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Plant disease resistance genes: Current status and future directions
Article in Physiological and Molecular Plant Pathology · January 2012
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Physiological and Molecular Plant Pathology 78 (2012) 51e65
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Physiological and Molecular Plant Pathology

journal homepage: www.elsevier.com/locate/pmpp
Plant disease resistance genes: Current status and future directions

Mayank Anand Gururani a,1, Jelli Venkatesh a,1, Chandrama Prakash Upadhyaya b, Akula Nookaraju c,
Shashank Kumar Pandey c, Se Won Park a, *
a
Dept. of Molecular Biotechnology, Konkuk University, 1 Hwayang-dong, Gwangjin-gu, Seoul 143-701, Republic of Korea
b
Dept. of Botany, Guru Ghasidas University, Bilaspur, CG 495009, India
c
Dept. of Bioenergy Science and Technology, Chonnam National University, 333 Yongbongro, Buk-gu, Gwangju 500-757, Republic of Korea
a r t i c l e i n f o a b s t r a c t
Article history: Plant diseases can drastically abate the crop yields as the degree of disease outbreak is getting severe
Accepted 15 January 2012 around the world. Therefore, plant disease management has always been one of the main objectives of
any crop improvement program. Plant disease resistance (R) genes have the ability to detect a pathogen
Keywords: attack and facilitate a counter attack against the pathogen. Numerous plant R-genes have been used with
Plant diseases varying degree of success in crop improvement programs in the past and many of them are being
Resistance genes
continuously exploited. With the onset of recent genomic, bioinformatics and molecular biology tech-
Plant pathogens
niques, it is quite possible to tame the R-genes for efficiently controlling the plant diseases caused by
Disease management
pathogens. This review summarizes the recent applications and future potential of R-genes in crop
disease management.
Ó 2012 Elsevier Ltd. All rights reserved.
1. Introduction such crops. However, in case of conventional breeding for resis-

tance, the introgression of resistance genes from one species into
Plant pathogen interaction is a well understood mechanism the gene pool of another by repeated backcrossing is a long-term
which involves the activation of signals sometimes resulting in process which usually takes many hybrid generations before the
a rapid defense response against an array of plant pathogens. This backcrossing occurs. It is assumed that the complete functional
response helps the host plant to avoid further infection of the studies, cloning, characterization and genetic transformation of
disease. Induction of plant defense signaling involves the recogni- plant resistance genes could help the researchers to overcome
tion of specific pathogen effectors by the products of specialized these problems in near future.
host genes called R-genes [13]. Numerous individual plant resis- Efficient and sustained control of pathogens such as bacteria,
tance (R) genes have already been characterized and are being fungi, oomycetes, viruses, nematodes and insects is an exigency for
efficiently used in crop improvement research programs. Using all agricultural systems. In spite of the continued release of new
plant resistance genes for developing disease-resistant varieties is resistant cultivars, the global yield losses caused by pathogens are
a convenient alternative to other measures like pesticides or other substantial [8,188]. Plant pathogens not only decrease the crop
chemical control methods employed to protect crops from diseases. yields, they also lower the crop quality by releasing toxins that
Benefits of using the plant resistance genes in resistance breeding affect human health. Moreover, pathogens are constantly becoming
programs include the efficient reduction of pathogen growth, resistant to existing resistance genes and pesticides. This situation
minimal damage to the host plant, zero input of pesticides from the therefore demands some alternate methods of disease control. Crop
farmers and most importantly the environment friendly nature of improvement programs based on plant disease resistance genes are
being optimized by incorporating molecular marker techniques
and biotechnology. Therefore, plant resistance genes need to be
Abbreviations: Avr, Avirulence; CC, Coiled coil; HR, Hypersensitive reaction; LRR, studied extensively to alleviate the existing problem of pest and
Leucine rich repeats; NBS, Nucleotide-binding site; Pto, Pseudomonas tomato diseases apart from the abiotic challenges [147]. Facing selective
resistance; R, Resistance; RPP5, Resistance to Peronospora parasitica 5; RPS2, pressure imposed by the pathogens, plants have evolved post
Resistance to Pseudomonas syringae 2; TIR, Toll/interleukin-1-receptor homology invasion resistance mechanisms, often controlled by dominant
region.
* Corresponding author. Tel.: þ82 10 2450 3310.
resistance genes, whose products directly or indirectly detect
E-mail addresses: [email protected], [email protected] (S.W. Park). specific pathogen effectors and trigger effective defense responses
1
Authors contributed equally. [40,122]. R protein-triggered resistance to various pathogens is
0885-5765/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.pmpp.2012.01.002
52 M.A. Gururani et al. / Physiological and Molecular Plant Pathology 78 (2012) 51e65
normally race-specific and only effective against pathogen strains change the physiological state of host plant in order to benefit
expressing the cognate effector protein (Avr protein) recognized by pathogen colonization or are used to interrupt the activation of host
the R protein. This resistance is often associated with a hypersen- plant defenses [44,91]. However, plants have subsequently devel-
sitive response (HR), which is manifested as rapid death of the oped a form of immunity that is based on perception of these
invaded cell and in some cases a few surrounding cells proteins [185] by host resistance proteins called R-gene mediated
[89,96,179,260]. The structural and functional analysis of plant pathogen resistance.
resistance genes and R-gene loci is relevant for assembling various In gene-for-gene relationships, a plant carrying a resistance
resistance sources effectively and for engineering new strategies for gene resists pathogen races with the corresponding effectors
disease resistance in agriculture. Apart from that, it is highly [67,132,281]. The effectors found in bacteria, virus, nematodes,
desirable to understand the plantepathogen interaction in order to fungus, oomycetes and insects cause a plant pathogen to elicit
achieve the said goals. These aspects have been discussed in detail a resistance response in a host plant (Fig. 1). The effector genes are
later in the present review which would be beneficial for defined by corresponding resistance genes of which a relatively
researchers engaged in plant disease control based projects. The large number have now been cloned [162]. This resistance response
present article also highlights the concernment of many recent is appended with another reaction called hypersensitive reaction
investigations regarding the plant resistance genes and their (HR) which is a form of programmed cell death. The signaling
dispensation in the field of plant disease management strategies. cascade behind the HR is triggered either when an appropriate
disease resistance gene recognizes an effector or by an elicitor of
2. Plant basal disease resistance plant defense responses recognized by a specific receptor [177,184].
Either of these signals accompanied by other factors like influx of
Plants possess two major types of disease resistance, basal Ca2þ ions from the extracellular space and/or anion flux results in
defense and R-gene mediated defense (Fig. 1). Basal defense, which an oxidative burst producing reactive oxygen intermediates (ROIs)
can be a constituent of both non-host and host resistance, provides and defense gene activation, finally resulting in development of
first line of defense to the infection by a wide range of pathogens. local and systemic disease resistance [233,316,318].
Often, the plant disease resistance is cultivar or accession specific A well characterized example of HR elicitation through gene-for-
which is referred as host resistance whereas non-host resistance is gene interaction is provided by the tomato (Solanum lycopersicon)
the resistance against pathogens throughout all members of Cf-9 gene, which confers resistance to races of the fungus Clado-
a plant species [95,97,254] that is expressed when a plant comes sporium fulvum expressing the Avr9 gene [279]. Treatment of leaves
into contact with a pathogen which is incapable of provoking any of Cf-9 tomato or transgenic Cf-9 tobacco (Nicotiana tabacum) with
disease [98]. Elicitors of basal defense can be plant cell wall- the Avr9 peptide induces HR [90] and Avr9-treated Cf-9 tobacco cell
derived components released by hydrolytic activity of enzymes cultures showed rapid production of ROS and activation of MAP
secreted by invading pathogens, but also common features of the (Mitogen Activated Protein) kinases and calcium-dependent
pathogen, referred as pathogen-associated molecular patterns protein kinases [220,221]. The interaction between rice (Oryza
(PAMPs), such as lipopolysaccharides, chitins, glucans and flagel- sativa) and the fungal pathogen Magnoporthe grisea (Hebert) Barr
lins [187,222,236,325]. Non-pathogens as well as pathogens can (anamorph Pyricularia grisea Sacc.) causing the devastating rice
trigger a basal resistance in plants due to the widespread presence blast disease is another example of well documented gene-for-gene
of these molecular components in their cells [69]. However, system [134,247,278]. M. grisea has the Avr-Pita gene containing the
adapted microbes express a suite of effector proteins that often act C-terminal 176 amino acids which functions as an elicitor molecule
to suppress these defenses. Subsequently, plants have evolved that directly binds the Pita protein of rice and triggers a signal
other receptors (R proteins) that detect these pathogen effectors cascade leading to resistance [113].
and activate strong defenses [19]. Despite several studies and intense efforts with numerous sets
of R and Avr proteins [113,266], the interaction between R and Avr
3. R-gene mediated pathogen resistance proteins remained inexplicit and the insufficiency of verifiable R-
Avr interactions led to the formulation of the ’guard hypothesis’
Phytopathogens produce certain molecules called ‘effectors’, [165,270,279,280]. According to this model, the R proteins activate
encoded by Avr (avirulence) genes, which are delivered directly into resistance when they interact with another plant protein known as
the plant cells during initial stage of infection. These effectors either guardee protein that is targeted and modified by the pathogen in
Fig. 1. Plant pathogen interaction and development of disease resistance.

