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Alternaria brassicicola , and Fusarium oxysporum in A . thaliana (Llorente et al.
2005 ; Trusov et al. 2006 ). In contrast, mutations in the Arabidopsis
β
subunit AGB1
and the Arabidopsis
subunit AGG1 enhanced plant susceptibility to necrotrophic
fungal pathogens (Llorente et al. 2005 ; Trusov et al. 2006 , 2007 ). These results
suggest that the G
γ
α
subunit is a negative regulator, while G
β
and G
γ
subunits are
positive regulators of disease resistance.
Besides the three subunits of the heterotrimeric G-proteins, Arabidopsis has three
unique G
-like proteins, known as Extra Large G-protein 1 (XLG1), XLG2, and
XLG3 (Zhu et al. 2009 ). XLG2 and XLG 3 were rapidly induced by infection with the
bacterial pathogen Pseudomonas syringae , whereas the XLG1 transcript level was not
affected by the pathogen infection. The xlg2 loss-of-function mutation caused
enhanced susceptibility to P . syringae (Zhu et al. 2009 ). The xlg2 mutation affected
pathogen-triggered induction of a small set of defense-related genes (Zhu et al. 2009 ).
Constitutive overexpression of XLG2 leads to the accumulation of transcripts from
multiple defense-related genes. These results suggest that XLG2 is involved in trig-
gering immune responses. In contrast, xlg1 and xlg3 mutants showed no difference
from wild-type in resistance to P . syringae , suggesting that XLG1 and XLG3 are not
involved in triggering defense responses against the pathogen (Zhu et al. 2009 ).
AGB1, the sole G
α
subunit in Arabidopsis has been found to be a positive regula-
tor in resistance against necrotrophic fungal pathogens. The agb1 mutant impaired
in the G
β
subunit shows enhanced susceptibility to these pathogens (Llorente et al.
2005 ; Trusov et al. 2006 , 2009 ). The G
β
β
subunit forms an obligate dimer with either
one of the Arabidopsis G
2/AGG2). The agg1 agg2 dou-
ble mutant is as susceptible as agb1 plants to Plectosphaerella cucumerina (Delgado-
Cerezo et al. 2012 ). This heteromeric G-protein-mediated resistance was found to
be independent of SA-, JA-, ethylene-, and abscisic acid-mediated signaling path-
ways. However, this G-protein-mediated resistance was found to be modulated by
cell wall defense responses. The xylose content was lower in agb1 and agg1 agg2
mutants than in wild-type plants, suggesting that cell wall modifi cations may be the
immune response triggered by G
γ
subunits (
γ
1/AGG1 and
γ
β
and G
γ
subunits of the heteromeric G-proteins
(Delgado-Cerezo et al. 2012 ).
The rice dl mutant, which is defi cient in the G
subunit, exhibited a reduced
hypersensitive response after infection by the rice blast pathogen Magnaporthe ory-
zae (formerly known as M. grisea ) (Suharsono et al. 2002 , Iwata et al. 2002 ) and
bacterial blight pathogen Xanthomonas oryzae pv. oryzae (Komatsu et al. 2004 ).
These results suggest that the heteromeric G
α
subunit may play an important role in
both fungal and bacterial disease resistance (Izawa et al. 2010 ).
α
3.6
Small G-Proteins Activate Plant Innate Immunity
The small G-proteins have been reported to play important role in activating immune
responses against fungal, oomycete, bacterial, and viral diseases in different plants
(Sano and Ohashi 1995 ; Ono et al. 2001 ; Suharsono et al. 2002 ; Shirasu and
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