Abstract
Abstract Heterophylly is regard as an important adaptive mechanism in response to different environments within plants. However, the genetic mechanisms responsible for heterophylly in woody plants are still poorly understood. Herein, the divergence of heterophyllous leaves was investigated at morphogenesis and using microdissection and physiological indexes in paper mulberry, and the genetic basis of heterophylly was further revealed combined with genome-wide association study (GWAS), transcriptome analysis and weighted gene coexpression network analysis (WGCNA). Our results revealed that the flavonoid content and antioxidant activity increased gradually from the entire leaf to the palmatisect leaf, while the hormone content and net photosynthetic rate decreased. Through GWAS and transcriptome analysis, a total of 98 candidate genes and 2338 differentially expressed genes associated with heterophylly were identified. Importantly, we uncovered critical variations in the candidate genes Bp07g0981 (WOX) and Bp07g0920 (HHO), along with significant differences in haplotypes and expression levels among heterophyllous leaves. Our results also suggested that the genes involved in hormone signaling pathways, antioxidant activity, and flavonoid metabolism might be closely related to the heterophylly of paper mulberry, which could account for the physiological data. Indeed, CR-wox mutant lines showed significant changes in leaf phenotypes, and differential expression profile analysis also highlighted the expression of genes related to phytohormones and transcription factors. Together, the genetic variations and candidate genes detected in this study provide novel insights into the genetic mechanism of heterophylly, and would improve the understanding of eco-adaptability in heterophyllous woody plants.
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Claims 11
Claim · model-written summary
Flavonoid content increased from entire to palmatisect paper mulberry leaves.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Organ / stage
- palmatisect leafe0
- Outcome
- flavonoid contente0
- Direction
- increased graduallye0
- Comparator
- entire leafe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the flavonoid content and antioxidant activity increased gradually from the entire leaf to the palmatisect leaf, while the hormone content and net photosynthetic rate decreased”
t0“in paper mulberry”
Claim · model-written summary
Antioxidant activity increased from entire to palmatisect leaves.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Organ / stage
- palmatisect leafe0
- Outcome
- antioxidant activitye0
- Direction
- increased graduallye0
- Comparator
- entire leafe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the flavonoid content and antioxidant activity increased gradually from the entire leaf to the palmatisect leaf, while the hormone content and net photosynthetic rate decreased”
t0“in paper mulberry”
Claim · model-written summary
Hormone content decreased from entire to palmatisect leaves.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Organ / stage
- palmatisect leafe0
- Outcome
- hormone contente0
- Direction
- decreasede0
- Comparator
- entire leafe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the flavonoid content and antioxidant activity increased gradually from the entire leaf to the palmatisect leaf, while the hormone content and net photosynthetic rate decreased”
t0“in paper mulberry”
Claim · model-written summary
Net photosynthetic rate decreased from entire to palmatisect leaves.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Organ / stage
- palmatisect leafe0
- Outcome
- net photosynthetic ratee0
- Direction
- decreasede0
- Comparator
- entire leafe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the flavonoid content and antioxidant activity increased gradually from the entire leaf to the palmatisect leaf, while the hormone content and net photosynthetic rate decreased”
t0“in paper mulberry”
Claim · model-written summary
GWAS and transcriptomics identified 98 candidate genes associated with heterophylly.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Outcome
- candidate genese0
- Value
- 98 candidate genese0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Through GWAS and transcriptome analysis, a total of 98 candidate genes and 2338 differentially expressed genes associated with heterophylly were identified”
t0“in paper mulberry”
Claim · model-written summary
GWAS and transcriptomics identified 2338 differentially expressed genes associated with heterophylly.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Outcome
- differentially expressed genese0
- Value
- 2338 differentially expressed genese0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Through GWAS and transcriptome analysis, a total of 98 candidate genes and 2338 differentially expressed genes associated with heterophylly were identified”
t0“in paper mulberry”
Claim · model-written summary
Bp07g0981 (WOX) showed critical variation and haplotype and expression differences among leaf forms.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Gene
- Bp07g0981 (WOX)e0
- Outcome
- critical variationse0
- Qualifier
- significant differences in haplotypes and expression levelse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“we uncovered critical variations in the candidate genes Bp07g0981 (WOX) and Bp07g0920 (HHO), along with significant differences in haplotypes and expression levels among heterophyllous leaves”
t0“in paper mulberry”
Claim · model-written summary
Bp07g0920 (HHO) showed critical variation and haplotype and expression differences among leaf forms.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Gene
- Bp07g0920 (HHO)e0
- Outcome
- critical variationse0
- Qualifier
- significant differences in haplotypes and expression levelse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“we uncovered critical variations in the candidate genes Bp07g0981 (WOX) and Bp07g0920 (HHO), along with significant differences in haplotypes and expression levels among heterophyllous leaves”
t0“in paper mulberry”
Claim · model-written summary
Genes in hormone signaling, antioxidant activity and flavonoid metabolism were suggested to relate to heterophylly.
- Taxon
- paper mulberry
Broussonetia papyriferae0 - Gene
- genes involved in hormone signaling pathways, antioxidant activity, and flavonoid metabolisme0
- Outcome
- heterophyllye0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the genes involved in hormone signaling pathways, antioxidant activity, and flavonoid metabolism might be closely related to the heterophylly of paper mulberry”
Claim · model-written summary
CR-wox mutant lines showed significant leaf phenotype changes.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Material
- CR-wox mutant linese0
- Outcome
- changes in leaf phenotypese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“CR-wox mutant lines showed significant changes in leaf phenotypes”
t0“in paper mulberry”
Claim · model-written summary
Differential expression analysis highlighted genes related to phytohormones and transcription factors.
- Taxon
- paper mulberry
Broussonetia papyriferat0 - Outcome
- expression of genes related to phytohormones and transcription factorse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“differential expression profile analysis also highlighted the expression of genes related to phytohormones and transcription factors”
t0“in paper mulberry”