M.A. Gururani et al. / Physiological and Molecular Plant Pathology 78 (2012) 51e65 53
order to create an appropriate environment. Resistance is initiated spanning domains (Fig. 3, Table 1). The LRRs (Leucine rich repeats)
when the R protein detects an attack of its guardee or, in some cases represents the components having an important role for recogni-
when the R protein recognizes the product of the pathogen attack tion specificity and these domains are present in the majority of R
[244], which might not necessarily involve direct interaction proteins [121].
between the R and Avr proteins [165], (Fig. 2). To date, the most First major class of R-genes include the genes encoding for
convincing evidence for the guard hypothesis has been found in cytoplasm proteins with a nucleotide-binding site (NBS), a C-
Arabidopsis thaliana bacterial R-Avr systems [158] where RIN4 terminal leucine rich repeat (LRR) and a putative coiled coil domain
(RPM1- interacting protein 4) was identified as a cellular protein (CC) at the N- terminus. The examples of this class of resistance
that is required for the resistance to Pseudomonas syringae pv. genes include the P. syringae RPS2 and RPM1 resistance genes of
tomato mediated by RPM1 and RPS2. The RIN4 (guardee) is modi- Arabidopsis and the tomato Fusarium oxysporum resistance gene I2.
fied in various ways, depending on the Avr that it associates with, The second class of resistance genes consists of cytoplasmic
and these modifications then serve to activate the corresponding R proteins which possess LRR and NBS motifs and an N-terminal
protein (guard). Another example is the cleavage of the A. thaliana domain with homology to the mammalian toll-interleukin-1-
kinase PBS1 (guardee) by the cysteine protease AvrPphB from receptor (TIR) domain. The tobacco N gene, flax L6 gene and RPP5
P. syringae pv. tomato, which results in activation of RPS5 (guard)- gene are a few examples categorized under this class [146]. Third
mediated resistance [244]. Recently, it was shown that AvrPphB, major class of resistance genes family devoid of NBS motif consists
a cysteine protease, binds PBS1 and cleaves it, which triggers RPS5- of extra cytoplasmic leucine rich repeats (eLRR), attached to
mediated resistance, indicating that RPS5 might sense the integrity a transmembrane domain (TrD). eLRRs are known to play an
of PBS1 [242,243]. important role for certain defense proteins such as, poly-
Several genes have been implicated in the regulation of resis- galacturonase inhibiting proteins (PGIPs) [119] even though they
tance gene function; of these,Rar1 and Sgt1 are among the most are not directly involved in pathogen recognition and activation of
extensively studied genes. It has been reported that Rar1 and Sgt1 defense genes [121,256]. The C. fulvum resistance genes (Cf-9, Cf-4
are required in multiple R-gene mediated and non-host resistance and Cf-2) having an extracellular LRR (eLRR), a membrane span-
responses to a variety of pathogens [198,199,234]. A notable ning domain, and a short cytoplasmic C terminus [150] are some
example is in barley where the regulation of Mla transcript accu- examples of this class of resistance genes. The rice Xa21 resistance
mulation is not constitutive and that induction is coordinately gene for Xanthomonas is an example of the fourth class of resistance
controlled by recognition-specific factors [88]. Rar1 from barley has genes which consists of an extracellular LRR domain, a trans-
been identified as a required component for resistance against membrane domain (TrD) and an intracellular serine-threonine
powdery mildew (Blumeria graminis f. sp. Hordei) mediated by kinase (KIN) domain [252].
Mla12 [274] which is required for a subset of R-gene mediated The fifth class of resistance genes contain the putative extra-
resistance responses in monocot and dicot plant species cellular LRRs, along with a PEST (Pro-Glu-Ser-Thr) domain for
[155,182,237,246]. Sgt1 interacts with Rar1, and contributes to R- protein degradation (found only in Ve2, and not Ve1), and short
gene mediated resistance [7,154,155] although recently, Bhaskar proteins motifs (ECS) that might target the protein for receptor
et al. [21] demonstrated that Sgt1, but not Rar1, is essential for the mediated endocytos (e.g. tomato Ve1 and Ve2 genes) However,
RB-mediated broad-spectrum resistance to potato late blight. these Ve1 and Ve2 proteins have recently been proposed as PAMP
Similarly, Hein et al. [99] reported that Hsp90 (heat shock protein receptors [270].
90), a molecular chaperone and one of the most abundant proteins The Arabidopsis RPW8 protein is an example of the sixth major
expressed in cells was found as a required component for Mla13- class of resistance genes which contains a membrane protein
mediated race-specific resistance. domain (TrD), fused to a putative coiled coil domain (CC) [299]
whereas, the seventh major class of resistance genes includes the
4. Major classes of R proteins Arabidopsis RRS1-R gene conferring resistance to the bacterial
phytopathogen Ralstonia solanacearum, and it is a new member of
Plant resistance genes can be broadly divided into eight groups the TIReNBSeLRR R protein class. RRS1-R has a C-terminal exten-
based on their amino acid motif organization and their membrane sion with a putative nuclear localization signal (NLS) and a WRKY
Fig. 2. Guard hypothesis e the plant R proteins (guard) are associated with the endogenous host protein (guardee) which are common target proteins for the pathogens. The
interaction of effector pathogen proteins with the host proteins, causes a change in their structure which is then recognized by the guard proteins. As a result, a pathogen response
signaling cascade is triggered against the microbial evasion.
Fig. 3. Major classes of plant resistance (R) genes based on the arrangement of the functional domains. LRR e Leucine rich repeats; NBS e Nucleotide-binding site; TIRToll/
Interleukin-1- receptors; C-C e Coiled coil; TrD e Transmembrane domain; PEST e Protein degradation domain (proline-glycine-serine-threonine); ECS e Endocytosis cell
signaling domain; NLS e Nuclear localization signal; WRKY e Amino acid domain; HM1 e Helminthosporium carbonum toxin reductase enzyme.
domain [52,53]. The WRKY domain is a 60 amino acid region that is Though most of the resistance genes show dominant inheri-
defined by the conserved amino acid sequence WRKYGQK at its N- tance, recessive resistance is fairly common in viral systems [130],
terminal end, together with a novel zinc-finger-like motif. (Section 4.5) Recessive resistance genes in bacterial and fungal
The eighth major class of resistance genes includes the enzy- plant pathogen interactions have also been reported, such as barley
matic R-genes which contain neither LRR nor NBS groups. For mlo [32], Arabidopsis RRS1-R [53], rice xa13 [42], and xa5 [106,116].
example the maize Hm1 gene which provides protection against With the onset of functional genomics approaches and complete
southern corn leaf blight caused by the fungal pathogen Cochlio- genome sequencing of some important crop plants, the identifi-
bolus carbonum [117]. Unlike other resistance genes, Hm1 encodes cation and deployment of R-genes has become easier. Numerous
the enzyme HC toxin reductase, which detoxifies a specific cyclic resistance genes conferring resistance against a range of pathogens
tetrapeptide toxin produced by the fungus (HC toxin) that is have been successfully used in development of transgenic crops.
essential for pathogenicity. Therefore, cereal resistance genes like Therefore, the possibility of discerning some novel classes of
Hm1 can be seen to encode a range of different proteins that in resistance genes in near future cannot be ruled out.
some cases have obviously very different functions. Another
notable example, Pto protein in P. syringae contains a Ser-Thr kinase 4.1. Bacterial resistance genes
domain without LRRs [161] whereas, the Rpg1 gene of barley which
confers resistance to stem rust encodes a receptor kinase-like A number of plant resistance genes conferring resistance against
protein with two tandem protein kinase (kinaseekinase) domains bacterial attack have been studied so far (Table 2) and for the
and does not contain a strong membrane-targeting motif and majority of plant diseases, the genetics of susceptibility are less
known receptor sequences [31]. tangible. It has been known that bacterial pathogens of both plants
Table 1
Major classes of plant resistance genes e LRR e Leucine rich repeats; NBS e Nucleotide-binding site; TIR e Toll/Interleukin-1- receptors; CC e Coiled coil; TrD e Trans-
membrane domain; PEST e Amino acid domain; ECS e Endocytosis cell signaling domain; NLS e Nuclear localization signal; WRKY e Amino acid domain; HC toxin reductase e
Helminthosporium carbonum toxin reductase enzyme.
S. no Major R-gene classes Domains Example
LRR NBS TIR Kinase CC TrD PEST ECS NLS WRKY

