least in the tomato species analyzed to date. Pto is the direct target of two unrelated P. syringae effectors, AvrPto and AvrPtoB, each of which contributes to pathogen virulence in Pto mutants. It is thus likely that Prf guards Pto. The Pto kinase is apparently not required for PTI, though there may be redundancy in its function because it is a member of a gene family.
3) Tomato Cf-2 – Cladosporium fulvum Avr2 (Rooney et al., 2005)
The transmembrane RLP Cf-2 guards the extracellular cysteine protease Rcr3. Cf-2 recognizes the C. fulvum extracellular effector Avr2, which encodes a cysteine protease inhibitor. Avr2 binds and inhibits the tomato Rcr3 cysteine protease. Mutations in Rcr3 result in the specific loss of Cf-2-dependent recognition of Avr2. Hence, Cf-2 seems to monitor the state of Rcr3, and activates defense if Rcr3 is inhibited by Avr2.
Indirect interaction (Guard hypothesis)
The R proteins interact, or guard, a protein known as the ‘guardee’ which is the target of the Avr protein. When it detects interference with the guardee protein, it activates resistance.
1. Arabidopsis RPM1(Resistance to Pseudomonas syringae ssp. maculicola 1) AvrRpm1 (Jonathan D. G et al.,2006)
Arabidopsis RPM1 is a peripheral plasma membrane NB-LRR protein. It is activated by either the AvrRpm1 or the AvrB effector proteins. AvrRpm1 enhances the virulence of some P. syringae strains on Arabidopsis as does AvrB on soybeans. AvrRpm1 and AvrB are modified by eukaryote-specific acylation once delivered into the cell by the type III secretion system (red syringe) and are thus targeted to the plasma membrane. The biochemical functions of AvrRpm1 and AvrB are unknown, although they target RIN4(RPM1 interacting 4), which becomes phosphorylated (1P), and activate RPM1. In the absence of RPM1, AvrRpm1 and AvrB presumably act on RIN4 and other targets to contribute to virulence.
How to Use the Gene-for-Gene Concept in Disease Management
R-GENE APPLICATIONS – REALISING THE POTENTIAL
There are two approaches through which R genes control diseases.
1) Classical method- This involves conventional breeding approach.
Perez et al., (2007) introgressed three bacterial blight resistance genes, Xa4, Xa7 and Xa21, in to a temperature-sensitive genetic male sterile (TGMS1) line through three way crosses
2) Transgenic method- This involves non-conventional method or Transgenic approach.
Tang et al., (2001) used particle bombardment to co-transform mature seed-derived rice callus (Oryza sativa L., ssp. japonica, cv.) with plasmids containing the linked marker genes gusA and hpt, and the ap1 gene encoding an amphipathic protein previously shown to delay the hypersensitive response induced in non-host plants by the pathogen Pseudomonas syringae pv. syringae (Pss). A bacterial blight inoculation test was carried out on ten lines. In each case, plants carrying the ap1 gene showed enhanced resistance to Xanthomonas oryzae pv. oryzae (Xoo) race 6 at various levels.
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2 . Douglas C. Boyes, Jaesung Nam, and Jefert L. Dangl (1998) The Arabidopsis thaliana RPM1 disease resistance gene product is a peripheral plasma membrane protein that is degraded coincident with the hypersensitive response Proc. Natl. Acad. Sci. USA Vol. 95, pp. 15849–15854.
3 . Jeffrey G. Ellis, Peter N. Dodds, and Gregory J. (2007) Lawrence Flax Rust Resistance Gene Specificity is Based on Direct Resistance-Avirulence Protein Interactions . Annu. Rev. Phytopathol.45:289–306
4 . Jonathan D. G. Jones & Jeffery L. Dang (2006) The plant immune system. NATURE. Vol 444.
5 . Kerry F. Pedley1 and Gregory B. Martin (2003) Molecular basis of Pto mediated resistance to bacterial speck disease in Tomato. Annu. Rev. Phytopathol. 41:215–43
6 . Kexuan Tang a, b, Xiaofen Sun a, Quanan Hu a, Aizhong Wu b, C. H. Lin c, (2001) Transgenic rice plants expressing the ferredoxin-like protein (AP1) from sweet pepper show enhanced resistance to Xanthomonas oryzae pv. Oryzae. Plant Science 160 : 1035–1042
7 . Loida M. Perez, Edilberto D. Redoña, Merlyn S. Mendioro, Casiana M. Vera Cruz, Hei Leung. (2008) Introgression of Xa4, Xa7 and Xa21 for resistance to bacterial blight in thermosensitive genetic male sterile rice (Oryza sativa L.) for the development of two-line hybrids. Euphytica, 164 :627-636
8 . MHAJ Joosten and PJGM de wit (1999) THE TOMATO–CLADOSPORIUM FULVUM INTERACTION: A Versatile Experimental System to Study Plant-Pathogen Interactions. Annu. Rev. Phytopathol. 37:335–67
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regulate specific plant immunity. Plant Cell, 18: 2792-2806
10 . Rooney, H. C., Van't Klooster J. W., van der Hoorn R. A., Joosten M. H., Jones J. D., de Wit P. J. (2005) Cladosporium Avr2 inhibits tomato Rcr3 protease required for Cf-2-dependent disease resistance. Science 308: 1783–1786.
11. Wu, A. J., Andriotis, V. M., Durrant, M. C. & Rathjen, J. P. (2004) A patch of surface exposed residues mediates negative regulation of immune signaling by tomato Pto kinase.
Plant Cell 16: 2809–2821
About Author / Additional Info:
I did Ph. D. with specialization in Plant Molecular biology and Biotechnology and 3 years of research experience in Agriculture biotechnology.
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