Paper · 2025 · Plant Cell & Environment

Trichoderma afroharzianum T22 Induces Rhizobia and Flavonoid‐Driven Symbiosis to Promote Tolerance to Alkaline Stress in Garden Pea

Asha Thapa; Md Rokibul Hasan; Ahmad H. Kabir

17
claims
34
verbatim quotes
13
citations
4
research areas
✓
Open access

Abstract

Soil alkalinity is a limiting factor for crops, yet the role of beneficial fungi in mitigating this abiotic stress in garden pea is understudied. In this study, Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner. In alkaline-exposed Sugar Snap, T22 improved growth parameters, consistent with increased tissue mineral content, particularly Fe and Mn, as well as enhanced rhizosphere siderophore levels. The split-root assay demonstrated that the beneficial effects of T22 on alkaline stress mitigation are the result of a whole-plant association rather than localised root-specific effects. RNA-seq analysis showed 575 and 818 differentially expressed genes upregulated and downregulated in the roots inoculated with T22 under alkaline conditions. The upregulated genes were mostly involved in the flavonoid biosynthetic pathway (monooxygenase activity, ammonia-lyase activity, 4-coumarate-CoA ligase), along with genes related to mineral transport and redox homoeostasis. Further, a flavonoid precursor restored plant health even in the absence of T22, confirming the role of microbial symbiosis in mitigating alkaline stress. Interestingly, T22 restored the abundance of rhizobia, particularly Rhizobium leguminosarum and Rhizobium indicum, along with the induction of NifA, NifD, and NifH in nodules, suggesting a connection between T22 and rhizobia under soil alkalinity. Further, the elevated rhizosphere siderophore, root flavonoid, expression of PsCoA (4-coumarate-CoA ligase) as well as the relative abundance of TaAOX1 and R. leguminosarum diminished when T22 was substituted with exogenous Fe. This suggests that exogenous Fe eliminates the need for microbiome-driven mineral mobilisation, while T22-mediated alkaline stress mitigation depends on flavonoid-driven symbiosis and R. leguminosarum abundance. It was further supported by the positive interaction of T22 on R. leguminosarum growth in alkaline media. Thus, the beneficial effect of T22 on rhizobia likely stems from their interactions, not solely from the improved mineral status, particularly Fe, in plants. This study provides the first mechanistic insights into T22 interactions with host and rhizobia, advancing microbiome strategies to alleviate soil alkalinity in peas and other legumes.

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Claims 17

Observationobservedoriginal resultmeasured

Claim · model-written summary

T22 colonised the roots of garden pea cultivars under soil alkalinity in a host-specific manner.

Taxon
garden peaLathyrus oleraceuse1
Interactor
Trichoderma afroharzianum T22Trichoderma afroharzianume1symbiont
Outcome
colonised the rootse1
Qualifier
in a host-specific mannere1

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

T22 improved growth parameters of alkaline-exposed Sugar Snap.

Taxon
garden peaLathyrus oleraceuse1
Material
Sugar Snape2
Treatment
T22e2
Outcome
growth parameterse2
Direction
improvede2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“In alkaline-exposed Sugar Snap, T22 improved growth parameters, consistent with increased tissue mineral content, particularly Fe and Mn, as well as enhanced rhizosphere siderophore levels”

✓ located verbatim in the source · abstract, characters 289–477 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

T22 increased tissue mineral content in alkaline-exposed Sugar Snap, particularly Fe and Mn.

Taxon
garden peaLathyrus oleraceuse1
Material
Sugar Snape2
Treatment
T22e2
Outcome
tissue mineral contente2
Direction
increasede2
Qualifier
particularly Fe and Mne2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“In alkaline-exposed Sugar Snap, T22 improved growth parameters, consistent with increased tissue mineral content, particularly Fe and Mn, as well as enhanced rhizosphere siderophore levels”

✓ located verbatim in the source · abstract, characters 289–477 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

T22 enhanced rhizosphere siderophore levels in alkaline-exposed Sugar Snap.

