Abstract
BACKGROUND: Lignin is an important structural component of plant cell wall that confers mechanical strength and tolerance against biotic and abiotic stressors; however it affects the use of biomass such as wheat straw for some industrial applications such as biofuel production. Genetic alteration of lignin quantity and quality has been considered as a viable option to overcome this problem. However, the molecular mechanisms underlying lignin formation in wheat biomass has not been studied. Combining molecular and biochemical approaches, the present study investigated the transcriptional regulation of lignin biosynthesis in two wheat cultivars with varying lodging characteristics and also in response to waterlogging. It also examined the association of lignin level in tissues with that of plant hormones implicated in the control of lignin biosynthesis. RESULTS: Analysis of lignin biosynthesis in the two wheat cultivars revealed a close association of lodging resistance with internode lignin content and expression of 4-coumarate:CoA ligase1 (4CL1), p-coumarate 3-hydroxylase1 (C3H1), cinnamoyl-CoA reductase2 (CCR2), ferulate 5-hydroxylase2 (F5H2) and caffeic acid O-methyltransferase2 (COMT2), which are among the genes highly expressed in wheat tissues, implying the importance of these genes in mediating lignin deposition in wheat stem. Waterlogging of wheat plants reduced internode lignin content, and this effect is accompanied by transcriptional repression of three of the genes characterized as highly expressed in wheat internode including phenylalanine ammonia-lyase6 (PAL6), CCR2 and F5H2, and decreased activity of PAL. Expression of the other genes was, however, induced by waterlogging, suggesting their role in the synthesis of other phenylpropanoid-derived molecules with roles in stress responses. Moreover, difference in internode lignin content between cultivars or change in its level due to waterlogging is associated with the level of cytokinin. CONCLUSION: Lodging resistance, tolerance against biotic and abiotic stresses and feedstock quality of wheat biomass are closely associated with its lignin content. Therefore, the findings of this study provide important insights into the molecular mechanisms underlying lignin formation in wheat, an important step towards the development of molecular tools that can facilitate the breeding of wheat cultivars for optimized lignin content and enhanced feedstock quality without affecting other lignin-related agronomic benefits.
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Claims 9
Claim · model-written summary
Lodging resistance was closely associated with wheat internode lignin content.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Organ / stage
- internodee0
- Outcome
- lodging resistancee0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“close association of lodging resistance with internode lignin content”
Claim · model-written summary
Lodging resistance was associated with expression of 4CL1, C3H1, CCR2, F5H2 and COMT2.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Gene
- 4-coumarate:CoA ligase1 (4CL1), p-coumarate 3-hydroxylase1 (C3H1), cinnamoyl-CoA reductase2 (CCR2), ferulate 5-hydroxylase2 (F5H2) and caffeic acid O-methyltransferase2 (COMT2)e0
- Outcome
- lodging resistancee0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“lodging resistance with internode lignin content and expression of 4-coumarate:CoA ligase1 (4CL1), p-coumarate 3-hydroxylase1 (C3H1), cinnamoyl-CoA reductase2 (CCR2), ferulate 5-hydroxylase2 (F5H2) and caffeic acid O-methyltransferase2 (COMT2)”
Claim · model-written summary
Waterlogging reduced wheat internode lignin content.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Treatment
- Waterlogginge0
- Outcome
- internode lignin contente0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“Waterlogging of wheat plants reduced internode lignin content”
Claim · model-written summary
Waterlogging repressed transcription of PAL6, CCR2 and F5H2 in wheat internodes.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Treatment
- Waterlogginge1
- Gene
- phenylalanine ammonia-lyase6 (PAL6), CCR2 and F5H2e0
- Outcome
- transcriptional repressione0
- Direction
- repressione0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“transcriptional repression of three of the genes characterized as highly expressed in wheat internode including phenylalanine ammonia-lyase6 (PAL6), CCR2 and F5H2”
e1“Waterlogging of wheat plants reduced internode lignin content”
Claim · model-written summary
Waterlogging decreased PAL activity in wheat.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Treatment
- waterlogginge1
- Gene
- PALe0
- Outcome
- activity of PALe0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“decreased activity of PAL”
e1“Waterlogging of wheat plants reduced internode lignin content”
Claim · model-written summary
Waterlogging induced expression of other lignin-related genes in wheat.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Treatment
- waterlogginge0
- Outcome
- Expression of the other genese0
- Direction
- inducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“Expression of the other genes was, however, induced by waterlogging”
Claim · model-written summary
Cultivar differences and waterlogging changes in internode lignin content were associated with cytokinin level.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Compound
- cytokinine0
- Outcome
- internode lignin contente0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“difference in internode lignin content between cultivars or change in its level due to waterlogging is associated with the level of cytokinin”
Claim · model-written summary
The authors infer that highly expressed lignin genes mediate deposition in wheat stems.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Outcome
- lignin depositione0
- Qualifier
- these genese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“implying the importance of these genes in mediating lignin deposition in wheat stem”
Claim · model-written summary
The authors suggest induced genes may synthesize other phenylpropanoid-derived stress response molecules.
- Taxon
- wheat (Triticum aestivum L.)
Triticum aestivumt0 - Outcome
- synthesis of other phenylpropanoid-derived moleculese0
- Qualifier
- roles in stress responsese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“wheat (Triticum aestivum L.)”
e0“suggesting their role in the synthesis of other phenylpropanoid-derived molecules with roles in stress responses”
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