Paper · 2026 · Pest Management Science

Harpin Ea protein confers top rot resistance in Rosa roxburghii Tratt. through synergistic activation of structural defenses and the phenylpropanoid–flavonoid pathway

Tingting Wei; Qiong Wu; Handinghong Zheng; et al.

12
claims
31
verbatim quotes
0
citations
5
research areas
—
Not open access

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

Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

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 fructicolaColletotrichum 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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗
Colletotrichum fructicola
Colletotrichum fructicola · InteractorMa X, Nontachaiyapoom S, Jayawardena RS, Hyde KD, Gentekaki E, Zhou S, Qian Y, Wen T, Kang J (2018) Endophytic Colletotrichum species from Dendrobium spp. in Ch · CC BY 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Harpin Ea protein under field conditions significantly reduced top rot incidence, with >75% control efficacy over consecutive growing seasons”

✓ located verbatim in the source · abstract, characters 350–491 · show in text ↑
t0

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
e1

“top rot disease caused by Colletotrichum fructicola”

✓ located verbatim in the source · abstract, characters 119–170 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

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%”

✓ located verbatim in the source · abstract, characters 350–525 · show in text ↑
t0

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“increased fruit yield by >9% without negatively affecting quality”

✓ located verbatim in the source · abstract, characters 497–562 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Harpin Ea exhibited negligible direct antifungal activity in vitro, with 3.39-8.25% inhibition.

Taxon
Colletotrichum fructicolaColletotrichum 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.

Colletotrichum fructicola
Colletotrichum fructicola · TaxonMa X, Nontachaiyapoom S, Jayawardena RS, Hyde KD, Gentekaki E, Zhou S, Qian Y, Wen T, Kang J (2018) Endophytic Colletotrichum species from Dendrobium spp. in Ch · CC BY 4.0 · Wikimedia Commons ↗

Verbatim

e0

“despite negligible direct antifungal activity in vitro (3.39-8.25% inhibition)”

✓ located verbatim in the source · abstract, characters 564–642 · show in text ↑
e1

“top rot disease caused by Colletotrichum fructicola”

✓ located verbatim in the source · abstract, characters 119–170 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“The elicitor enhanced antioxidant enzyme activities”

✓ located verbatim in the source · abstract, characters 644–695 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“elevated levels of defense-related compounds”

✓ located verbatim in the source · abstract, characters 697–741 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“improved photosynthetic performance”

✓ located verbatim in the source · abstract, characters 746–781 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Harpin Ea protein reinforced the fruit epidermis and cell walls, increasing key structural components (cellulose, lignin, carbohydrates) by 102-200%”

✓ located verbatim in the source · abstract, characters 813–961 · show in text ↑
t0

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

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)”

✓ located verbatim in the source · abstract, characters 1033–1274 · show in text ↑
t0

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“promoting the accumulation of critical defense-related metabolites including p-cinnamic acid, caffeic acid, phenylalanine”

✓ located verbatim in the source · abstract, characters 1280–1401 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“promoting the accumulation of critical defense-related metabolites including p-cinnamic acid, caffeic acid, phenylalanine, naringenin, quercetin and hesperidin”

✓ located verbatim in the source · abstract, characters 1280–1439 · show in text ↑
t0

“Harpin Ea protein”

✓ located verbatim in the source · title, characters 0–17 · show in text ↑
t1

“Rosa roxburghii Tratt.”

✓ located verbatim in the source · title, characters 48–70 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Harpin-induced resistance is mediated by synergistic physical barrier enhancement and activation of phenylpropanoid-flavonoid metabolism.

Taxon
R. roxburghiiRosa 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.

Rosa roxburghii
Rosa roxburghii · TaxonAgnieszka Kwiecień, Nova · CC BY-SA 4.0 · Wikimedia Commons ↗

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.”

✓ located verbatim in the source · abstract, characters 1453–1625 · show in text ↑