Taxon
garden peaLathyrus oleraceuse1
Material
Sugar Snape2
Treatment
T22e2
Outcome
rhizosphere siderophore levelse2
Direction
enhancede2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“In alkaline-exposed Sugar Snap, T22 improved growth parameters, consistent with increased tissue mineral content, particularly Fe and Mn, as well as enhanced rhizosphere siderophore levels”

✓ located verbatim in the source · abstract, characters 289–477 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

The beneficial effect of T22 on alkaline stress mitigation reflects a whole-plant association rather than localised root-specific effects.

Taxon
garden peaLathyrus oleraceuse1
Treatment
T22e3
Outcome
alkaline stress mitigatione3
Qualifier
whole-plant association rather than localised root-specific effectse3

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e3

“The split-root assay demonstrated that the beneficial effects of T22 on alkaline stress mitigation are the result of a whole-plant association rather than localised root-specific effects”

✓ located verbatim in the source · abstract, characters 479–665 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Under alkaline conditions, 575 genes were upregulated in roots inoculated with T22.

Taxon
garden peaLathyrus oleraceuse1
Treatment
roots inoculated with T22e4
Outcome
differentially expressed genese4
Direction
upregulatede4
Value
575e4

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e4

“575 and 818 differentially expressed genes upregulated and downregulated in the roots inoculated with T22 under alkaline conditions”

✓ located verbatim in the source · abstract, characters 691–822 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Under alkaline conditions, 818 genes were downregulated in roots inoculated with T22.

Taxon
garden peaLathyrus oleraceuse1
Treatment
roots inoculated with T22e4
Outcome
differentially expressed genese4
Direction
downregulatede4
Value
818e4

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e4

“575 and 818 differentially expressed genes upregulated and downregulated in the roots inoculated with T22 under alkaline conditions”

✓ located verbatim in the source · abstract, characters 691–822 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Upregulated genes under T22 were mostly involved in the flavonoid biosynthetic pathway.

Taxon
garden peaLathyrus oleraceuse1
Gene
4-coumarate-CoA ligasee5
Outcome
involved in the flavonoid biosynthetic pathwaye5

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e5

“The upregulated genes were mostly involved in the flavonoid biosynthetic pathway (monooxygenase activity, ammonia-lyase activity, 4-coumarate-CoA ligase)”

✓ located verbatim in the source · abstract, characters 824–977 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Genes related to mineral transport and redox homoeostasis were also among the T22-regulated genes.

Taxon
garden peaLathyrus oleraceuse1
Outcome
genes related to mineral transport and redox homoeostasise6

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e6

“along with genes related to mineral transport and redox homoeostasis”

✓ located verbatim in the source · abstract, characters 979–1047 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

A flavonoid precursor restored plant health without T22.

Taxon
garden peaLathyrus oleraceuse1
Treatment
flavonoid precursore7
Outcome
plant healthe7
Direction
restorede7
Comparator
absence of T22e7

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e7

“a flavonoid precursor restored plant health even in the absence of T22”

✓ located verbatim in the source · abstract, characters 1058–1128 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

T22 restored the abundance of rhizobia, particularly R. leguminosarum and R. indicum.

Taxon
garden peaLathyrus oleraceuse1
Treatment
T22e2
Interactor
rhizobiae2symbiont
Outcome
abundance of rhizobiae2
Direction
restorede2
Qualifier
particularly Rhizobium leguminosarum and Rhizobium indicume2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“T22 restored the abundance of rhizobia, particularly Rhizobium leguminosarum and Rhizobium indicum”

✓ located verbatim in the source · abstract, characters 1219–1317 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

T22 induced NifA, NifD and NifH expression in nodules.

