Abstract
WRKY transcription factors serve as key regulators in plants, playing important roles in growth and development, secondary metabolism, and stress responses. Here, a comprehensive genome-wide analysis identified 58 WRKY genes (LcWRKYs) in litchi for the first time. All LcWRKY proteins were predicted to be hydrophilic and localized in the nucleus. Phylogenetic analysis classified them into three major groups (Groups I, II, and III), with a pronounced expansion of Group II, which contained 42 members divided into five subgroups. Members within the same phylogenetic clade exhibited highly similar exon–intron structures and conserved motif compositions, indicating strong evolutionary conservation. LcWRKYs were unevenly distributed across the litchi chromosomes, with chromosome 3 showing the highest gene density. Collinearity analysis suggested that both segmental and tandem duplications contributed to the evolutionary expansion of this family. Notably, promoter cis-acting element analysis revealed that LcWRKYs are enriched with light-responsive, hormone-responsive (e.g., ABA, MeJA, SA), and stress-responsive elements, suggesting their potential involvement in integrating light signaling, hormonal pathways, and environmental stress responses. Integrative expression analysis further revealed that multiple LcWRKYs were significantly up-regulated during the middle and late stages of fruit development in cultivars such as ‘Feizixiao’ and ‘Nuomici’. Consistent with these patterns, qRT-PCR validation demonstrated a pronounced induction of four representative genes (LITCHI004628.m1, LITCHI018082.m1, LITCHI021964.m1, and LITCHI030932.m1) at 40 days post-anthesis, followed by gene-specific expression trajectories at later stages, indicating their potential involvement in regulating fruit development, particularly during the mid-developmental stage. Altogether, the results of this study provide insight into the expansion and potential functional diversification of WRKY transcription factors in litchi and identify candidate regulators associated with fruit development, offering valuable targets for future functional studies and genetic improvement.
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Claims 11
Claim · model-written summary
A genome-wide analysis identified 58 WRKY genes (LcWRKYs) in litchi for the first time.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“a comprehensive genome-wide analysis identified 58 WRKY genes (LcWRKYs) in litchi for the first time”
Claim · model-written summary
Phylogenetic analysis classified the LcWRKY proteins into three major groups (I, II and III).
- Outcome
- three major groupse0
Not reportedGene
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“classified them into three major groups (Groups I, II, and III)”
Claim · model-written summary
Group II was markedly expanded, containing 42 members divided into five subgroups.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“a pronounced expansion of Group II, which contained 42 members divided into five subgroups”
Claim · model-written summary
LcWRKY members within the same phylogenetic clade had highly similar exon-intron structures and conserved motif compositions.
- Outcome
- highly similar exon–intron structures and conserved motif compositionse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“Members within the same phylogenetic clade exhibited highly similar exon–intron structures and conserved motif compositions”
Claim · model-written summary
LcWRKYs were unevenly distributed across litchi chromosomes, with chromosome 3 showing the highest gene density.
- Gene
- LcWRKYse0
- Outcome
- unevenly distributed across the litchi chromosomese0
- Qualifier
- chromosome 3 showing the highest gene densitye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“LcWRKYs were unevenly distributed across the litchi chromosomes, with chromosome 3 showing the highest gene density”
Claim · model-written summary
Collinearity analysis suggested that segmental and tandem duplications contributed to the evolutionary expansion of the WRKY family.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“Collinearity analysis suggested that both segmental and tandem duplications contributed to the evolutionary expansion of this family”
Claim · model-written summary
Promoter analysis showed LcWRKYs are enriched with light-, hormone- (e.g., ABA, MeJA, SA) and stress-responsive cis-acting elements.
- Gene
- LcWRKYse0
- Outcome
- enriched with light-responsive, hormone-responsive (e.g., ABA, MeJA, SA), and stress-responsive elementse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“LcWRKYs are enriched with light-responsive, hormone-responsive (e.g., ABA, MeJA, SA), and stress-responsive elements”
Claim · model-written summary
Multiple LcWRKYs were significantly up-regulated during the middle and late stages of fruit development.
- Organ / stage
- middle and late stages of fruit developmente0
- Gene
- LcWRKYse0
- Outcome
- up-regulatede0
- Direction
- up-regulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“multiple LcWRKYs were significantly up-regulated during the middle and late stages of fruit development”
Claim · model-written summary
qRT-PCR showed pronounced induction of four representative LcWRKY genes at 40 days post-anthesis.
- Organ / stage
- 40 days post-anthesise0
- Gene
- LITCHI004628.m1, LITCHI018082.m1, LITCHI021964.m1, and LITCHI030932.m1e0
- Outcome
- pronounced inductione0
- Direction
- inductione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“qRT-PCR validation demonstrated a pronounced induction of four representative genes (LITCHI004628.m1, LITCHI018082.m1, LITCHI021964.m1, and LITCHI030932.m1) at 40 days post-anthesis”
Claim · model-written summary
The four representative genes are suggested to be potentially involved in regulating fruit development, particularly mid-development.
- Organ / stage
- mid-developmental stagee1
- Gene
- four representative genese0
- Outcome
- regulating fruit developmente1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“four representative genes”
e1“their potential involvement in regulating fruit development, particularly during the mid-developmental stage”
Claim · model-written summary
All LcWRKY proteins were predicted to be hydrophilic and nuclear-localized.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Litchi”
e0“All LcWRKY proteins were predicted to be hydrophilic and localized in the nucleus”