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Article

Genome-Wide Identification and Expression Analysis of the CsCAMTA Gene Family in Tieguanyin Tea Plants Under Heat Stress

by
Zijia Cui
1,
Hua Wu
2,
Zhicheng Yang
3,
Bohao Xu
1,
Fan Jiang
1,
Rien Lai
1,
Lu Han
1,
Ciding Lu
1,
Dandan Li
1,* and
Kehui Zheng
3,*
1
School of Modern Forestry, Jiangxi Environmental Engineering Vocational College, Ganzhou 341000, China
2
College of Food Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China
3
College of Computer and Information Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, China
*
Authors to whom correspondence should be addressed.
Curr. Issues Mol. Biol. 2026, 48(6), 597; https://doi.org/10.3390/cimb48060597
Submission received: 27 April 2026 / Revised: 25 May 2026 / Accepted: 28 May 2026 / Published: 5 June 2026
(This article belongs to the Topic Big Data Mining in Plant Stress Resistance Evolution and Germplasm Innovation)
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Abstract

Tieguanyin (Camellia sinensis cv. Tieguanyin) is an important oolong tea cultivar in China, and heat stress has become a major environmental constraint affecting its growth and productivity. Calmodulin-binding transcription activators (CAMTAs) are important transcription factors involved in calcium/calmodulin-mediated signaling and plant responses to environmental stresses. However, systematic knowledge of the CAMTA gene family in Tieguanyin remains limited. In this study, 20 CsCAMTA genes were identified from the Tieguanyin genome and characterized based on their physicochemical properties, phylogenetic relationships, conserved motifs, gene structures, chromosomal distribution, collinearity, promoter cis-acting elements, and functional annotation. The 20 CsCAMTA genes were unevenly distributed across eight chromosomes, and collinearity analysis suggested that segmental duplication may have contributed to the expansion of this gene family. Conserved motif and domain analyses indicated that CsCAMTA proteins retained typical structural features of CAMTA transcription factors, including CG-1, ANKYR, TIG, and CaMBD/IQ-related regions. Promoter analysis showed that CsCAMTA genes harbored multiple cis-acting elements related to hormone responsiveness, stress response, light response, and growth regulation. Furthermore, qRT-PCR analysis of 18 representative CsCAMTA genes under 40 °C heat treatment revealed distinct temporal expression patterns, suggesting that different CsCAMTA members may respond to heat stress at different stages. Several genes, such as CsCAMTA2, CsCAMTA10, and CsCAMTA16, showed marked transcriptional changes and may represent candidate heat-responsive genes in Tieguanyin. These results provide a systematic overview of the CsCAMTA gene family and lay a foundation for further functional studies of heat stress responses in Tieguanyin.

