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
Claim · model-written summary
T22 colonised the roots of garden pea cultivars under soil alkalinity in a host-specific manner.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
Claim · model-written summary
T22 improved growth parameters of alkaline-exposed Sugar Snap.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
Claim · model-written summary
T22 increased tissue mineral content in alkaline-exposed Sugar Snap, particularly Fe and Mn.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
Claim · model-written summary
T22 enhanced rhizosphere siderophore levels in alkaline-exposed Sugar Snap.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
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 pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
Claim · model-written summary
Under alkaline conditions, 575 genes were upregulated in roots inoculated with T22.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e4“575 and 818 differentially expressed genes upregulated and downregulated in the roots inoculated with T22 under alkaline conditions”
Claim · model-written summary
Under alkaline conditions, 818 genes were downregulated in roots inoculated with T22.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e4“575 and 818 differentially expressed genes upregulated and downregulated in the roots inoculated with T22 under alkaline conditions”
Claim · model-written summary
Upregulated genes under T22 were mostly involved in the flavonoid biosynthetic pathway.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e5“The upregulated genes were mostly involved in the flavonoid biosynthetic pathway (monooxygenase activity, ammonia-lyase activity, 4-coumarate-CoA ligase)”
Claim · model-written summary
Genes related to mineral transport and redox homoeostasis were also among the T22-regulated genes.
- Taxon
- garden pea
Lathyrus oleraceuse1 - Outcome
- genes related to mineral transport and redox homoeostasise6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e6“along with genes related to mineral transport and redox homoeostasis”
Claim · model-written summary
A flavonoid precursor restored plant health without T22.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e7“a flavonoid precursor restored plant health even in the absence of T22”
Claim · model-written summary
T22 restored the abundance of rhizobia, particularly R. leguminosarum and R. indicum.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e2“T22 restored the abundance of rhizobia, particularly Rhizobium leguminosarum and Rhizobium indicum”
Claim · model-written summary
T22 induced NifA, NifD and NifH expression in nodules.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e2“T22 restored the abundance of rhizobia, particularly Rhizobium leguminosarum and Rhizobium indicum”
e3“the induction of NifA, NifD, and NifH in nodules”
Claim · model-written summary
Exogenous Fe lowered rhizosphere siderophore, root flavonoid and PsCoA expression relative to T22.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
Claim · model-written summary
Exogenous Fe lowered the relative abundance of TaAOX1 and R. leguminosarum relative to T22.
- Taxon
- garden pea
Lathyrus oleraceuse1 - Treatment
- exogenous Fee2
- Interactor
- R. leguminosarum
Rhizobium 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.


Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
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”
Claim · model-written summary
Exogenous Fe removes the need for microbiome-driven mineral mobilisation.
- Taxon
- garden pea
Lathyrus oleraceuse1 - Treatment
- exogenous Fee2
- Outcome
- need for microbiome-driven mineral mobilisatione2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e2“exogenous Fe eliminates the need for microbiome-driven mineral mobilisation”
Claim · model-written summary
T22-mediated alkaline stress mitigation depends on flavonoid-driven symbiosis and R. leguminosarum abundance.
- Taxon
- garden pea
Lathyrus 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.

Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e2“T22-mediated alkaline stress mitigation depends on flavonoid-driven symbiosis and R. leguminosarum abundance”
Claim · model-written summary
T22 had a positive interaction with R. leguminosarum growth in alkaline media.
- Taxon
- garden pea
Lathyrus oleraceuse1 - Treatment
- T22e2
- Interactor
- R. leguminosarum
Rhizobium 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.


Verbatim
e1“Trichoderma afroharzianum T22 colonised the roots of garden pea cultivars exposed to soil alkalinity in a host-specific manner”
e2“the positive interaction of T22 on R. leguminosarum growth in alkaline media”
The same questions in other papers
iron atom · molecular: expression · Lathyrus oleraceus (pea)Positive and negative results
1 reporting no difference from an alternative
- positive
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.