I NBSeLRReTIR U U U X X X X X X X N, L6, RPP5
II NBSeLRReCC U U X X U X X X X X I2, RPS2, RPM1
III LRReTrD U X X X X U X X X X Cf-9, Cf-4, Cf-2
IV LRReTrDeKinase U X X U X U X X X X Xa21
V TrDeCC X X X X U U X X X X RPW8
VI TIReNBSeLRReNLS- WRKY U U U X X X X X U U RRS1R
VII LRReTrDePESTeECS U X X X X U U U X X Ve1, Ve2
VIII Enzymatic R-genes X X X U X X X X X X Pto, Rpg1
X X X X X X X X X X Hm1
U ¼ present.
X ¼ absent.
Table 2 toward incompatible pathogens involves the differential expression

Bacterial pathogens and interacting Avr-genes and R-genes. of the resistance gene in presence of the AvrXa27 effector [85]. A
Pathogen Host Avr-gene R-gene Reference dominant rice gene Os8N3 is an exception as it is up- regulated by
Xanthomonas Capsicum AvreBs2 Bs2 [177,259] a bacterial type-III effector protein, and that confers gene-for-gene-
campestris annumm specified disease susceptibility [126].
Xanthomonas Oryza sativa e NPR1 [39] Some bacterial resistant plant resistance genes may confer
oryzae AvreXa1 Xa1 [321]
resistance against unrelated or distantly related pathogens. Zhao
AvreXa21 Xa21 [252]
Pseudomonas Lycopersicum AvrePto, Pto [1,135,161,223] et al. [323] demonstrated the feasibility of non-host resistance gene
syringae pv esculentum AvrePtoB transfer between two cereal crops maize and rice. They proposed
tomato that a maize non-host resistance gene Rxo1 recognizes a rice
P. syringae Arabidopsis AvrRpm1, RPM1 [50,82,105,263] pathogen, Xanthomonas oryzae pv. oryzicola and causes bacterial
thaliana AvrB
AvrRpt2 RPS2 [17,102,176,304]
streak disease. Interestingly, Rxo1 was also found to confer resis-
AvrPphB RPS5 [108,257,301] tance to the unrelated pathogen Burkholderia andropogonis, known
AvrRps4 RPS4 [77,101] to cause bacterial stripe of sorghum and maize indicating that the
same gene controls resistance to both pathogens and non-patho-
gens of maize. The function of Rxo1 in rice thus demonstrates that
and animals deliver virulence proteins into the host cytoplasm via an NBS-LRR type of resistance gene can be effectively transferred
the type-III secretion system (T3SS), also called injectisome [54] between distantly related cereals [323].
which enables Gram negative bacteria to secrete and inject path-
ogenicity proteins into the cytosol of eukaryotic host cells [71,94]. 4.2. Fungal resistance genes
The T3SS is encoded by hrp (HR and pathogenicity) and hrc (HR and
conserved) genes, whose mutations eliminate bacterial pathoge- Fungal diseases are rated either the most important or second
nicity in susceptible host plants and the ability to elicit HR in non- most important factor contributing to yield losses in almost all the
host or cultivar-specific resistant plants. Many of the T3SS effector major crops [300]. So far, several fungal resistance genes (Table 3)
proteins have been shown to be dependent on molecular chaper- have been reported and used in crop improvement programs.
ones, which keep the effector in a partially unfolded form in the However, the sequence variation occurring within the central LRR
bacterial cytoplasm [255]. The emergent results on their role in domain and the variation in LRR copy number of the gene plays an
pathogenesis have indicated that they act as molecular double important role in determining recognition specificity [27,141]. For
agents betraying the pathogen to plant defenses in some interac- example, the sequence variations in tomato Cf-4 and Cf-9 genes
tions and suppressing host defenses in others [181]. play an important role in determining recognition specificity,
In rice, resistance and susceptible alleles of Xa27 encode iden- which confer resistance to biotrophic leaf mold pathogen Clado-
tical proteins however, expression of only the resistance allele sporium and induce a hypersensitive response (HR) upon recogni-
occurs when a rice plant is challenged by bacteria harboring tion of the fungus-encoded Avr4 and Avr9 peptides [27]. In tomato,
AvrXa27, whose product is a nuclear localized T3SS effector. Ve is involved in race-specific resistance to infection by Verticillium
Induction of Xa27 occurs only in the immediate vicinity of infected species [126]. The Ve1-mediated resistance signaling only partially
tissue, whereas ectopic expression of Xa27 results in resistance to overlaps with signaling mediated by Cf- proteins [191]. Recently,
otherwise compatible strains of the pathogen. The Xa27 specificity a virus induced gene silencing approach for the characterization of
Table 3
Fungal pathogens and interacting R-genes.
Pathogen Host Avr- gene R-gene Reference