1. Introduction

Tieguanyin (Camellia sinensis cv. Tieguanyin) is a representative oolong tea cultivar in China. Owing to its distinctive aroma profile and favorable processing characteristics, Tieguanyin has become an important resource for the high-value development of the tea industry [1,2,3,4]. However, in recent years, the intensification of global climate change has increased the frequency of extreme weather events, including drought, high temperature, and strong radiation, which seriously threaten the yield and quality stability of Tieguanyin in its major production areas [5]. Abiotic stresses can induce the overproduction of reactive oxygen species (ROS), impair the photosynthetic system, and disrupt secondary metabolic networks, thereby altering the balance of tea polyphenols and amino acids and ultimately affecting tea flavor quality [6]. Therefore, elucidating the molecular regulatory networks underlying stress responses in Tieguanyin, especially the core components of stress signal transduction, is important for variety improvement and the sustainable development of the tea industry.
Calcium signaling is a central regulatory system in plant responses to environmental stresses. As key components of this system, calmodulin-binding transcription activators (CAMTAs) play important roles in regulating plant environmental adaptation [7,8,9]. CAMTA proteins typically contain conserved CG-1 DNA-binding domains, ANK repeats, and IQ motifs [10]. They can recognize the cis-acting element (A/C) CGCG box and participate in stress responses by regulating downstream gene expression networks associated with abscisic acid (ABA), salicylic acid (SA), and heat shock proteins (HSPs) [11]. In model plants, AtCAMTA3 enhances cold tolerance in Arabidopsis by activating the CBF/DREB1 cold-response module [12,13,14]. In soybean, GmCAMTA12 regulates stomatal movement through an ABA-dependent pathway and improves drought resistance [15,16]. GhCAMTA11 in cotton has been identified as a key regulatory factor involved in heat stress responses, and its overexpression significantly enhances plant heat tolerance [17]. In addition, SlCAMTA4 in tomato affects plant resistance to pathogens by regulating the jasmonic acid (JA) signaling pathway, while OsCAMTA1 in rice is involved in salt stress responses [18,19]. These studies indicate the evolutionary conservation and functional diversity of the CAMTA family in plant stress signaling networks.
In recent years, with the development of tea plant genomics and transcriptomics, several CsCAMTA members have been identified in tea plants, and their expression patterns have shown tissue-specific and treatment-dependent characteristics under low-temperature, drought, and ABA treatments [10]. For example, the expression levels of CsCAMTA3 and CsCAMTA5 in the cold-tolerant cultivar “Longjing 43” were higher than those in the cold-sensitive cultivar “Da Bai Mian”, suggesting their potential involvement in tea plant cold adaptation. Furthermore, studies on woody plants have expanded our understanding of CAMTA function. For example, in Phoebe bournei, 17 PbCAMTA genes form a dynamic co-expression network under drought, high-temperature, and salt stresses, with PbCAMTA2, PbCAMTA12, and PbCAMTA16 identified as core stress-response nodes [9]. In Populus trichocarpa, PtCAMTA7 enhances drought tolerance by regulating antioxidant enzyme activity and increasing ROS-scavenging capacity [20]. These findings provide useful references for understanding stress adaptation mechanisms in woody plants, including tea plants.
Although CAMTA genes have been reported in several plant species, systematic identification and heat-responsive expression analysis of the CAMTA gene family in Tieguanyin tea plants remain limited. In this study, we performed genome-wide identification of CsCAMTA genes in Tieguanyin and analyzed their phylogenetic relationships, physicochemical properties, conserved motifs, gene structures, chromosomal distribution, duplication events, promoter cis-acting elements, and functional annotations. In addition, 18 representative CsCAMTA genes were selected for qRT-PCR analysis under heat stress. This study aimed to clarify the basic characteristics of the CsCAMTA gene family in Tieguanyin and to identify potential candidate genes involved in heat stress responses.

2. Materials and Methods

2.1. Data Sources

The genome assembly and genome annotation files of Tieguanyin tea plant were retrieved from the Genome Warehouse of the China National GeneBank Database under accession number GWHASIV00000000 [21]. To support comparative analyses, genome sequences and corresponding annotation files of Arabidopsis thaliana, Triticum aestivum, Zea mays, Nicotiana tabacum, Glycine max (L.) Merr. and Solanum lycopersicum were downloaded from public genome databases, including TAIR (https://www.arabidopsis.org/, accessed on 3 May 2025) and Ensembl Plants (https://plants.ensembl.org/index.html, accessed on 3 May 2025). The reported CAMTA protein sequences of A. thaliana were collected from PlantTFDB (https://planttfdb.gao-lab.org/, accessed on 25 May 2025) and used as query sequences for subsequent identification. The hidden Markov model profile of the CAMTA-associated conserved domain was downloaded from the Pfam database (http://pfam.xfam.org/, accessed on 3 May 2025) and used for domain-based screening of candidate CAMTA members.

2.2. Identification and Physicochemical Properties of CAMTA Gene Family

To identify CAMTA family members in Tieguanyin, two complementary strategies were used. First, the known A. thaliana CAMTA protein sequences were used as queries to search against the Tieguanyin protein database using the BLAST module in TBtools (version 2.310). Second, the HMM profile of the CAMTA conserved domain was employed to perform an HMMER-based search against the Tieguanyin proteome. Candidate proteins identified by both approaches were merged and redundant sequences were removed. The conserved domains of the remaining candidates were further examined using the NCBI Conserved Domain Database to confirm the presence and completeness of characteristic CAMTA domains. Sequences lacking essential conserved regions or showing obvious annotation abnormalities were excluded from further analysis. Multiple sequence alignment of confirmed CsCAMTA proteins was performed using DNAMAN (version 9.0) to examine sequence conservation among family members [22]. The amino acid length, molecular weight, theoretical isoelectric point, instability index, and other physicochemical parameters of CsCAMTA proteins were predicted using the ExPASy ProtParam (https://web.expasy.org/protparam/, accessed on 3 May 2025) [23], and the resulting data were summarized using WPS Excel (version 12.1.0).

