Plant Immune System
The plant innate immunity system is generally initiated via recognition
of conserved pathogen / microbe-associated molecular pattern (PAMP / MAMP)
ligands, including bacterial flagellin or peptidoglycan, and fungal chitin.
These MAMPs are perceived by pattern recognition receptors (PRRs), leading
to activation of a series of immune responses (PTI: pattern-triggered immunity).
PRRs are classified into two types of receptors, receptor-like kinase (RLKs)
and receptor-like proteins (RLP). Both types of PRRs contain ligand-binding
domains and a transmembrane domain, but only RLKs have an intercellular
kinase domain. The well-characterized ligand-binding domains are LRR (leucine-rich
repeat) and LysM (lysine rich motif).
Successful pathogens are able to secrete or directly deliver effector proteins into the host cytoplasm which results in inhibition of host PTI. To overcome PTI inhibition by the effectors, plants have developed intracellular immune receptors of the nucleotide-binding leucine-rich repeat (NB-LRR) protein family, which directly or indirectly recognize the effectors, and activate strong immune responses (ETI: effector-triggered immunity).
Perception of PAMPs with PRRs induces a rapid intracellular activation
of mitogen-activated protein kinase (MAPK) cascades consisting of highly
conserved modules in eukaryotes. Each MAPK cascade is composed of three
consecutively acting protein kinases: a MAPK kinase kinase (MAPKKK), a
MAPK kinase (MAPKK) and a MAPK. The MAPK cascades induce robust immune
responses, however how the MAPK cascades are activated downstream of PRRs
is largely unknown. We are interested in understanding how PRRs and NB-LRR
receptors transmit the immune signals to the downstream components to induce
PTI and ETI, respectively, and how the pathogen effectors inhibit host
immune response.
Current Projects
1)Elucidation of molecular mechanism of pattern-triggered immunity in plants.
PRRs recognize PAMPs at plasma membrane, which triggers the intracellular immune response. However, the molecular mechanisms remain to be identified. Our research focuses on immune signaling induced by recognition of fungal chitin and bacterial peptidoglycan. Rice CEBiP and LYP4/LYP6 are RLPs with the extracellular LysM domains responsible for recognition of chitin and PGN, respectively. Perception of chitin with CEBiP induces a complex formation with a RLK OsCERK1, which activates the intracellular immune responses.
We have identified that a receptor-like cytoplasmic kinase (RLCK) OsRLCK185
interacts with the cytoplasmic kinase domain of OsCERK1. In addition, chitin
perception triggers OsCERK1-mediated phosphorylation of OsRLCK185. Silencing
of OsRLCK185 suppresses immune responses induced by recognition of chitin
and PGN, indicating that OsRLCK185 plays a key role in OsCERK1-mediated
immune signaling (Yamaguchi et al. Cell Host & Microbe 2013: the paper
was selected as the featured article).
We also identified PBL27, Arabidopsis homolog of OsRLCK185. PBL27 interacts
with CERK1 at plasma membrane, and is phosphorylated by CERK1 in response
to chitin. Our data indicate the chitin-signaling pathways are conserved
in rice and Arabidopsis (Fig. 2). These results were obtained with collaboration
of Dr. Shibuya group (Meiji Univ) (Shinya and Yamaguchi et al. Plant J.
2014). Currently, we are analyzing how CERK1 activates PBL27 and how PBL27
induces intracellular immune responses.
2)Understanding the inhibitory mechanisms by pathogen effectors
Pathogen effectors are secreted into host cells and inhibit the immune
responses. In fact, expression of Xanthomonas oryzae pv. oryzae (Xoo) effectors
in rice cells results in inhibition of PTI, suggesting that the effectors
target important immune factors. We have identified several immune factors
interacted with the effectors by yeast two hybrid screening. We are currently
analyzing the functions of these factors in plant immune response, and
inhibitory mechanism of host immunity by the pathogen effectors (Fig. 3).
We have identified a rice receptor-like cytoplasmic kinase (OsRLCK185) targeted by Xoo effector XopY (Xoo1488). Our research indicates that XopY suppresses chitin-induced immunity by inhibiting the phosphorylation of OsRLCK185 by OsCERK1 (Yamaguchi et al. Cell Host & Microbe 2013).
We also found that a Xoo effector, XopP (Xoo3222) targets OsPUB44, one
of rice U-box type ubiquitin E3 ligases. OsPUB44 functions as a positive
regulator in chitin- and PGN-induced immunity. The U-box domain of OsPUB44
possesses the ubiquitin ligase activity. XopP interacts with the U-box
domain, which results in suppression of the OsPUB44 enzymatic activity.
These data indicate that XopP modulates host ubiquitination system to inhibit
the immunity (Ishikawa et al. Nature Communications 2014).
3)Elucidation of molecular mechanism of effector-triggered immunity in
plants
The intracellular NB-LRR receptors recognize pathogen effectors in the
cells, and induce immune responses often accompanied with hyper-sensitive
cell death. Rice Xa1 is one of rice NB-LRR receptors (Yoshimura et al.
Proc. Natl. Acad. Sci. USA 1998), which possibly recognizes directly or
indirectly Xoo AvrXa1. However, AvrXa1 has not been identified yet. We
are analyzing Xa1-mediated pathogen recognition and immune signaling by
isolation of rice factors interacted with Xa1.
4)Understanding of MAPK activation in plant immunity
The MAPK cascades are activated by PAMP perception with PRRs. However, it is unknown how PRRs transmit immune signals to MAPK cascades. As mentioned above, OsRLCK185 and PBL27 regulate chitin-induced MAPK activation. Recently, we isolated MAPKKKs interacted with OsRLCK185 and PBL27. We are studying the molecular mechanisms of RLCK-mediated MAPK activation in plants.
5) Screening of chemicals that activate host immunity or inhibit the virulence of pathogens
Based upon our data, we developed the screening system to identify new pesticides that induce host immunity or inhibit the proliferation of the pathogens. The use of these chemicals likely contributes to the environmental conservation.
6) New technology to improve plant disease resistance
During the past several decades, plant Immune research has identified
many important genes regulating host immunity from model plants such as
Arabidopsis and rice. Recently, the genes were used to develop broad-spectrum
and durable resistance to pathogens. This type of research is termed as
‘translational research’.
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