Blumeria graminis Hordeum vulgarae AvrMla Mla [324]
e Mlo [32]
Cochliobolus carbonum Zea mays e Hm1 [117]
Cladospoium fulvum Lycopersicum Avr2 Cf-2 [157,224,251,284]
esculentum Avr4 Cf-4 [27,123,269]
Avr5 Cf-5 [55]
Avr9 Cf-9d [120]
Erysiphe orontii, E. cichoracearum Arabidopsis thaliana e RPW8.1, [299,314,317]
and Oidium lycopersici RPW8.2
Fusarium oxysporium Lycopersicum esculentum Avr1 I2 [189,248]
Melamspora lini Linum usitatissimum AyrL L [56,57,146]
AvrM M
AvrN AvrL567 genes, N
whose products are
recognized by the
L5, L6, and L7
Magnoporthe grisea Oryza sativa Avr-Pita Pieta [113,134]
Puccinia sorghi Zea mays AvrRPeIeD Rp1 [43]
Puccinia triticina Triticum aestivum e Lr46 [164,168]
Puccinia graminis f sp. tritici Hordeum vulgarae AvreRpg1 Rpg1, [31,103,136]
Rpg4,
Rpg5
Verticillium alboeatrum Lycopersicum esculentum e Ve1, Ve2 [2,131,293]
Mentha arvensis mVe1
Mentha longifolia
Verticillium dahliae Lycopersicum esculentum e Ve1 [68]
Ve mediated signaling revealed that signaling cascade downstream homolog of the C. fulvum resistance genes of the Vf region), of
of Ve1 requires two genes EDS1 (Enhanced Disease Susceptibility 1) which HcrVfa1, HcrVfa2 and HcrVfa4 encode typical RLPs while
and NDR1 (non race-specific disease resistance 1). Moreover, the HcrVfa3 contains an insertion at the end of the LRR motif, resulting
results showed that the locus Ve consists of two closely linked in truncated transcripts [292,315]. Only expression of HcrVfa1 or
inversely oriented genes, Ve1 and Ve2 encoding cell surface HcrVfa2 in susceptible apple cultivars provided resistance against
receptor proteins of the extracellular LRR receptor-like protein. Out V. inaequalis strains [12,159].
of them, only Ve1 provides resistance in tomato against race 1
strains of Verticillium dahliae and Verticillium albo-atrum and not 4.3. Oomycetes resistance genes
against race 2 strain. Based on the sequence analysis and the
expression study, Ve1 and Ve2 expression is induced in resistant as Phytopathogenic oomycetes are responsible for economically
well as susceptible tomato genotypes and that no single mutation important diseases, such as late blight of potato and sudden oak
in the CDS of Ve2 discriminates resistant and susceptible tomato death caused by Phytophthora infestans and Phytophthora ramorum
genotypes. However, a single point mutation in Ve1, resulting in respectively. The oomycetes (Pseudofungi) have been classified
a premature stop codon, was found in all susceptible genotypes and within the phylum Heterokontophyta comprising a number of
was absent in all resistant genotypes. This suggested that Ve1, but microbial lineages with phenotypic similarities to true fungi [216].
not Ve2, governs Verticillium resistance in tomato [68]. It was only with the use of molecular phylogenetic methods
A disease epidemic broke out in oats in the 1940’s due to the starting with small subunit rDNA analysis [34,35] followed by
extensive planting of ’’Victoria-type’’ oats carrying the Pc-2 gene multiple concatenated gene phylogenies [9] that the oomycetes
for resistance against the rust fungus, Puccinia coronata. Oats were demonstrated to group within the heterokont radiation [216].
carrying Pc-2 were highly susceptible to another disease, Victoria Several functional resistance genes from potato conferring
blight, caused by a fungus Cochliobolus victoriae [151,169]. Patho- resistance to late blight have been cloned and all of them
genicity of C. victoriae is dependent on the production of a toxin belong to the NBS-LRR class of plant resistance genes
called victorin, and in oats, both toxin sensitivity and Victoria [10,14e16,104,190,250,282,283]. In addition to the resistance to
blight disease susceptibility are conferred by the dominant Vb P. infestans genes Rpi-blb1 (RB) and Rpi-blb2, Solanum bulbocas-
gene. Despite extensive efforts, rust resistance (Pc-2) and Victoria tanum appears to harbor Rpi-blb3 located at a major late blight
blight susceptibility (Vb) have not been genetically separated and resistance locus on LG IV, which also harbors Rpi-abpt, R2, R2-
are suspected to share identity [298,312] thus suggesting an like, and Rpi-mcd1 in other Solanum spp [156]. Vleeshouwers
unexpected relationship between plant disease resistance and et al. [294] used a candidate gene approach for the rapid cloning
susceptibility. of S. stoloniferum Rpi-sto1 and S. papita Rpi-pta1, which are
Stem rust-susceptible barley cv. Golden Promise was trans- functionally equivalent to Rpi-blb1. Cloning and functional anal-
formed into a highly resistant one to pathotype Pgt-MCC of the yses of four Rpi genes, Rpi-blb3, Rpi-abpt, R2, and R2-like revealed