2.3. Analysis of Gene Structure and Conserved Motifs

The gene structure information of CsCAMTA genes was extracted from the genome annotation files in GFF3/GTF format. The exon–intron organization of each CsCAMTA gene was visualized using the Gene Structure View module in TBtools. To identify conserved motifs in CsCAMTA proteins, the full-length amino acid sequences of CsCAMTA family members were submitted to the MEME online program (https://meme-suite.org/meme/, accessed on 6 May 2025), with the maximum number of motifs set to 10. The distribution of conserved motifs was further visualized using TBtools. Conserved protein domains were analyzed using the NCBI Conserved Domain Database (CDD) (https://www.be-md.ncbi.nlm.nih.gov/cdd/, accessed on 6 May 2025), and the domain organization of CsCAMTA proteins was displayed using the Visualize NCBI CDD Domain Pattern function in TBtools. The phylogenetic tree, conserved motifs, conserved domains, and gene structures were integrated to compare the structural conservation and divergence among CsCAMTA family members.

2.4. Chromosome Mapping and Collinearity Analysis

The chromosomal locations of CsCAMTA genes were obtained from the Tieguanyin genome annotation file and plotted using the Gene Location Visualize function in TBtools. The physical positions of CsCAMTA genes on chromosomes were adjusted according to chromosome length and gene coordinate information. Gene duplication events within the Tieguanyin genome were analyzed using MCScanX of TBtools, and duplicated CsCAMTA gene pairs were visualized using the Circos function in TBtools [24]. To further explore the evolutionary relationships of CAMTA genes across species, interspecific synteny analyses were performed between Tieguanyin and A. thaliana, T. aestivum, Z. mays, and N. tabacum using MCScanX. The collinear gene pairs were then visualized to evaluate the conservation and divergence of CAMTA genes among different plant species.

2.5. Phylogenetic Tree Construction

The full-length CAMTA protein sequences from Tieguanyin, soybean, tobacco, tomato, and Arabidopsis thaliana were used for phylogenetic analysis. Multiple sequence alignment was first conducted, and the resulting alignment file was used to construct a phylogenetic tree with the One Step Build a NJ Tree module in MEGA12. Bootstrap analysis was performed with 1000 replicates to evaluate the reliability of each branch. The final phylogenetic tree was exported and further refined using Adobe Illustrator CC 2018 [25]. Based on the clustering relationship between CsCAMTA and AtCAMTA proteins, CsCAMTA members were classified into different subgroups.

2.6. Promoter Cis-Acting Element Analysis

For promoter analysis, the 2000 bp upstream sequences of CsCAMTA genes were extracted from the Tieguanyin genome using the Sequence Extract function in TBtools. These upstream sequences were submitted to PlantCARE to predict putative cis-acting regulatory elements. The identified cis-elements were classified according to their functional annotations, including elements related to hormone responsiveness, abiotic stress response, light response, and growth regulation. The distribution of major cis-elements in CsCAMTA promoter regions was visualized using the Simple BioSequence Viewer module in TBtools.

2.7. Tissue-Specific Expression Analysis

To investigate the expression characteristics of CsCAMTA genes in different tissues, transcriptome-derived FPKM values of roots, stems, and leaves of Tieguanyin were collected and analyzed. The expression matrix of CsCAMTA genes was organized and normalized before visualization. Expression heat maps were generated using the HeatMap module in TBtools, and the final figures were adjusted for clarity. Differences in expression levels among roots, stems, and leaves were used to infer the potential tissue-specific functions of CsCAMTA genes.

