Abstract
Ethylene signaling pathways regulate several physiological alterations that occur during tomato fruit ripening, such as changes in colour and flavour. The mechanisms underlying the transcriptional regulation of genes in these pathways remain unclear, although the role of the MADS-box transcription factor RIN has been widely reported. Here, we describe a bHLH transcription factor, SlbHLH95, whose transcripts accumulated abundantly in breaker+4 and breaker+7 fruits compared with rin (ripening inhibitor) and Nr (never ripe) mutants. Moreover, the promoter activity of SlbHLH95 was regulated by RIN in vivo. Suppression of SlbHLH95 resulted in reduced sensitivity to ethylene, decreased accumulation of total carotenoids, and lowered glutathione content, and inhibited the expression of fruit ripening- and glutathione metabolism-related genes. Conversely, up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch. Notably, overexpression of SlbHLH95 in rin led to the up-regulated expression of fruit ripening-related genes (FUL1, FUL2, SAUR69, ERF4, and CNR) and multiple glutathione metabolism-related genes (GSH1, GSH2, GSTF1, and GSTF5). These results clarified that SlbHLH95 participates in the regulation of fruit ripening and affects ethylene sensitivity and multiple metabolisms targeted by RIN in tomato.
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Claims 14
Claim · model-written summary
SlbHLH95 transcripts accumulated more abundantly in breaker+4 and breaker+7 fruits than in rin and Nr mutants.
- Taxon
- tomato
Solanum lycopersicumt0 - Organ / stage
- breaker+4 and breaker+7 fruitse0
- Gene
- SlbHLH95e0
- Outcome
- transcripts accumulated abundantlye0
- Direction
- abundantlye0
- Comparator
- rin (ripening inhibitor) and Nr (never ripe) mutantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“SlbHLH95, whose transcripts accumulated abundantly in breaker+4 and breaker+7 fruits compared with rin (ripening inhibitor) and Nr (never ripe) mutants.”
t0“tomato”
Claim · model-written summary
RIN regulated SlbHLH95 promoter activity in vivo.
- Taxon
- tomato
Solanum lycopersicumt0 - Gene
- RINe0
- Outcome
- promoter activity of SlbHLH95e0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the promoter activity of SlbHLH95 was regulated by RIN in vivo.”
t0“tomato”
Claim · model-written summary
Suppression of SlbHLH95 caused reduced sensitivity to ethylene in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Suppression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- sensitivity to ethylenee0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Suppression of SlbHLH95 resulted in reduced sensitivity to ethylene, decreased accumulation of total carotenoids, and lowered glutathione content, and inhibited the expression of fruit ripening- and glutathione metabolism-related genes.”
t0“tomato”
Claim · model-written summary
Suppression of SlbHLH95 caused decreased accumulation of total carotenoids in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Suppression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- accumulation of total carotenoidse0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Suppression of SlbHLH95 resulted in reduced sensitivity to ethylene, decreased accumulation of total carotenoids, and lowered glutathione content, and inhibited the expression of fruit ripening- and glutathione metabolism-related genes.”
t0“tomato”
Claim · model-written summary
Suppression of SlbHLH95 caused lowered glutathione content in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Suppression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- glutathione contente0
- Direction
- lowerede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Suppression of SlbHLH95 resulted in reduced sensitivity to ethylene, decreased accumulation of total carotenoids, and lowered glutathione content, and inhibited the expression of fruit ripening- and glutathione metabolism-related genes.”
t0“tomato”
Claim · model-written summary
Suppression of SlbHLH95 caused inhibited expression of fruit ripening- and glutathione metabolism-related genes in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Suppression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- expression of fruit ripening- and glutathione metabolism-related genese0
- Direction
- inhibitede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Suppression of SlbHLH95 resulted in reduced sensitivity to ethylene, decreased accumulation of total carotenoids, and lowered glutathione content, and inhibited the expression of fruit ripening- and glutathione metabolism-related genes.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused higher sensitivity to ethylene in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- sensitivity to ethylenee0
- Direction
- highere0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused increased accumulation of total carotenoids in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- accumulation of total carotenoidse0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused premature ripening in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- ripeninge0
- Direction
- prematuree0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused elevated accumulation of glutathione in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- accumulation of glutathionee0
- Direction
- elevatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused elevated soluble sugar in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- soluble sugare0
- Direction
- elevatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
up-regulation of SlbHLH95 caused elevated starch in tomato.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- up-regulation of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- starche0
- Direction
- elevatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“up-regulation of SlbHLH95 in wild-type tomato resulted in higher sensitivity to ethylene, increased accumulation of total carotenoids, slightly premature ripening, and elevated accumulation of glutathione, soluble sugar, and starch.”
t0“tomato”
Claim · model-written summary
SlbHLH95 overexpression in rin up-regulated expression of fruit ripening-related genes (FUL1, FUL2, SAUR69, ERF4, and CNR).
- Taxon
- tomato
Solanum lycopersicumt0 - Material
- rine0
- Treatment
- overexpression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- fruit ripening-related genes (FUL1, FUL2, SAUR69, ERF4, and CNR)e0
- Direction
- up-regulatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“overexpression of SlbHLH95 in rin led to the up-regulated expression of fruit ripening-related genes (FUL1, FUL2, SAUR69, ERF4, and CNR) and multiple glutathione metabolism-related genes (GSH1, GSH2, GSTF1, and GSTF5).”
t0“tomato”
Claim · model-written summary
SlbHLH95 overexpression in rin up-regulated expression of glutathione metabolism-related genes (GSH1, GSH2, GSTF1, and GSTF5).
- Taxon
- tomato
Solanum lycopersicumt0 - Material
- rine0
- Treatment
- overexpression of SlbHLH95e0
- Gene
- SlbHLH95e0
- Outcome
- glutathione metabolism-related genes (GSH1, GSH2, GSTF1, and GSTF5)e0
- Direction
- up-regulatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“overexpression of SlbHLH95 in rin led to the up-regulated expression of fruit ripening-related genes (FUL1, FUL2, SAUR69, ERF4, and CNR) and multiple glutathione metabolism-related genes (GSH1, GSH2, GSTF1, and GSTF5).”
t0“tomato”
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