R gene mediated
response
(a)
gene AvrR
S
gene AvrS
S
Host cellHost cell Host cell Host cell Avirulent pathogen Virulent pathogen Virulent pathogen Avirulent pathogen R-gene R-gen
Gene for gene model (Flor, 1971)
- specific response
- usually used in plant breeding
- mechanism widely known
Maize
Hm1/Hm2
against
Cochliobolus carbonum
race 1
encode HCTR (toxin reductase) that inactivates
Pto
Ser/Thr PKase
N, L6
TIR/NBS/LRR
Rps2, Rpm1
LZ/NBS/LRR
Cf-2, Cf-9
eLRR
Xa21
eLRR, quinase
TIR
NBS
LRR
N
C
Main types of R genes
CC
NBS
LRR
N
C
quinase
quinase
TM
LRR
N
C
TM
LRR
N
C
N
C
Signal
transduction
Protein-protein
interaction
signaling
Transmembrane
domain
Nucleotide
binding
protein-protein
interaction, dimers
Protein domains present in
different types of R genes
Domain swaps between tomato
Cf-9 and Cf-4
identified
LRR as responsible of the
specificity of the pathogen
race....
... But in flax
L
gene is the
TIR domain
Protein
degradation
Zinc finger
Membrane
anchor
Recognition
specificity
Gen Especie Patógeno Avirulencia Tipo R-gen Referencia
N tabaco TMV replicasa TIR-NB-LRR Witham et al. 1994
Rps2 Arabidopsis Pseudomonas syringae AvrRpt2 CC-NB-LRR Bent et al. 1994
Rpp5 Arabidopsis Peronospora parasitica AvrRpp5 TIR-NB-LRR Parker et al., 1997
Mi tomate Meloidogyne incognita Macrosiphum euphorbia
- CC-NB-LRR Milligan et al., 1998
Tabla 3.1. Algunos ejemplos de R-genes del tipo NB-LRR activos frente a distintos grupos de patógenos. Se incluye un gen de resistencia activo frente a un virus, una bacteria, un hongo y un nemátodo. Adicionalmente el gen Mitambién confiere resistencia frente a un insecto. Se añade la avirulencia correspondiente si es conocida.
TMV: virus del mosaico del tabaco; Pseudomonas syringae: bacteria; Peronospora parasitica: hongo; Meloidogyne incógnita: nemátodo; Macrosiphum euphorbia: insecto
Gen Especie Patógeno Avirulencia Tipo R-gen Referencia
Pto tomate Pseudomonas syringaepv. tomato AvrPto Serina/treonina quinasa Martin et al. 1993
N tabaco TMV replicasa TIR-NB-LRR Witham et al. 1994
Rps2 Arabidopsis Pseudomonas syringae AvrRpt2 CC-NB-LRR Bent et al. 1994
Cf-9 tomate Cladosporium fulvum Avr9 eLRR-TM Jones et al. 1994
Xa21 arroz Xanthomonas oryzaepv. oryzae AvrXa21 eLRR-TM-quinasa Song et al., 1995
Ve1, Ve2
tomate Verticillium albo-atrum - CC-eLRR-TM eLRR-TM-PEST
Kawchuk et al. 2001
Rrs1 Arabidopsis Ralstonia solanacearum PopP2 NB-LRR-WRKY Deslandes et al. 2002
Rpw8 Arabidopsis Erysiphe cichoracearum - CC Xiao et al., 2001
Rpg1 cebada Puccinia graminisf. sp. tritici - Receptor quinasa Brueggeman et al. 2002
Tabla 3.2. Algunos ejemplos de distintos tipos de R-genes. Se incluye solamente un ejemplo para cada tipo de R-gen, además de algunos R-genes adicionales que presentan algunas particularidades estructurales que los diferencian de los grupos anteriores, como hemos comentado en el texto. Se añade la avirulencia correspondiente si es conocida.
