Abstract
Plants react to pathogen attack via recognition of, and response to, pathogen-specific molecules at the cell surface and inside the cell. Pathogen effectors (virulence factors) are monitored by intracellular nucleotide-binding leucine-rich repeat (NB-LRR) sensor proteins in plants and mammals. Here, we study the genetic requirements for defense responses of an autoactive mutant of ADR1-L2, an Arabidopsis coiled-coil (CC)-NB-LRR protein. ADR1-L2 functions upstream of salicylic acid (SA) accumulation in several defense contexts, and it can act in this context as a "helper" to transduce specific microbial activation signals from "sensor" NB-LRRs. This helper activity does not require an intact P-loop. ADR1-L2 and another of two closely related members of this small NB-LRR family are also required for propagation of unregulated runaway cell death (rcd) in an lsd1 mutant. We demonstrate here that, in this particular context, ADR1-L2 function is P-loop dependent. We generated an autoactive missense mutation, ADR1-L2D484V, in a small homology motif termed MHD. Expression of ADR1-L2D848V leads to dwarfed plants that exhibit increased disease resistance and constitutively high SA levels. The morphological phenotype also requires an intact P-loop, suggesting that these ADR1-L2D484V phenotypes reflect canonical activation of this NB-LRR protein. We used ADR1-L2D484V to define genetic requirements for signaling. Signaling from ADR1-L2D484V does not require NADPH oxidase and is negatively regulated by EDS1 and AtMC1. Transcriptional regulation of ADR1-L2D484V is correlated with its phenotypic outputs; these outputs are both SA-dependent and -independent. The genetic requirements for ADR1-L2D484V activity resemble those that regulate an SA-gradient-dependent signal amplification of defense and cell death signaling initially observed in the absence of LSD1. Importantly, ADR1-L2D484V autoactivation signaling is controlled by both EDS1 and SA in separable, but linked pathways. These data allows us to propose a genetic model that provides insight into an SA-dependent feedback regulation loop, which, surprisingly, includes ADR1-L2.
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Claims 15
Claim · model-written summary
ADR1-L2 acts upstream of salicylic acid accumulation in defense contexts.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- ADR1-L2e0
- Outcome
- salicylic acid (SA) accumulatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ADR1-L2 functions upstream of salicylic acid (SA) accumulation in several defense contexts”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
ADR1-L2 can act as a helper for microbial activation signals from sensor NB-LRRs.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- NB-LRRse0
- Outcome
- transduce specific microbial activation signalse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“it can act in this context as a "helper" to transduce specific microbial activation signals from "sensor" NB-LRRs”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
Helper activity does not require an intact P-loop.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- P-loope0
- Outcome
- helper activitye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“This helper activity does not require an intact P-loop”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
ADR1-L2 and a related family member are required for runaway cell death in lsd1.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- ADR1-L2e0
- Outcome
- propagation of unregulated runaway cell death (rcd)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ADR1-L2 and another of two closely related members of this small NB-LRR family are also required for propagation of unregulated runaway cell death (rcd) in an lsd1 mutant”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
ADR1-L2 function in runaway cell death requires its P-loop.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- ADR1-L2e0
- Outcome
- ADR1-L2 functione0
- Qualifier
- P-loop dependente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in this particular context, ADR1-L2 function is P-loop dependent”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
Expression of the stated autoactive variant produced dwarfed plants.
- Taxon
- Arabidopsis
Arabidopsise1 - Treatment
- Expression of ADR1-L2D848Ve0
- Outcome
- dwarfed plantse0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Expression of ADR1-L2D848V leads to dwarfed plants”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
Expression of the stated autoactive variant increased disease resistance.
- Taxon
- Arabidopsis
Arabidopsise1 - Treatment
- Expression of ADR1-L2D848Ve0
- Outcome
- disease resistancee0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Expression of ADR1-L2D848V leads to dwarfed plants that exhibit increased disease resistance”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
Expression of the stated autoactive variant caused constitutively high salicylic acid levels.
- Taxon
- Arabidopsis
Arabidopsise1 - Treatment
- Expression of ADR1-L2D848Ve0
- Outcome
- SA levelse0
- Direction
- highe0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Expression of ADR1-L2D848V leads to dwarfed plants that exhibit increased disease resistance and constitutively high SA levels”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
The morphological phenotype requires an intact P-loop.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- P-loope0
- Outcome
- morphological phenotypee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The morphological phenotype also requires an intact P-loop”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
ADR1-L2D484V signaling does not require NADPH oxidase.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- NADPH oxidasee0
- Outcome
- Signaling from ADR1-L2D484Ve0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Signaling from ADR1-L2D484V does not require NADPH oxidase”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
EDS1 negatively regulates ADR1-L2D484V signaling.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- EDS1e0
- Outcome
- Signaling from ADR1-L2D484Ve0
- Direction
- negatively regulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Signaling from ADR1-L2D484V does not require NADPH oxidase and is negatively regulated by EDS1 and AtMC1”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
AtMC1 negatively regulates ADR1-L2D484V signaling.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- AtMC1e0
- Outcome
- Signaling from ADR1-L2D484Ve0
- Direction
- negatively regulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Signaling from ADR1-L2D484V does not require NADPH oxidase and is negatively regulated by EDS1 and AtMC1”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
ADR1-L2D484V transcriptional regulation correlated with phenotypic outputs.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- ADR1-L2D484Ve0
- Outcome
- phenotypic outputse0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Transcriptional regulation of ADR1-L2D484V is correlated with its phenotypic outputs”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
The phenotypic outputs include salicylic acid dependent and independent components.
- Taxon
- Arabidopsis
Arabidopsise1 - Outcome
- outputse0
- Qualifier
- both SA-dependent and -independente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“these outputs are both SA-dependent and -independent”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
Claim · model-written summary
EDS1 and salicylic acid control autoactive signaling through separable linked pathways.
- Taxon
- Arabidopsis
Arabidopsise1 - Gene
- EDS1e0
- Outcome
- ADR1-L2D484V autoactivation signalinge0
- Qualifier
- separable, but linked pathwayse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ADR1-L2D484V autoactivation signaling is controlled by both EDS1 and SA in separable, but linked pathways”
e1“an Arabidopsis coiled-coil (CC)-NB-LRR protein”
The same questions in other papers
transgenic · disease · ArabidopsisPositive and negative results
35 papers in the same direction as this paper · 19 in the opposite direction
- no effect
- negative
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