Abstract
• Cytokinin (CK) has been reported to prime plant immunity, however, previous research on CK hormopriming has largely focused on classical synthetic cytokinins like kinetin or 6-benzylaminopurine (BAP). Here, we asked whether synthetic aromatic CK derivatives could offer enhanced efficacy in priming immunity and inhibiting pathogens due to their unique chemical structure. • Our work investigated how CK arabinosides compare to classical CKs in terms of immunity priming efficacy and undesirable effects on plant growth, uncovering the underlying molecular mechanisms and signalling pathways involved in CK arabinoside-mediated immunity in plants and their inhibitory effects on fungi. • Our findings show that CK arabinosides are substantially more effective at promoting immunity and reducing disease caused by the necrotrophic fungal pathogen Botrytis cinerea and the hemibiotrophic bacterium Xanthomonas euvesicatoria in tomato compared to the classical CK 6-BAP. • Crucially, unlike classical CKs, CK arabinosides achieve these benefits without strongly activating canonical cytokinin signalling pathways in plants. Thus, they possess immunity-promoting and fungal inhibitory activities with minimal unwanted effects on plant growth and development. • On the mechanistic level, our work reveals that CK arabinoside-mediated immunity in tomato primarily operates via a salicylic acid (SA)-dependent mechanism and is generally jasmonic acid (JA)-independent; and that CK arabinosides alter sugar metabolism and virulence pathways in B. cinerea . • Importantly, unlike classical CKs, CK arabinosides do not reverse growth inhibition under sugar-limited conditions in fungi, contributing to their stronger antifungal effects and distinguishing their mechanism of action from classical CKs. Cytokinins (CKs) regulate numerous biological processes in plants, and are involved in plant growth and development as well as responses to biotic and abiotic stresses. Although CKs have been reported to prime plant immunity, thus far, research on CK hormopriming has focused on classical synthetic CKs such as kinetin or 6-benzylaminopurine (BAP). Here, we show that synthetic aromatic CK derivatives—specifically, 3-methoxy-6-benzylaminopurine-9-arabinosides—effectively induce immune responses and promote resistance to Botrytis cinerea in tomato ( Solanum lycopersicum ) primarily via a salicylic acid (SA)-dependent mechanism. Additionally, like 6-BAP, these compounds directly inhibit B. cinerea . Transcriptomic analyses of B. cinerea treated with CK arabinosides revealed significant changes in expressed gene profiles, notably in pathways associated with sugar metabolism. Unlike classical CKs, CK arabinosides do not alleviate growth inhibition under sugar-limited conditions, contributing to their stronger antifungal effects. Furthermore, CK arabinosides induce tomato immune responses more robustly than classical CKs, yet without activating canonical cytokinin signaling pathways, likely owing to their exceptional chemical stability which prevents release of the free active CK base, and due to their inability to directly bind CK receptors. Altogether, our findings suggest that CK arabinosides act via a dual mechanism—immune priming and direct pathogen inhibition—and hold promise as next-generation hormopriming agents for sustainable crop protection.
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Claims 9
Claim · model-written summary
Cytokinin arabinosides were substantially more effective than 6-BAP at promoting immunity and reducing Botrytis cinerea disease in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese1
- Interactor
- Botrytis cinerea
Botrytis cinereae1pathogen - Outcome
- reducing diseasee1
- Direction
- more effectivee1
- Comparator
- classical CK 6-BAPe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e1“CK arabinosides are substantially more effective at promoting immunity and reducing disease caused by the necrotrophic fungal pathogen Botrytis cinerea and the hemibiotrophic bacterium Xanthomonas euvesicatoria in tomato compared to the classical CK 6-BAP”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Cytokinin arabinosides induce immunity without strongly activating canonical cytokinin signalling, unlike classical cytokinins.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese2
- Outcome
- canonical cytokinin signalling pathwayse2
- Qualifier
- without strongly activatinge2
- Comparator
- classical CKse2
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e2“Crucially, unlike classical CKs, CK arabinosides achieve these benefits without strongly activating canonical cytokinin signalling pathways in plants”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Cytokinin arabinoside-mediated immunity in tomato operates primarily via a salicylic acid-dependent, jasmonic acid-independent mechanism.