stem rust fungus Puccinia graminis f. sp. tritici by Agrobacterium- that these genes contain all signature sequences characteristic of
mediated transformation with the dominant Rpg1 gene. A single leucine zipper nucleotide-binding site leucine rich repeat (LZ-
copy of Rpg1 against stem rust, and progenies from several trans- NBS-LRR) proteins, and share 34.9% of amino acid sequences
formants segregated in a 3:1 ratio for resistance: susceptibility as similar to RPP13 from A. thaliana [149,193e195]. So far, a number
expected for Mendelian inheritance and unequivocally demon- of Hyaloperonospora parasitica resistance (RPP) genes against the
strated that the DNA segment isolated by map-based cloning is the downy mildew have been cloned from Arabidopsis which belong
functional Rpg1 gene for resistance to stem rust and the trans- to the NBS-LRR class of resistance genes [119,264]. These resis-
formants exhibited a higher level of resistance than the original tance genes are distinguished by their N-terminal regions,
sources of Rpg1 like cvs. Chevron and Peatland [103]. Another showing homology to the TIR domain (RPP1 and RPP5 clusters)
fungal resistance plant resistance gene RUS1 from Setaria italica and leucine zipper motifs (RPP8 cluster) [25,166,172]. Another
Beauv. cv. Shilixiang resistant to Uromyces S. italica, was cloned and example of oomycetes resistance genes with NBS-LRR motifs is
it was found to contain an NB- ARC (nucleotide-binding adapter downy mildew resistance gene, Dm3 [45,244,245] in Bremia lac-
shared by APAF-1, R proteins, and CED-4) domain as well as three tucae which is a member of the large RGC2 (Resistance Gene
conserved motifs P-loop, kinase 2, and kinase 3, having the char- Candidate2) multigene family similar to the genes cloned from
acteristics of NBS-LRR type resistance gene of plant [303]. other species for resistance to downy mildews and other patho-
Another notable example of fungal resistance genes is the gens [167].
broad-spectrum mildew resistance gene RPW8.2 from Arabidopsis Several oomycete effector genes (Table 4) encoding products
thaliana which is induced by powdery mildew [299] and is assumed that are recognized by R proteins situated in the plant cytoplasm
to be involved in enhancing the formation of a callosic encasement have been discovered which indicate toward a mechanism of
of the haustorial complex (EHC) with onsite accumulation of H2O2, transporting fungal and oomycete effectors into plant cells
in order to constrain the haustorium while reducing oxidative [5,241,271,273,294]. This mechanism has recently been character-
damage to the host cell. Targeting of RPW8.2 to the EHM (Extra ized using gene ontology by Torto-Alalibo et al. [275] while the
haustorial membrane) requires normal function of the actin cyto- motifs in their amino acid sequence have already been identified in
skeleton while microtubules are not involved in the process. the past [8,13,16]. The identification of the first effectors from
Despite its critical role for the defense function, SA signaling is oomycetes, together with whole genome sequencing projects has
dispensable for targeting RPW8.2 to the EHM and both EHM revealed a special class of secreted effector proteins, RXLR that are
localization and defense activation are required for RPW8.2 to delivered into host cells [4,6,81,83,212,277]. The RXLR effectors
induce resistance against powdery mildew [314]. constitute large super families of rapidly evolving proteins in all
The majority of resistance genes reside in clusters, and the oomycete genomes [58,115] and include Avr1b-1, Avr1a and Avr3a
frequency of recombination between clustered genes can vary from Phytophthora sojae [207,241], Avr3a, Avr4, and Avrblb1 from
remarkably, even within a single cluster. The Apple Vf locus, derived P. infestans [5,6,286,294], ATR1 and ATR13 from Hyaloperonospora
from the crab apple species Malus floribunda, confers resistance to arabidopsidis [5,212] and IpiO and IpiB from certain Phytophthora
five races of the apple scab fungus Venturia inaequalis. The Vf locus species including P. infestans [36,203,294]. While the majority of
comprises a cluster of four RLP genes, HcrVfa1 to HcrVfa4 (for IPI-O proteins are recognized by RB gene to elicit host resistance,
Table 4
Oomycetes pathogens and interacting Avr-genes and R-genes.
Pathogen Host Avr-gene R-gene Reference

Bremia lactucae Lactuca sativa Avr3 Dm3 [173,174]
Hyaloperonospora arabidopsis Arabidopsis thaliana ATR1 RPP1-Nd/WsB [212]
ATR13 RPP13eNd [5,23]
Perenospora parasitica A. thaliana AvrB, AvrRPP1A, RPP1, [25,166,196,197,280]
AvrRPP1B, AvrRPP1C, AvrRPP2, RPP2
AvrRPP4, AvrRPP5, AvrRPP8 RPP4, RPP5, RPP8
Phytophthora infestans Solanum tuberosum Avr1 R1 [10]