2.8. Prediction of Secondary and Tertiary Protein Structures

The secondary structures of CsCAMTA proteins were predicted using SOPMA (https://npsa.lyon.inserm.fr/cgi-bin/npsa_automat.pl?page=/NPSA/npsa_sopma.html, accessed on 20 December 2025), including the proportions of α-helix, extended strand, β-turn, and random coil. For tertiary structure prediction, the amino acid sequences of CsCAMTA proteins were submitted to SWISS-MODEL for homology modeling. The predicted three-dimensional structures were compared among different CsCAMTA subgroups to evaluate potential structural differences and functional divergence within the family.

2.9. Plant Materials and Heat Treatment

One-year-old Tieguanyin tea seedlings with uniform growth status were used as experimental materials. The seedlings were obtained from Xiping Town, Anxi County, Fujian Province, China, and maintained under controlled conditions in an artificial climate chamber before stress treatment. For heat treatment, healthy and consistently growing seedlings were transferred to a growth chamber set at 40 °C. Leaf samples were collected at 0, 4, 8, 12, and 24 h after treatment, with the 0 h samples serving as the control. For each time point, three independent biological replicates were prepared, and each biological replicate consisted of pooled leaves collected from at least three individual seedlings. All samples were immediately frozen in liquid nitrogen after collection and stored at −80 °C until RNA extraction.

2.10. RNA Extraction, cDNA Synthesis and qRT-PCR Analysis

Total RNA was extracted from heat-treated Tieguanyin tea leaf samples using an RNA extraction kit (RA106-01, Beijing Biomarker Technologies Co., Ltd., Beijing, China) according to the manufacturer’s instructions. The concentration and purity of RNA were assessed using a NanoDrop spectrophotometer by Thermo Fisher Scientific (Waltham, MA, USA), and RNA integrity was checked by agarose gel electrophoresis. High-quality RNA was then used for first-strand cDNA synthesis using the MT403-01 reverse transcription kit (Beijing Biomarker Technologies Co., Ltd., Beijing, China) following the manufacturer’s protocol.
To examine the transcriptional responses of CsCAMTA genes under heat stress, 18 representative CsCAMTA genes were selected for qRT-PCR analysis. Gene-specific primers were designed in the non-conserved regions of the target genes using TBtools software, and the primers were synthesized by Fuzhou Boshan Biotechnology Co., Ltd. (Fuzhou, China).
qRT-PCR was performed using SYBR Green qPCR Master Mix by Thermo Fisher Scientific (Waltham, MA, USA) on a real-time PCR detection system. The amplification program was as follows: initial denaturation at 95 °C for 30 s, followed by 40 cycles of denaturation at 95 °C for 5 s and annealing/extension at 60 °C for 30 s. Melting curve analysis was performed after amplification to verify the specificity of PCR products. CsGAPDH (accession number: GE651107) was used as the internal reference gene. The relative expression levels of CsCAMTA genes were calculated using the 2−ΔΔCt method. Each sample included three biological replicates and three technical replicates.

2.11. Statistical Analysis

All qRT-PCR data were obtained from three independent biological replicates, and each biological replicate included three technical replicates. Data are presented as the mean ± standard deviation (SD). Differences among different heat treatment time points were analyzed by one-way analysis of variance (ANOVA) using GraphPad Prism 10. Statistical significance was indicated as follows: * p ≤ 0.05, ** p ≤ 0.01, *** p ≤ 0.001, and **** p ≤ 0.0001.

3. Results

3.1. Phylogenetic Analysis of the CAMTA Gene Family in Tieguanyin

This study conducted a phylogenetic analysis of CAMTA proteins from tea plant (Camellia sinensis var. assamica, Cs), soybean (Glycine max, Gm), tobacco (Nicotiana tabacum, Nt), tomato (Solanum lycopersicum, Sl), and Arabidopsis thaliana (At). Previous studies have reported the identification and phylogenetic analysis of CAMTA genes in tea plants and other plant species [26,27]. In the present study, the phylogenetic tree was constructed using the Neighbor-Joining (NJ) method with 1000 bootstrap replicates. The resulting phylogenetic tree is shown in Figure 1. In this tree, CAMTA proteins from different species were color-coded in the inner circle, while the outer circle distinguished the four CAMTA subgroups using different colors.
As shown in Figure 1, CAMTA proteins from the five species could be divided into four major subgroups. Several CsCAMTA proteins clustered closely with homologous CAMTA proteins from soybean, tobacco, tomato, or Arabidopsis, suggesting a certain degree of evolutionary conservation among CAMTA members from different plant species. Meanwhile, some CsCAMTA members were located in relatively independent branches, indicating possible sequence divergence within the Tieguanyin CAMTA gene family.
Overall, the phylogenetic analysis provides a framework for understanding the evolutionary relationships of CsCAMTA proteins and lays a foundation for subsequent analyses of gene structure, conserved domains, chromosomal distribution, and expression patterns under heat stress.

