Abstract
In compatible interactions, biotrophic microbial phytopathogens rely on the supply of assimilates by the colonized host tissue. In rice, phloem localized SWEET sucrose transporters can be reprogrammed by bacterial effectors to establish compatibility. We observed that sweet11/sweet12 double mutants, but not single mutants, exhibited increased resistance toward the fungal hemibiotroph Colletotrichum higginsianum (Ch), both in the biotrophic and the necrotrophic colonization phase. We therefore investigated if the phloem localized transporters AtSWEET11 and AtSWEET12 represent additive susceptibility factors in the interaction of Arabidopsis with Ch. AtSWEET12-YFP fusion protein driven by the endogenous promoter strongly accumulated at Ch infection sites and in the vasculature upon challenge with Ch. However, susceptibility of sweet12 single mutants to Ch was comparable to wild type, indicating that the accumulation of AtSWEET12 at Ch infection sites does not play a major role for compatibility. AtSWEET12-YFP reporter protein was not detectable at the plant pathogen interface, suggesting that AtSWEET12 is not targeted by Ch effectors. AtSWEET11-YFP accumulation in pAtSWEET11:AtSWEET11-YFP plants was similar in Ch infected and mock control leaves. A close inspection of major carbohydrate metabolism in non-infected control plants revealed that soluble sugar and starch content were substantially elevated in sweet11/sweet12 double mutants during the entire diurnal cycle, that diurnal soluble sugar turnover was increased more than 2-fold in sweet11/sweet12, and that accumulation of free hexoses and sucrose was strongly expedited in double mutant leaves compared to wild type and both single mutants during the course of Ch infection. After 2 days of treatment, free and conjugated SA levels were significantly increased in infected and mock control leaves of sweet11/sweet12 relative to all other genotypes, respectively. Induced genes in mock treated sweet11/sweet12 leaves were highly significantly enriched for several GO terms associated with SA signaling and response compared to mock treated wild type leaves, indicating sugar-mediated priming of the SA pathway in the double mutant. Infection assays with salicylic acid deficient sweet11/sweet12/sid2 triple mutants demonstrated that reduced susceptibility observed in sweet11/sweet12 was entirely dependent on the SA pathway. We suggest a model how defects in phloem loading of sucrose can influence SA priming and hence, compatibility.
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Claims 15
Claim · model-written summary
The sweet11/sweet12 double mutants, but not the single mutants, showed increased resistance to Colletotrichum higginsianum in both biotrophic and necrotrophic phases.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12 double mutantse1
- Interactor
- Colletotrichum higginsianumColletotrichum higginsianume1pathogen
- Outcome
- resistancee1
- Direction
- increasede1
- Qualifier
- both in the biotrophic and the necrotrophic colonization phasee1
- Comparator
- single mutantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“sweet11/sweet12 double mutants, but not single mutants, exhibited increased resistance toward the fungal hemibiotroph Colletotrichum higginsianum (Ch), both in the biotrophic and the necrotrophic colonization phase”
Claim · model-written summary
Susceptibility of sweet12 single mutants to Ch was comparable to wild type.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet12 single mutantse2
- Interactor
- Che2pathogen
- Outcome
- susceptibility of sweet12 single mutants to Che2
- Direction
- comparablee2
- Comparator
- wild typee2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e2“susceptibility of sweet12 single mutants to Ch was comparable to wild type”
Claim · model-written summary
Accumulation of AtSWEET12 at Ch infection sites does not play a major role for compatibility.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- AtSWEET12e3
- Outcome
- compatibilitye3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e3“However, susceptibility of sweet12 single mutants to Ch was comparable to wild type, indicating that the accumulation of AtSWEET12 at Ch infection sites does not play a major role for compatibility”
Claim · model-written summary
AtSWEET12-YFP accumulated at Ch infection sites and in the vasculature upon Ch challenge.
- Taxon
- Arabidopsis
Arabidopsise0 - Interactor
- Che4pathogen
- Gene
- AtSWEET12-YFPe4
- Outcome
- accumulated at Ch infection sites and in the vasculaturee4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e4“AtSWEET12-YFP fusion protein driven by the endogenous promoter strongly accumulated at Ch infection sites and in the vasculature upon challenge with Ch”
Claim · model-written summary
AtSWEET12-YFP reporter protein was not detectable at the plant-pathogen interface.
- Taxon
- Arabidopsis
Arabidopsise0 - Gene
- AtSWEET12-YFPe5
- Outcome
- not detectable at the plant pathogen interfacee5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e5“AtSWEET12-YFP reporter protein was not detectable at the plant pathogen interface, suggesting that AtSWEET12 is not targeted by Ch effectors”
Claim · model-written summary
AtSWEET12 is suggested not to be targeted by Ch effectors.