P. infestans Solanum tuberosum Avr-blb1 Rpieblb1 [294]
P. infestans Solanum tuberosum PiAvr2 Rpi [156,296]
P. infestans Solanum demissum Avr3a R3a [6]
P. infestans Solanum bulbocastanum Ipio, Ipib, Ipieo4 RB [36,287]
P. infestans Solanum tuberosum Avr3beAvr10eAvr11 locus, R3b, R10, R11 [114]
Phytophthora sojae Glycine max Avr1a, Avr3a and Avr3c, Rps1a [58,160,207]
Rps3a
Rps3c
some variants exist that are able to elude detection (e.g. Ipi-O4) Numerous sources of nematode resistance have been identified and
[87]. Intriguingly, few oomycete effectors that do not encode RXLR several of the responsible genes have been genetically mapped
effectors have also been proposed, such as Avr3b, Avr10 and Avr11 in [125,276,289,309,316].
P. infestans [114,208] and Avr1b-2 in P. sojae [241]. So far, the host Resistance to root-knot nematode was first identified in Lyco-
targets of RXLR effectors have not been well described in the persicum peruvianum Mill., a wild relative of cultivated tomato
literature [268], while the target proteins of several oomycete [302]. The single dominant Mi gene of tomato confers resistance to
apoplastic effectors have been determined [128,178,271,272]. three major root-knot nematodes Meloidogyne arenaria, Meloido-
P. sojae encodes numerous putative host cytoplasmic effectors gyne incognita and Meloidogyne javanica [79] but it does not confer
[1,24,59] with conserved FLAK (F, Phe; L, Leu; A, Ala; and K, Lys) resistance to Meloidogyne hapla, a nematode present in overlapping
motifs following signal peptides, termed crinkling- and necrosis- geographic locations [218]. Mi gene encodes a protein with CC-NBS-
inducing proteins (CRN) or Crinkler. Recently, the functional LRR motifs [175] was introduced into cultivated tomato using
studies of CRN revealed that two functional genes, PsCRN63 and embryo culture of an interspecific cross between Lycopersicum
PsCRN115 encode proteins that induce contrasting responses when esculentum and L. peruvianum [249], followed by extensive back-
expressed in Nicotiana benthamiana and soybean (Glycine max). crossing with L. esculentum. Later this gene was isolated by posi-
Silencing of the PsCRN63 and PsCRN115 genes in P. sojae stable tional cloning approach [175]. Mi-1 confers resistance to the root-
transformants exhibited a reduction of virulence on soybean and knot nematodes. The mechanism of resistance to nematodes
a loss of ability to suppress host cell death and callose deposition on conferred by Mi appeared to involve a hypersensitive response on
inoculated plants. These results suggested a role for CRN effectors the part of the host [60,61]. Mi-1 remains the only cloned root-knot
in the suppression of host defense responses [152]. In future, more nematode resistance gene [310] and the resistance mediated by Mi-
studies on oomycete effectors and their cognate host targets will 1 acts in a gene-for-gene manner.
undoubtedly explore novel plant immune pathways. Several common components that interact with R proteins or
required for resistance gene function have been recently identified
[235]. Bhattarai et al. [22] demonstrated the role of Hsp90, Sgt1, and
4.4. Nematode resistance genes
Rar1 in Mi-1-mediated aphid and nematode resistance. Studies
with approaches however identified the requirement of Rme1 gene
Plant parasitic nematodes are obligate parasites that obtain
for Mi-1-mediated resistance to nematodes, aphids, and whiteflies
nutrition from the cytoplasm of living plant cells and comprise
[22,163]. In addition to Rme1, Mi-1 resistance requires the salicylic
many species including ectoparasites and endoparasites. Nematode
acid (SA) signaling pathway and mitogen activated protein kinase
resistance genes are present in several crop species (Table 5) and
(MAPK) cascades [26,148]. The tomato MAPK kinases MKK2 and
form an important component in many breeding programs
MAPKs LeMPK1, LeMPK2, andLeMPK3 are required for Mi-1-
including those for tomato, potato, soybeans and cereals [276].
mediated aphid resistance [148]. However, their role in root-knot
nematode resistance has not yet been identified.
Table 5 The first nematode resistance gene to be cloned was Hs1pro-1,
Nematodes and interacting R-genes. a gene from a wild relative of sugar beet conferring resistance
Pathogen Host Avr-gene R-gene Reference against Heterodera schachtii, the beet cyst nematode [33]. Hs1pro-1
Melidogyne Lycopersicum e Mi [175,239]
cloned under the control of the CaMV35S promoter, was shown to
incognita esculentum confer nematode resistance to susceptible sugar beet roots trans-
Globodera Solanum e Hero, Gpa2 [66,227,297] formed with Agrobacterium rhizogenes [65] however, the resistance
pallida tuberosum mediated by Hs1pro-1, does not appear to involve a hypersensitive
Globodera Solanum e Hero, Gro1e4 [310]
response [124]. Complementation analysis by stable potato trans-
rostochiensis tuberosum
Heterodera Beta e HS1pro-1 [33] formation showed that the gene Gro1-4 conferred resistance to
schachtii vulgaris Globodera rostochiensis pathotype Ro1 and it encodes a protein of
