Abstract
Waterlogging stress has been found to have adverse impacts on plant growth, subsequently reducing crop yields. Spermidine (Spd), a second messenger, positively affects the growth of plants under waterlogging stress. However, the molecular mechanisms of exogenous Spd application alleviating waterlogging stress remained unclear. In this study, we performed physiological analysis and multi-omics to underlying the effect of Spd application on waterlogging stress. Spd application increased genes expression level of light-harvesting complex (LHC) and photosynthesis-related and starch-related pathway, inhibited chlorophyll degradation and maintained higher photosynthetic rate, thus increased biomass accumulation under waterlogging stress. The activation of genes related trehalose and Spd biosynthesis would result in high accumulation of trehalose and endogenous Spd. Inhibiting 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase (ACO,) expression contributed to reduced ethylene emission. All those changes increased maize resistance to waterlogging. After Spd sparying, auxin-related genes up-regulated and IAA content was increased, those favor cell elongation in maize and thus keep normal growth after Spd application under waterlogging stress. Most of genes involved in lipids were up-regulated and thus increased lipids content and protected cell membranes in maize after Spd application under waterlogging conditions. All those changes contributed to increasing the resistance to waterlogging stress. These findings broaden our understanding of the regulatory roles of Spd in alleviating waterlogging damage and will provide evidence for breeding waterlogging-tolerant maize varieties.
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Claims 14
Claim · model-written summary
Spd application increased expression of light-harvesting complex (LHC), photosynthesis-related and starch-related pathway genes in maize under waterlogging stress.
- Treatment
- exogenous spermidine applicationt0
- Gene
- light-harvesting complex (LHC)e1
- Outcome
- genes expression levele1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“Spd application increased genes expression level of light-harvesting complex (LHC) and photosynthesis-related and starch-related pathway”
Claim · model-written summary
Spd application inhibited chlorophyll degradation in maize under waterlogging stress.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e2“inhibited chlorophyll degradation”
Claim · model-written summary
Spd application maintained a higher photosynthetic rate in maize under waterlogging stress.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e3“maintained higher photosynthetic rate”
Claim · model-written summary
Spd application increased biomass accumulation in maize under waterlogging stress.
- Treatment
- exogenous spermidine applicationt0
- Outcome
- biomass accumulatione4
- Direction
- increasede4
- Qualifier
- under waterlogging stresse4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e4“increased biomass accumulation under waterlogging stress”
Claim · model-written summary
Activation of genes related to trehalose and Spd biosynthesis would result in high accumulation of trehalose and endogenous Spd in maize.
- Gene
- genes related trehalose and Spd biosynthesise5
- Compound
- trehalosee5
- Outcome
- high accumulation of trehalose and endogenous Spde5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e5“The activation of genes related trehalose and Spd biosynthesis would result in high accumulation of trehalose and endogenous Spd”
Claim · model-written summary
Inhibiting ACC oxidase (ACO) expression reduced ethylene emission in maize under Spd application.
- Treatment
- Inhibiting 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase (ACO,) expressione6
- Gene
- 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase (ACO,)e6
- Outcome
- ethylene emissione6
- Direction
- reducede6
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e6“Inhibiting 1-aminocyclopropane-1-carboxylic acid (ACC) oxidase (ACO,) expression contributed to reduced ethylene emission”
Claim · model-written summary
Spd spraying up-regulated auxin-related genes in maize under waterlogging stress.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e7“After Spd sparying, auxin-related genes up-regulated”
Claim · model-written summary
Spd application increased IAA content in maize under waterlogging stress.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“IAA content was increased”
Claim · model-written summary
Spd application kept normal growth of maize under waterlogging stress.
- Treatment
- exogenous spermidine applicationt0
- Outcome
- normal growthe1
- Qualifier
- under waterlogging stresse1
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“keep normal growth after Spd application under waterlogging stress”
Claim · model-written summary
Auxin-related genes and IAA are proposed to favour cell elongation in maize under Spd application.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e7“After Spd sparying, auxin-related genes up-regulated”
e2“IAA content was increased”
e3“those favor cell elongation in maize”
Claim · model-written summary
Most genes involved in lipids were up-regulated in maize under Spd application and waterlogging.
- Treatment
- exogenous spermidine applicationt0
- Gene
- genes involved in lipidse1
- Outcome
- genes involved in lipidse1
- Direction
- up-regulatede1
- Qualifier
- Most ofe1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“Most of genes involved in lipids were up-regulated”
Claim · model-written summary
Spd application increased lipids content in maize under waterlogging conditions.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“increased lipids content”
Claim · model-written summary
Spd application protected cell membranes in maize under waterlogging conditions.
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“protected cell membranes”
Claim · model-written summary
Spd-induced changes increased maize resistance to waterlogging.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Integrated transcriptomics and metabolomics analysis provide insights into the alleviation of waterlogging stress in maize by exogenous spermidine application”
e1“All those changes increased maize resistance to waterlogging”
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