- Taxon
- Arabidopsis
Arabidopsise0 - Interactor
- Che5pathogen
- Gene
- AtSWEET12e5
- Outcome
- not targeted by Ch effectorse5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e5“AtSWEET12-YFP reporter protein was not detectable at the plant pathogen interface, suggesting that AtSWEET12 is not targeted by Ch effectors”
Claim · model-written summary
AtSWEET11-YFP accumulation was similar in Ch infected and mock control leaves.
- Taxon
- Arabidopsis
Arabidopsise0 - Treatment
- Ch infectede6
- Interactor
- Che6pathogen
- Gene
- AtSWEET11-YFPe6
- Outcome
- AtSWEET11-YFP accumulatione6
- Direction
- similare6
- Comparator
- mock control leavese6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e6“AtSWEET11-YFP accumulation in pAtSWEET11:AtSWEET11-YFP plants was similar in Ch infected and mock control leaves”
Claim · model-written summary
Soluble sugar and starch content were substantially elevated in sweet11/sweet12 double mutants throughout the diurnal cycle.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12 double mutantse7
- Organ / stage
- Leavest0
- Outcome
- soluble sugar and starch contente7
- Direction
- elevatede7
- Qualifier
- during the entire diurnal cyclee7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e7“soluble sugar and starch content were substantially elevated in sweet11/sweet12 double mutants during the entire diurnal cycle”
t0“Sugar Accumulation in Leaves of Arabidopsis sweet11/sweet12 Double Mutants”
Claim · model-written summary
Diurnal soluble sugar turnover increased more than 2-fold in sweet11/sweet12.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12e1
- Outcome
- diurnal soluble sugar turnovere1
- Direction
- increasede1
- Value
- more than 2-folde1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“diurnal soluble sugar turnover was increased more than 2-fold in sweet11/sweet12”
Claim · model-written summary
Free hexoses and sucrose accumulated more strongly in double mutant leaves than in wild type and single mutants during Ch infection.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- double mutante1
- Organ / stage
- leavese1
- Treatment
- Ch infectione1
- Interactor
- Che1pathogen
- Outcome
- accumulation of free hexoses and sucrosee1
- Direction
- strongly expeditede1
- Comparator
- wild type and both single mutantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“accumulation of free hexoses and sucrose was strongly expedited in double mutant leaves compared to wild type and both single mutants during the course of Ch infection”
Claim · model-written summary
Free and conjugated SA levels were significantly increased in sweet11/sweet12 relative to all other genotypes.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12e1
- Outcome
- free and conjugated SA levelse1
- Direction
- increasede1
- Qualifier
- After 2 days of treatmente1
- Comparator
- all other genotypese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“After 2 days of treatment, free and conjugated SA levels were significantly increased in infected and mock control leaves of sweet11/sweet12 relative to all other genotypes”
Claim · model-written summary
Induced genes in mock-treated sweet11/sweet12 leaves were highly significantly enriched for GO terms associated with SA signalling and response.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12e1
- Organ / stage
- leavese1
- Outcome
- enriched for several GO terms associated with SA signaling and responsee1
- Direction
- enrichede1
- Comparator
- mock treated wild type leavese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“Induced genes in mock treated sweet11/sweet12 leaves were highly significantly enriched for several GO terms associated with SA signaling and response compared to mock treated wild type leaves”
Claim · model-written summary
Sugar-mediated priming of the SA pathway occurs in the double mutant.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- double mutante1
- Outcome
- priming of the SA pathwaye1
- Qualifier
- sugar-mediatede1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“indicating sugar-mediated priming of the SA pathway in the double mutant”
Claim · model-written summary
Reduced susceptibility of sweet11/sweet12 is entirely dependent on the SA pathway, shown with SA-deficient sweet11/sweet12/sid2 triple mutants.
- Taxon
- Arabidopsis
Arabidopsise0 - Material
- sweet11/sweet12/sid2 triple mutantse1
- Outcome
- reduced susceptibilitye1
- Qualifier
- entirely dependent on the SA pathwaye1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“Infection assays with salicylic acid deficient sweet11/sweet12/sid2 triple mutants demonstrated that reduced susceptibility observed in sweet11/sweet12 was entirely dependent on the SA pathway”
Claim · model-written summary
Defects in phloem loading of sucrose may influence SA priming and hence compatibility.
- Taxon
- Arabidopsis
Arabidopsise0 - Outcome
- SA priming and hence, compatibilitye1
- Qualifier
- defects in phloem loading of sucrosee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“in the interaction of Arabidopsis with Ch”
e1“We suggest a model how defects in phloem loading of sucrose can influence SA priming and hence, compatibility”
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