Heterodera Triticum spp. e Cre3 [144,238] 1136 amino acids containing the TIR, NBS and LRR homology
avenae domains along with a C-terminal domain with unknown function
Melidogyne Capsicum e CaMi [37]
incognita annuum
[190]. The Gpa2 gene that confers resistance against some isolates
of the potato cyst nematode Globodera pallida, is a member of the
NBS-LRR-gene family and contains a possible LZ near its amino monogenic dominant resistance (Table 6). Although, these R
terminus. Gpa2 is highly similar in predicted amino acid sequence proteins appear to be similar, they confer resistance to highly
to the Rx1 gene which confers extreme resistance to Potato Virus X divergent viruses. For example, A. thaliana RCY1 (resistance to C
[227]. strain Y1) and HRT (HR to turnip crinkle virus) are allelic, encode
The Cre3 gene confers a high level of resistance to the root proteins that share 91% similarity [261] but confer resistance to
endoparasitic nematode Heterodera avenae in wheat. As a result of unrelated viruses such as cucumber mosaic virus (CMV, a cucumo-
map-based cloning of a disease resistance gene family at the Cre3 virus) and turnip crinkle virus (TCV, a carmovirus), respectively
locus, two genes related to members of the cytoplasmic NBS-LRR [253].
class of plant disease resistance genes have been analyzed. One The viral R protein-Avr system that strongly justifies the guard
encodes a polypeptide with a nucleotide-binding site (NBS) and hypothesis is the HRT-TCV pair. The TCV coat protein is the Avr
a leucine rich region; this member of the disease resistance gene determinant for HRT-mediated resistance responses and its inter-
family is expressed in roots. The second Cre3 gene sequence action with a host transcription factor, TCV-interacting protein (TIP)
appears to be a pseudo gene, with a frame shift caused by a deletion is required for HRT-elicited defense responses [214]. Although,
event [144]. Based on the conserved regions of known resistance a direct interaction between HRT and TIP has not been reported,
genes, an NBSeLRR-type CCN (cereal cyst nematode) resistance TCV coat protein inhibits the nuclear localization of TIP [215],
gene analog was isolated from the CCN resistant Ee10 near isogenic however it is possible that HRT detects the altered cellular distri-
lines (NILs) of wheat, designated as CreZ. The expression profiling of bution of TIP which might therefore be the guardee of the guard
CreZ indicated that it was specifically expressed in the roots of protein HRT. However knock out mutation studies [112] showed
resistant plants and expression levels drastically increased when that loss of TIP does not alter HR or resistance to TCV. Moreover, the
the plants were inoculated with cereal cyst nematodes [322]. In mutation in TIP neither impaired the salicylic acidemediated
addition, the wheat and barley resistance gene analogs (RGAs) induction of HRT expression nor the enhanced resistance
contain other conserved motifs present in known resistance genes conferred by overexpression of HRT. Noticeably, the mutation in TIP
from other plants and share between 55 and 99% amino acid resulted in increased replication of TCV and Cucumber mosaic virus,
sequence identity to the NBS-LRR sequence at the Cre3 locus and suggesting that TIP may play a role in basal resistance but is not
have been found to be associated with CCN and aphid resistance in required for HRT-mediated signaling. Resistance to Tomato Spotted
barley [238]. Wilt Virus (TSWV) in tomato is conferred by Sw-5 gene which was
In another example, a candidate root-knot nematode resistance introgressed from Solanum peruvianum into tomato, and has
gene (designated as CaMi) was isolated from the resistant pepper demonstrated broad and stable resistance [225]. The positional
line PR 205 which was highly expressed in roots, leaves, and cloning of Sw-5 locus was revealed that the resistance allele
flowers, and at a lower level in stems, and not detectable at all in encodes a CC-NBS-LRR R protein and is remarkably similar to the
fruits. Transgenic plants expressing CaMi gene triggered a hyper- tomato Mi gene for nematode resistance with the exception of four
sensitive response (HR) as well as many necrotic cells around leucine zippers at the N terminus [29].
nematodes and thus conferred significant resistance to root-knot In cultivated tomato, ToMV (Tomato mosaic virus) infections are
nematodes when compared to susceptible control plants [37]. controlled by the introgressed Tm-1, Tm-2 and Tm-22 genes. The
Tm-22 resistance gene was shown to be strikingly durable [86,202]
4.5. Viral resistance genes and it has been cloned and well characterized by Lanfermeijer et al.
[145]. The susceptible tomato plants, which were transformed with
The majority of characterized viral resistance genes from plants the Tm-22 gene, displayed resistance against ToMVinfection and the
fall into the NBS-LRR class of resistance genes, providing resistance was conserved in all transgenic lines. Similarly, Rai [209],
Table 6
Viral pathogens and interacting R-genes.
Pathogen Host Avr-gene R-gene Reference