Taxon
garden peaLathyrus oleraceuse1
Treatment
T22e2
Gene
NifA, NifD, and NifHe3
Outcome
NifA, NifD, and NifHe3
Direction
inductione3

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“T22 restored the abundance of rhizobia, particularly Rhizobium leguminosarum and Rhizobium indicum”

✓ located verbatim in the source · abstract, characters 1219–1317 · show in text ↑
e3

“the induction of NifA, NifD, and NifH in nodules”

✓ located verbatim in the source · abstract, characters 1330–1378 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Exogenous Fe lowered rhizosphere siderophore, root flavonoid and PsCoA expression relative to T22.

Taxon
garden peaLathyrus oleraceuse1
Treatment
exogenous Fee2
Gene
PsCoAe2
Outcome
rhizosphere siderophore, root flavonoid, expression of PsCoAe2
Direction
diminishede2
Comparator
T22e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“the elevated rhizosphere siderophore, root flavonoid, expression of PsCoA (4-coumarate-CoA ligase) as well as the relative abundance of TaAOX1 and R. leguminosarum diminished when T22 was substituted with exogenous Fe”

✓ located verbatim in the source · abstract, characters 1461–1678 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Exogenous Fe lowered the relative abundance of TaAOX1 and R. leguminosarum relative to T22.

Taxon
garden peaLathyrus oleraceuse1
Treatment
exogenous Fee2
Interactor
R. leguminosarumRhizobium leguminosarume2symbiont
Gene
TaAOX1e2
Outcome
relative abundance of TaAOX1 and R. leguminosarume2
Direction
diminishede2
Comparator
T22e2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗
Rhizobium leguminosarum
Rhizobium leguminosarum · Interactor · picture from its Wikimedia Commons categoryQaswarah 21 · CC BY 4.0 · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“the elevated rhizosphere siderophore, root flavonoid, expression of PsCoA (4-coumarate-CoA ligase) as well as the relative abundance of TaAOX1 and R. leguminosarum diminished when T22 was substituted with exogenous Fe”

✓ located verbatim in the source · abstract, characters 1461–1678 · show in text ↑
Mechanismauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Exogenous Fe removes the need for microbiome-driven mineral mobilisation.

Taxon
garden peaLathyrus oleraceuse1
Treatment
exogenous Fee2
Outcome
need for microbiome-driven mineral mobilisatione2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“exogenous Fe eliminates the need for microbiome-driven mineral mobilisation”

✓ located verbatim in the source · abstract, characters 1699–1774 · show in text ↑
Mechanismauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

T22-mediated alkaline stress mitigation depends on flavonoid-driven symbiosis and R. leguminosarum abundance.

Taxon
garden peaLathyrus oleraceuse1
Treatment
T22e2
Outcome
alkaline stress mitigatione2
Qualifier
depends on flavonoid-driven symbiosis and R. leguminosarum abundancee2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“T22-mediated alkaline stress mitigation depends on flavonoid-driven symbiosis and R. leguminosarum abundance”

✓ located verbatim in the source · abstract, characters 1782–1890 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

T22 had a positive interaction with R. leguminosarum growth in alkaline media.

Taxon
garden peaLathyrus oleraceuse1
Treatment
T22e2
Interactor
R. leguminosarumRhizobium leguminosarume2symbiont
Outcome
R. leguminosarum growth in alkaline mediae2
Direction
positive interactione2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Lathyrus oleraceus
Lathyrus oleraceus · TaxonRenee Comet (photographer) AV Number: AV-9400-4164 · Public domain · Wikimedia Commons ↗
Rhizobium leguminosarum
Rhizobium leguminosarum · Interactor · picture from its Wikimedia Commons categoryQaswarah 21 · CC BY 4.0 · Wikimedia Commons ↗

Verbatim

e1

“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”

✓ located verbatim in the source · abstract, characters 161–287 · show in text ↑
e2

“the positive interaction of T22 on R. leguminosarum growth in alkaline media”

✓ located verbatim in the source · abstract, characters 1920–1996 · show in text ↑

iron atom · molecular: expression · Lathyrus oleraceus (pea)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 reporting no difference from an alternative

  1. positive
See all 2 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.