Abstract
ABSTRACT Dwarfing is a pivotal agronomic trait affecting both yield and quality. Citrus species exhibit substantial variation in plant height, among which internode length is a core element. However, the molecular mechanism governing internode elongation remains unclear. Here, we unveiled that the transcriptional cascade consisting of B‐BOX DOMAIN PROTEIN 22 ( BBX22 ) and ELONGATED HYPOCOTYL 5 ( HY5 ) finely tunes plant height and internode elongation in citrus. Loss‐of‐function mutations of BBX22 in an early‐flowering citrus ( Citrus hindsii “SJG”) promoted internode elongation and reduced pigment accumulation, whereas ectopic expression of BBX22 in SJG, sweet orange ( C. sinensis ), pomelo ( C. maxima ) or heterologous expression of BBX22 in tomato ( Solanum lycopersicum ) significantly decreased internode length. Furthermore, exogenous application of gibberellin A3 (GA 3 ) rescued the shortened internode and dwarf phenotype caused by BBX22 overexpression. Additional experiments revealed that BBX22 played a dual role in regulation internode elongation and pigmentation in citrus. On the one hand, it directly bound to and activated the expression of HY5 , GA metabolism gene ( GA2 OXIDASE 8 , GA2ox8 ), carotenoid biosynthesis gene ( PHYTOENE SYNTHASE 1 , PSY1 ) and anthocyanin regulatory gene ( Ruby1, a MYB DOMAIN PROTEIN). On the other hand, it acted as a cofactor of HY5, enhancing the ability of HY5 to regulate target genes expression. Together, our results reveal the critical role of the transcriptional cascade consisting of BBX22 and HY5 in controlling internode elongation and pigment accumulation in citrus. Unraveling the crosstalk regulatory mechanism between internode elongation and fruit pigmentation provides key genes for breeding of novel types with both dwarf and health‐beneficial fortification in citrus.
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Claims 11
Claim · model-written summary
Loss-of-function mutations of BBX22 in citrus promoted internode elongation.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e1“Loss‐of‐function mutations of BBX22 in an early‐flowering citrus ( Citrus hindsii “SJG”) promoted internode elongation and reduced pigment accumulation”
Claim · model-written summary
Loss-of-function mutations of BBX22 in citrus reduced pigment accumulation.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e1“Loss‐of‐function mutations of BBX22 in an early‐flowering citrus ( Citrus hindsii “SJG”) promoted internode elongation and reduced pigment accumulation”
Claim · model-written summary
Ectopic expression of BBX22 in citrus (SJG, sweet orange, pomelo) significantly decreased internode length.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e2“ectopic expression of BBX22 in SJG, sweet orange ( C. sinensis ), pomelo ( C. maxima ) or heterologous expression of BBX22 in tomato ( Solanum lycopersicum ) significantly decreased internode length”
Claim · model-written summary
Heterologous expression of BBX22 in tomato significantly decreased internode length.
- Taxon
- tomato
Solanum lycopersicume2 - Treatment
- heterologous expression of BBX22e2
- Outcome
- internode lengthe2
- Direction
- decreasede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e2“ectopic expression of BBX22 in SJG, sweet orange ( C. sinensis ), pomelo ( C. maxima ) or heterologous expression of BBX22 in tomato ( Solanum lycopersicum ) significantly decreased internode length”
Claim · model-written summary
Exogenous gibberellin A3 rescued the shortened internode and dwarf phenotype caused by BBX22 overexpression in citrus.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e3“exogenous application of gibberellin A3 (GA 3 ) rescued the shortened internode and dwarf phenotype caused by BBX22 overexpression”
Claim · model-written summary
BBX22 directly bound to and activated the expression of HY5.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e4“On the one hand, it directly bound to and activated the expression of HY5 , GA metabolism gene ( GA2 OXIDASE 8 , GA2ox8 ), carotenoid biosynthesis gene ( PHYTOENE SYNTHASE 1 , PSY1 ) and anthocyanin regulatory gene ( Ruby1, a MYB DOMAIN PROTEIN)”
Claim · model-written summary
BBX22 directly bound to and activated the expression of GA2ox8.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e4“On the one hand, it directly bound to and activated the expression of HY5 , GA metabolism gene ( GA2 OXIDASE 8 , GA2ox8 ), carotenoid biosynthesis gene ( PHYTOENE SYNTHASE 1 , PSY1 ) and anthocyanin regulatory gene ( Ruby1, a MYB DOMAIN PROTEIN)”
Claim · model-written summary
BBX22 directly bound to and activated the expression of PSY1.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e4“On the one hand, it directly bound to and activated the expression of HY5 , GA metabolism gene ( GA2 OXIDASE 8 , GA2ox8 ), carotenoid biosynthesis gene ( PHYTOENE SYNTHASE 1 , PSY1 ) and anthocyanin regulatory gene ( Ruby1, a MYB DOMAIN PROTEIN)”
Claim · model-written summary
BBX22 directly bound to and activated the expression of the anthocyanin regulatory gene Ruby1.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e4“On the one hand, it directly bound to and activated the expression of HY5 , GA metabolism gene ( GA2 OXIDASE 8 , GA2ox8 ), carotenoid biosynthesis gene ( PHYTOENE SYNTHASE 1 , PSY1 ) and anthocyanin regulatory gene ( Ruby1, a MYB DOMAIN PROTEIN)”
Claim · model-written summary
BBX22 acts as a cofactor of HY5, enhancing the ability of HY5 to regulate target gene expression.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Citrus species exhibit substantial variation in plant height”
e5“On the other hand, it acted as a cofactor of HY5, enhancing the ability of HY5 to regulate target genes expression”
Claim · model-written summary
The BBX22-HY5 transcriptional cascade controls internode elongation and pigment accumulation in citrus.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e6“Together, our results reveal the critical role of the transcriptional cascade consisting of BBX22 and HY5 in controlling internode elongation and pigment accumulation in citrus”
The same questions in other papers
loss of function · growth · CitrusPositive and negative results
3 papers in the same direction as this paper · 2 in the opposite direction
- positive
- positive
- negative
2 papers more in this group
See all 6 papers →overexpression · growth · Solanum lycopersicum (tomato)Positive and negative results
3 papers in the same direction as this paper · 7 in the opposite direction
- negative
- no effect
- negative
7 papers more in this group
See all 11 papers →loss of function · physiology: pigments · CitrusResults agree
2 papers in the same direction as this paper
- negative
- negative
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.