TIR: Dominio ‘TOLL-Interleukin-1 resistance’; NB: dominio de unión a nucleótidos; LRR: repetición rica en leucina; CC: dominio coiled-coil; eLRR: repetición rica en leucina extracelular; PEST: dominio señal de degradación proteolítica; WRKY: dominio de unión a DNA
Melon BAC containing an NBS-LRR cluster
Gene
Homology
MRGH (7.5x)
TIR-NBS-LRR resistance genes
MRGH fragment (3x)
TIR-NBS-LRR resistance genes
WD-40
WD40 domain, signal transduction/pre-mRNA processing/cytoskeleton assembly
Dynamin
Dynamin GTPase domain, signalling proteins
Exostosin
Exostosin family, limonene cyclase, tumor suppressor genes
Unknown1
Hypothetical and unknown proteins rice and
A. thaliana
PDI (4x)
Thioredoxin domain, postranslational modification, protein turnover
Ulp1 (2x)
Ulp1 protease family, postranslational modification, protein turnover
PPR
Pentatricopeptide repeat protein, plants, RNA stabilisation/processing
RLK
Serine/Threonine protein kinase and legume lectins beta domain
RTP-Copia (3x)
Retrotransposable elements, Copia type, 2 full length, 1 fragment
LINE (3x)
LINE transposable elements, 1 full length, 2 fragments
Pathogen factor Host factor Pathogen factor Host factor
Recognition,
susceptibility
Pathogen factor Pathogen factorNo recognition,
resistance
Host factor Host factorRecessive resistance genes
-
Do not follow gene-for-gene
-
Mutations in host genes
-
Necessary for susceptibility
-
Negative regulators of defence
•
Genus
Carmovirus
, Family
Tombusviridae
•
Transmitted by
Olpidium bornovanus
and
through the
seed
•
Affects
cucurbit crop
species grown under
greenhouse
conditions
•
Endemic
in cucurbit crops world-wide
•
yield losses
Disease Control
•
Certified seeds
•
Control of fungal vector
•
graffting melon in wattermelon
resistance stock
•
Use of
genetic resistance
(restricted to melon, recessive
and monogenic gene:
nsv
)
MNSV-264
2
MNSV-M
5
2
MNSV-Dutch
1
Nsv/-(Susceptible)
nsv/nsv
(Resistant)
1
Rivière and Rochon (1990) J. Gen. Virol 71: 3033-3042
2
Díaz et al (2004) MPMI 17: 668-675
Isolation of MNSV strain that
7K20 14 1 1 5 1 1 2 2
M132
65nsv
D08
24O20 38B12 U 41I23 SP6 U 5A6 SP6 U 30L11 U SP6 34H17 5B3 U SP6 10G6 SP6 20E15 SP6 U 9N8 U 10O16 SP6 U 20F8b SP6 59I22b U SP6 5 2 1 0 4 0 34 52K20 SP6 100 kb 1-21-10 L R X 15 M 132 30L11s p6 5B 3sp 6 108 7 13 59nsv
ns1 (M29)
ns2 (5A6sp6)
5A 6U 38B 12UF2
BC1
Map-based cloning of major genes: the
nsv
locus
for resistance to MNSV
Morales et al. TAG (2006)
- Based on 3.000 F2 and
BC1 individuals
-
BAC libraries from
WMR29 and PIT92
Nieto et al. Plant J (2006)
Eukaryotic translation initation complex
nsv
is a mutant allele of the Cm-eIF4E gene
Co-bombardment of MNSV
and Cm-eIF4E in melon leaves
Study of complex traits: the melon NIL collection
•
PI 161375 x T111
•
57 NILs
•
Average
introgresion size: 47
cM (3.4% genome)
•
Each linkage group
is represented by
4.75 NILs
•
Genetic resolution:
49 bins, 18.9 cM/bin
•
Subset of 28 NILs
cover the whole
genome
QTLs for fruit shape
Piel de sapo
SC3-5
QTLs for fruit ripening
QTLs for CMV resistance
Genetic dissection of complex traits using
the melon genetic map and NILs
cmv1,
a recessive gene that confers
resistance to some CMV strains
Essafi et al. TAG (2008)
S
S
S
S
R
S
S
S
S
M730
S
S
R
S
FNY
S
S
R
S
TL
S
S
R
S
M6
R
S
R
S
LS
R
S
R
S
P104.82
R
S
R
S
P9
12-1
12-3
PI
161375
PS
Strains
R
S
-
Few sources of resistance to
CMV in melon. One of them in
PI 161375
- Oligogenic recessive (
Karchi
et al
, 1975
)
- Several QTLs detected in PI 161375,
with a major QTL in GLXII, common
to 3 CMV strains (
Dogimont
et al
, 2000
)
eIF candidate genes do not
map in the
cmv1
interval
A fine map of the
cmv1
region has been started
84.7 11.2 18.5 38.8 56.1 61.9 89.4 5A6U CSAT425A TJ29 CMTCN14 ECM105 CMN21_55 CMN61_44 PSI 22B02 CMAGN33 CMGAN80 ECM67 0 6.1 8.3
S
S
R
S
R
cmv1
Pathogen inoculation
S
R
DNA extraction
PCR
Marker Assisted
Selection (MAS)
R
M
1 cM
R R S S R R R S R R
SCAR marker for resistance gene
to
Verticillium
in tomato
SCAR marker for resistance gene
to
Fusarium oxysporum
in melon
P1 P2 H
F2
R S R R R R R R R S R R R R R S R S S S R R