- Taxon
- tomato
Solanum lycopersicume3 - Compound
- salicylic acid (SA)e3
- Outcome
- CK arabinoside-mediated immunitye3
- Qualifier
- generally jasmonic acid (JA)-independente3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e3“CK arabinoside-mediated immunity in tomato primarily operates via a salicylic acid (SA)-dependent mechanism and is generally jasmonic acid (JA)-independent”
Claim · model-written summary
Cytokinin arabinosides alter sugar metabolism and virulence pathways in Botrytis cinerea.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese4
- Interactor
- B. cinerea
Botrytis cinereae4pathogen - Outcome
- sugar metabolism and virulence pathwayse4
- Direction
- altere4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e4“CK arabinosides alter sugar metabolism and virulence pathways in B. cinerea”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Transcriptomic analysis of Botrytis cinerea treated with cytokinin arabinosides showed significant changes in gene expression profiles, notably in sugar metabolism.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese5
- Interactor
- B. cinerea
Botrytis cinereae5pathogen - Outcome
- expressed gene profilese5
- Qualifier
- notably in pathways associated with sugar metabolisme5
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e5“Transcriptomic analyses of B. cinerea treated with CK arabinosides revealed significant changes in expressed gene profiles, notably in pathways associated with sugar metabolism”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Unlike classical cytokinins, cytokinin arabinosides do not alleviate growth inhibition under sugar-limited conditions, which contributes to their stronger antifungal effects.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese6
- Outcome
- growth inhibition under sugar-limited conditionse6
- Direction
- do not alleviatee6
- Comparator
- classical CKse6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
e6“Unlike classical CKs, CK arabinosides do not alleviate growth inhibition under sugar-limited conditions, contributing to their stronger antifungal effects”
Claim · model-written summary
Like 6-BAP, cytokinin arabinosides directly inhibit Botrytis cinerea.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese1
- Interactor
- B. cinerea
Botrytis cinereae7pathogen - Outcome
- B. cinereae7
- Direction
- directly inhibite7
- Comparator
- 6-BAPe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e1“CK arabinosides are substantially more effective at promoting immunity and reducing disease caused by the necrotrophic fungal pathogen Botrytis cinerea and the hemibiotrophic bacterium Xanthomonas euvesicatoria in tomato compared to the classical CK 6-BAP”
e7“Additionally, like 6-BAP, these compounds directly inhibit B. cinerea”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Cytokinin arabinosides induce tomato immune responses more robustly than classical cytokinins.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- CK arabinosidese1
- Outcome
- tomato immune responsese1
- Direction
- more robustlye1
- Comparator
- classical CKse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Furthermore, CK arabinosides induce tomato immune responses more robustly than classical CKs, yet without activating canonical cytokinin signaling pathways”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
Claim · model-written summary
Exceptional chemical stability of cytokinin arabinosides likely prevents release of the free active cytokinin base, explaining the lack of canonical signalling.
- Taxon
- tomato
Solanum lycopersicumt0 - Compound
- free active CK basee2
- Outcome
- without activating canonical cytokinin signaling pathwayse1
- Qualifier
- exceptional chemical stabilitye2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Furthermore, CK arabinosides induce tomato immune responses more robustly than classical CKs, yet without activating canonical cytokinin signaling pathways”
e2“likely owing to their exceptional chemical stability which prevents release of the free active CK base”
t0“priming tomato immunity and inhibiting the fungal phytopathogen B. cinerea”
The same questions in other papers
class · disease · Solanum lycopersicum (tomato)Positive and negative results
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- no effect
- no effect
- positive
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See all 9 papers →class · physiology: hormones · Solanum lycopersicum (tomato)Positive and negative results
2 papers in the same direction as this paper · 6 in the opposite direction
- no effect
- negative
- negative
5 papers more in this group
See all 9 papers →Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.