3.2. Collinearity and Physicochemical Characterization of the CsCAMTA Gene Family

As shown in Figure 2, the black curves in the figure connect CsCAMTA genes on different chromosomes, and these links represent intraspecific collinear blocks (segmental duplication), indicating that these genes are homologous copies generated via chromosomal segmental duplication [28,29].
This collinear relationship directly supports the evolutionary conclusion that “gene duplication events drive family expansion” and clearly demonstrates the molecular mechanism underlying the expansion of the CsCAMTA family via segmental duplication in the Tieguanyin genome [10,28].
As summarized in Table 1, the CsCAMTA gene family comprises 20 members (CsCAMTA1–CsCAMTA20), which exhibit significant variations in sequence length (164–1158 amino acids) and molecular weight (17.96–128.46 kDa). This length diversity suggests differential distribution of functional domains, with longer members (e.g., CsCAMTA3, CsCAMTA6, CsCAMTA12) potentially possessing more complex regulatory functions. Isoelectric point analysis revealed a broad pI range (4.54–8.73), where acidic members (e.g., CsCAMTA2, CsCAMTA4) may preferentially localize to nuclear or chloroplast stroma environments, while alkaline members (e.g., CsCAMTA9, CsCAMTA13) are likely involved in DNA binding or membrane interactions. Hydrophilicity analysis indicated that the vast majority of proteins exhibit hydrophilic properties (GRAVY = −0.655 to 0.055), with CsCAMTA17 and CsCAMTA18 showing positive GRAVY values, suggesting the presence of potential hydrophobic domains that may participate in membrane-associated functions.
Protein stability analysis classifies family members into stable (7 members, e.g., CsCAMTA2, CsCAMTA9) and unstable (13 members, e.g., CsCAMTA3, CsCAMTA7) types, with these differences potentially arising from post-translational modifications or chaperone protein regulation. Subcellular localization predictions reveal distinct functional partitioning: nuclear-localized members (6) may primarily regulate transcription, chloroplast-localized members (7) could participate in photosynthesis and stress responses, cytoplasmic members (5) might mediate signal transduction, while the uniquely plasma membrane-localized CsCAMTA14 may be involved in transmembrane transport. This multi-layered localization pattern fully demonstrates the functional versatility of the CsCAMTA family in tea plant growth, development, and environmental adaptation, providing critical insights for deeper understanding of its molecular mechanisms.

3.3. Structural Characterization of the CsCAMTA Gene Family in Tieguanyin

To investigate the structural characteristics of the CsCAMTA gene family in Tieguanyin tea plants, conserved motifs, conserved domains, and exon–intron structures were systematically analyzed. As shown in Figure 3, the CsCAMTA proteins displayed different motif compositions, conserved domain arrangements, and gene structure patterns, suggesting structural conservation and divergence among family members. Using the MEME tool, a total of 10 conserved motifs (Motif 1–10) were identified. Among them, Motif 2 was present in most family members except CsCAMTA7–9 and CsCAMTA13, suggesting that this motif may represent a relatively conserved structural feature of CsCAMTA proteins. Motif 3 was detected in several members, indicating possible structural conservation among specific CsCAMTA proteins. Motifs 5 and 8 were found in CsCAMTA3 and CsCAMTA6, whereas Motifs 7 and 9 were mainly distributed in CsCAMTA10 and CsCAMTA15, suggesting potential structural divergence among different members.
Domain analysis revealed the differential distribution of domains—ANKYR (16 members), CG-1 (11 members), TIG (7 members), and DHHC zinc finger (1 member) —and this distribution pattern reflected functional divergence. In particular, the synergistic interaction between the CG-1 domain and IQ motifs facilitated the coupling of calcium signaling with transcriptional regulation.
Gene structure analysis showed that the number of exons (1–15) varied greatly among family members, and significant variations were also observed in non-coding regions. The absence of 3′UTR in some CsCAMTA gene members may enhance mRNA stability, whereas the longer UTRs (>500 bp) in CsCAMTA17 and CsCAMTA20 likely contain abundant regulatory elements. Evolutionary analysis indicated that gene duplication events drove family expansion. While the conserved core motifs (Motif 1–3) and CG-1 domain maintained basic functions, the combination of specific motifs and domains, such as ANKYR and TIG domains, promoted functional divergence, enabling Tieguanyin to adapt to complex environments such as high altitude and low temperature. These findings provide important insights for further elucidating the role of CsCAMTA genes in environmental adaptation and quality formation in tea plants.
Overall, the integrated analysis of conserved motifs, conserved domains, and exon–intron structures indicated that the CsCAMTA gene family exhibits both structural conservation and divergence. These results provide useful information for subsequent expression analysis and functional characterization of CsCAMTA genes.