Bean dwarf mosaic virus Phaseolus vulgaris Bdm BV1 protein [76]
Cucumber mosaic virus Arabidopsis thaliana Coat protein RCY1 [262]
Cucumber mosaic virus A. thaliana Vpg (viral genomeelinked protein) At-eIF4E1 (cum1) [72,320]
At-eIF4G (cum2)
Lettuce mosaic virus (LMV) Lettuce (Lactuca sativa) 3’half of genome mo1(1), mo1(2) [183,211]
Pea seed borne mosaic virus Pea (Pisum sativum) Vpg sbm1 [73,133]
P3 and 6K1 cistron sbm2 [118]
Potato virus X Solanum tuberosum Coat protein Rx1, Rx2 [14,15,205]
Potato virus Y Capsicum annuum VPg pvr1, pvr12 [129,180,228]
Potato virus X Solanum tuberosum Nla proteae Ry [170]
Potato virus Y Solanum tuberosum e Ye1 [291]
Potato virus Y, Tobacco etch virus Tomato (Lycopsersicon spp.) Vpg pot-1 [180]
Rice yellow mottle virus Oryza sativa Vpg eIF(iso)4G1 [100]
Soybean mosaic virus Glycine max HcePro and P3 cistron Rsv1 [64]
Tobacco etch virus Arabidopsis thaliana e RTM1, RTM2 [41,307]
Tobacco mosaic virus Solanum lycopersicon e N gene [305,306]
Replicase Tm1 [145]
30 kD movement protein Tm2, Tm22
Turnip mosaic virus, Arabidopsis thaliana VPg At-eIF(iso)4E [311]
Turnip mosaic virus Brassica napus TuRBO1, CI [109e111]
TuRBO1b,
TuRBO3,
TuRBO4, P3
TuRBO5, P3
TuMV P3 CI
Turnip mosaic virus Capsicum annumm Coat protein L1, L2, L3 [20,48,78]
cloned a single dominant gene Ctv-R present in the trifoliate rela- (Nasanova ribisnigri) [213], Sd1 gene confers resistance rosy leaf
tive of Citrus, Poncirus trifoliata conferring broad-spectrum resis- curling aphid (Dysaphis devecta) in apple [219] and the melon Vat
tance against Citrus tristeza virus (CTV), a major pathogen of citrus gene against the melon/cotton aphid Aphis gossypii [126,192].
[11,74,75,80,171,209]. Transgenic grapefruit plants carrying Citrus Triticum aestivum resistance to Hessian fly, Mayetiola destructor
Ctv-R gene were developed and it was found that two of the (Say), has also been demonstrated to be a gene-for-gene mecha-
candidate resistance genes, R-2 and R-3 were exclusively expressed nism [92], although no genes have been cloned yet, 26 resistance
resulting in either an absence of initiation of infection or its slow genes have been described as being effective against 13 biotypes of
spread in R-2 plant lines or an initial appearance of infection and its Hessian fly [63]. The occurrence of a hypersensitive response (HR)
subsequent eradication in some R-1 and R-4 plant lines [209]. in case of an insect attack still remains dubious, since both presence
Seo et al. identified the TIR-NBS-LRR gene RT4-4 involved in and absence of HR have been reported in incompatible interactions
a viral resistance response in common bean (Phaseolus vulgari cv. between wheat and Hessian fly [84,93,308].
Othello) [240] which functions across two plant families. The Recently, Klingler and co workers reported the presence of an
functional analysis revealed that the RT4-4 gene in transgenic N. HR response to bluegreen aphid and pea aphid in Medicago trun-
benthamiana lines is up- regulated in a non-virus-specific manner, catula [138]. A single gene AIN was found responsible to trigger HR
although RT4-4 did not confer resistance to the reporter virus, response against those two pathogens. However, it was also
it activated a resistance-like response (systemic necrosis) to concluded that although the HR response is triggered in both cases,
Cucumber Mosaic Virus (CMV). the resistance is conferred only to bluegreen aphid [138]. Irre-
Recent molecular cloning of recessive resistance genes to spective of presence or absence of HR, a common mechanism of R-
potyviruses led to the identification of resistance genes corre- gene mediated resistance to piercing, sucking insects appears to be
sponding to mutations in translation initiation factors, eukaryotic limited phloem-feeding [127,137,285]. A detailed description on
initiation factors 4E (eIF4E) and to a lesser extent, the eukaryotic planteaphid interactions along with a summary of recent studies
initiation factor 4G (eIF4G) [204]. The eIF4E gene provides resis- has recently been reviewed by Tagu et al. [258].
tance to several Potyviridae family viruses and has been identified
in the dicots, pepper (pvr1), pea (sbm1), lettuce (mo1 (1), mol (2)), 4.7. R-genes with broad range host resistance
tomato (pot1), and melon (nsv) and in the monocot barley (rym4/5)
[130,217,229]. Similarly, translation initiation factor eIF4G is A common strategy proposed to achieve broad-range host
responsible for resistance of rice to Yellow mottle virus [3] and in resistance is to modify the narrow pathogen specificity of R-gene
Arabidopsis to Cucumber mosaic virus and Turnip crinkle virus [320]. mediated resistance. Therefore, elucidation of R protein domains
that control recognition of specific pathogens and subsequent
4.6. Insect resistance genes activation of the downstream defense response has been the
subject of intense research [290]. The function of a particular
Studies using the model plant Arabidopsis have contributed resistance gene totally depends on the pathogen’s genotype
greatly to our understanding of R-gene mediated plant defense, [4,47,49,132,140] but there are some resistance genes which confer
especially against pathogens [103], as well as the basal defense resistance against a broad range of pathogens. For instance, the Mi-
mechanisms against aphid feeding [46,143,200,201]. Resistance to 1 gene in tomato confers resistance to root-knot nematodes
insects has been identified in various plant species since long back (Meloidogyne spp.), potato aphid M. euphorbiae [175,226,239,295],
[18,51,62,191,206] and a number of single dominant R-genes have whitefly B. tabaci [186], viruses [28], bacteria [231] and fungi
been mapped, and molecular markers linked to these loci have [189,248]. Tomato Pto-overexpressing plants show resistance not
been identified [30,107,139,153,155,288,319]. The majority of these only to P. syringae pv. tomato but also to Xanthomonas campestris pv.
mapped genes (Table 7) are in staple crops like wheat and rice. In vesicatoria and to the fungal pathogen C. fulvum [267,314]. Simi-
addition to these mapped genes, several single dominant aphid larly, the lettuce Dm3 gene confers resistance to lettuce downy
resistance genes have been identified that confer resistance to mildew (B. lactucae) as well as to lettuce root aphid [172]. Moreover,
a single species of insects [213]. Cloning of number of insect several other Dm specificities as well as resistance to lettuce root
resistance genes has been accelerated with the advent of high aphids have been shown to be conferred by members of the RGC2
throughput molecular tools, such as genome mapping, sequencing, family using RNAi approach [142,313].
and gene cloning.
To date, only few insect resistance genes belonging to NBS-LRR 5. Challenges and future directions
group of plant resistance genes have been cloned and character-
ized. For example, The tomato Mi-1 confers resistance to the potato With the advent of high throughput techniques and efficient
aphid (Macrosiphum euphorbiae) and whitefly (Bemisia tabaci), genomic approaches, researchers have managed to produce a large
Lettuce Nr-gene confers resistance to a single species of aphid amount of experimental data in the form of ESTs, whole genome
sequences, gene expression data etc. Still, the progress in under-
standing the functional mechanism of resistance genes has been
Table 7 moderate. For instance, little is known about the structural basis of
Insects and interacting R-genes.
pathogen recognition. Furthermore, there is still an inadequacy of
Pathogen Host Avr-gene Regene Reference a reference set of sequences to be used as model for resistance
Macrosiphum Lycopersicum e Mi [226] genes that usually cluster in genomic regions with a high number of
euphorbiae esculentum homologs and pseudo genes. The difficulties in performing the
Nasanova Lactuca sativa e Nr [285]
plantepathogen interaction studies pose another obstacle [70].
ribisnigri
Dysaphis Malus domestica e Sd1 [219] Nevertheless, efficacious applications are being continuously
devecta developed based on our rather finite knowledge base. For example,
Sogatella Oryza sativa e Qbp1, [265] recently PRGdb, a web accessible open source database providing
furcifera, Qbp2 a comprehensive overview of resistance genes has been developed
Nilaparvata
lugens
[232], which is definitely going to help filling some gaps in the
models of the plant defense signal transduction network.
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Plant disease resistance genes: Current status and future directions

Article in Physiological and Molecular Plant Pathology · January 2012

Mayank Anand Gururani

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Physiological and Molecular Plant Pathology

Plant disease resistance genes: Current status and future directions

1. Introduction such crops. However, in case of conventional breeding for resis-

Fig. 1. Plant pathogen interaction and development of disease resistance.

S. no Major R-gene classes Domains Example

LRR NBS TIR Kinase CC TrD PEST ECS NLS WRKY

Table 2 toward incompatible pathogens involves the differential expression

Pathogen Host Avr- gene R-gene Reference

Pathogen Host Avr-gene R-gene Reference

Phytophthora infestans Solanum tuberosum Avr1 R1 [10]

Pathogen Host Avr-gene R-gene Reference

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