Abstract
BACKGROUND: Rosa roxburghii Tratt. is an economically important fruit crop whose production is severely constrained by top rot disease caused by Colletotrichum fructicola. Although protein elicitors such as Harpin Ea represent a promising sustainable alternative to chemical fungicides, their mode-of-action in this crop is largely unknown. RESULTS: Harpin Ea protein under field conditions significantly reduced top rot incidence, with >75% control efficacy over consecutive growing seasons, and increased fruit yield by >9% without negatively affecting quality, despite negligible direct antifungal activity in vitro (3.39-8.25% inhibition). The elicitor enhanced antioxidant enzyme activities, elevated levels of defense-related compounds and improved photosynthetic performance. Structural analyses show that Harpin Ea protein reinforced the fruit epidermis and cell walls, increasing key structural components (cellulose, lignin, carbohydrates) by 102-200%. Integrated transcriptomic and metabolomic profiling demonstrates that Harpin Ea coordinately upregulated the phenylpropanoid-flavonoid biosynthetic pathway, enhancing the expression of genes encoding key enzymes such as phenylalanine ammonia-lyase (PAL), 4-coumarate-CoA ligase (4CL) and chalcone synthase (CHS), and promoting the accumulation of critical defense-related metabolites including p-cinnamic acid, caffeic acid, phenylalanine, naringenin, quercetin and hesperidin. CONCLUSION: Harpin Ea-induced resistance in R. roxburghii is mediated by a synergistic enhancement of physical barriers and systemic activation of phenylpropanoid-flavonoid metabolism. This work provides the first mechanistic evidence linking this dual-layer defense to Harpin Ea-induced disease resistance and establishes a molecular basis for the sustainable application of protein elicitors in fruit crop protection. © 2026 Society of Chemical Industry.
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Claims 12
Claim · model-written summary
Harpin Ea significantly reduced top rot incidence with over 75% control across consecutive seasons.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit0 - Treatment
- Harpin Ea proteine0
- Interactor
- Colletotrichum fructicola
Colletotrichum fructicolae1pathogen - Outcome
- top rot incidencee0
- Direction
- reducede0
- Value
- >75%e0
- Qualifier
- over consecutive growing seasonse0
- Setting
- field conditionse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

_Figure_8.jpg?width=500)
Verbatim
e0“Harpin Ea protein under field conditions significantly reduced top rot incidence, with >75% control efficacy over consecutive growing seasons”
t0“Rosa roxburghii Tratt.”
e1“top rot disease caused by Colletotrichum fructicola”
Claim · model-written summary
Harpin Ea increased fruit yield by over 9%.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit0 - Treatment
- Harpin Ea proteine0
- Outcome
- fruit yielde0
- Direction
- increasede0
- Value
- >9%e0
- Setting
- field conditionse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Harpin Ea protein under field conditions significantly reduced top rot incidence, with >75% control efficacy over consecutive growing seasons, and increased fruit yield by >9%”
t0“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea did not negatively affect fruit quality.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- qualitye0
- Direction
- without negatively affectinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“increased fruit yield by >9% without negatively affecting quality”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea exhibited negligible direct antifungal activity in vitro, with 3.39-8.25% inhibition.
- Taxon
- Colletotrichum fructicola
Colletotrichum fructicolae1 - Treatment
- Harpin Ea proteint0
- Outcome
- antifungal activitye0
- Value
- 3.39-8.25% inhibitione0
- Setting
- in vitroe0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
_Figure_8.jpg?width=500)
Verbatim
e0“despite negligible direct antifungal activity in vitro (3.39-8.25% inhibition)”
e1“top rot disease caused by Colletotrichum fructicola”
t0“Harpin Ea protein”
Claim · model-written summary
Harpin Ea enhanced antioxidant enzyme activities.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- antioxidant enzyme activitiese0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The elicitor enhanced antioxidant enzyme activities”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea elevated defense-related compound levels.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- levels of defense-related compoundse0
- Direction
- elevatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“elevated levels of defense-related compounds”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea improved photosynthetic performance.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- photosynthetic performancee0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“improved photosynthetic performance”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea reinforced fruit epidermis and cell walls and increased cellulose, lignin and carbohydrates by 102-200%.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit0 - Organ / stage
- fruit epidermis and cell wallse0
- Treatment
- Harpin Ea proteine0
- Outcome
- key structural components (cellulose, lignin, carbohydrates)e0
- Direction
- increasinge0
- Value
- 102-200%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Harpin Ea protein reinforced the fruit epidermis and cell walls, increasing key structural components (cellulose, lignin, carbohydrates) by 102-200%”
t0“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea upregulated the phenylpropanoid-flavonoid pathway and enhanced expression of PAL, 4CL and CHS genes.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit0 - Treatment
- Harpin Eae0
- Gene
- phenylalanine ammonia-lyase (PAL), 4-coumarate-CoA ligase (4CL) and chalcone synthase (CHS)e0
- Outcome
- expression of genese0
- Direction
- enhancinge0
- Qualifier
- upregulated the phenylpropanoid-flavonoid biosynthetic pathwaye0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Harpin Ea coordinately upregulated the phenylpropanoid-flavonoid biosynthetic pathway, enhancing the expression of genes encoding key enzymes such as phenylalanine ammonia-lyase (PAL), 4-coumarate-CoA ligase (4CL) and chalcone synthase (CHS)”
t0“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea promoted accumulation of p-cinnamic acid, caffeic acid and phenylalanine.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- p-cinnamic acid, caffeic acid, phenylalaninee0
- Direction
- promotinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“promoting the accumulation of critical defense-related metabolites including p-cinnamic acid, caffeic acid, phenylalanine”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin Ea promoted accumulation of naringenin, quercetin and hesperidin.
- Taxon
- Rosa roxburghii Tratt.
Rosa roxburghiit1 - Treatment
- Harpin Ea proteint0
- Outcome
- naringenin, quercetin and hesperidine0
- Direction
- promotinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“promoting the accumulation of critical defense-related metabolites including p-cinnamic acid, caffeic acid, phenylalanine, naringenin, quercetin and hesperidin”
t0“Harpin Ea protein”
t1“Rosa roxburghii Tratt.”
Claim · model-written summary
Harpin-induced resistance is mediated by synergistic physical barrier enhancement and activation of phenylpropanoid-flavonoid metabolism.
- Taxon
- R. roxburghii
Rosa roxburghiie0 - Outcome
- Harpin Ea-induced resistancee0
- Qualifier
- synergistic enhancement of physical barriers and systemic activation of phenylpropanoid-flavonoid metabolisme0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Harpin Ea-induced resistance in R. roxburghii is mediated by a synergistic enhancement of physical barriers and systemic activation of phenylpropanoid-flavonoid metabolism.”