3.4. Chromosomal Distribution and Evolutionary Analysis of the CsCAMTA Gene Family in Tieguanyin

In this study, a total of 20 CsCAMTA genes were identified in Tieguanyin and unevenly distributed across eight chromosomes. The densest distributions were observed on chromosomes 1 and 2, each harboring four CsCAMTA members, namely CsTGYCAMTA01–CsTGYCAMTA04 and CsTGYCAMTA05–CsTGYCAMTA08, respectively. Chromosomes 5, 6, and 11 each contained three CsCAMTA genes, whereas chromosomes 9, 10, and 13 each carried only one CsCAMTA gene. This uneven distribution suggests that CsCAMTA genes were not uniformly retained across the Tieguanyin genome.
Collinearity analysis identified three segmental duplication groups within the CsCAMTA gene family. As shown in Table 2, Group 1 included CsTGYCAMTA03, CsTGYCAMTA05, and CsTGYCAMTA06; Group 2 included CsTGYCAMTA10, CsTGYCAMTA11, and CsTGYCAMTA15; and Group 3 included CsTGYCAMTA12 and CsTGYCAMTA16. These duplicated gene groups suggest that segmental duplication may have contributed to the expansion of the CsCAMTA gene family in Tieguanyin.
Previous studies have reported that tandem and segmental duplications are important mechanisms involved in the expansion and diversification of plant gene families. Consistent with these findings, the collinearity results in this study provide useful information for understanding the evolutionary expansion of CsCAMTA genes in Tieguanyin. Overall, the chromosomal distribution and collinearity analyses establish a basis for subsequent analyses of CsCAMTA gene structure, conserved domains, and expression patterns.

3.5. Cross-Species Collinearity Analysis of CAMTA Genes Between Tea Plants and Other Plant Species

In this study, multi-dimensional synteny analysis was performed to elucidate the evolutionary characteristics of the calmodulin-binding transcription activator gene family (CsCAMTA) in tea plants. As shown in Figure 4, interspecific synteny analysis revealed significant syntenic relationships between CsCAMTA genes and their homologs in species including wheat (Triticum aestivum), maize (Zea mays), Arabidopsis (Arabidopsis thaliana), and tobacco (Nicotiana tabacum) [30,31,32,33]. Specifically, syntenic genes between Arabidopsis and Tieguanyin were mainly distributed on Chr01-Chr03 and Chr05 of Arabidopsis; syntenic genes between wheat and Tieguanyin were concentrated on Chr03-Chr04 and Chr19-Chr21 of wheat; syntenic genes between maize and Tieguanyin were concentrated on Chr01-Chr02 and Chr07-Chr09 of maize; while syntenic regions between tobacco and Tieguanyin were widely distributed on Chr01-Chr04, Chr09, Chr10, and Chr12-Chr14 of tobacco. The Chr01, Chr02, and Chr06 chromosomes of Tieguanyin exhibited synteny with all four aforementioned species, suggesting that these three chromosomes are evolutionarily highly conserved core segments in the tea plant genome. The CsCAMTA genes carried on these chromosomes may be associated with fundamental biological processes such as calcium signal transduction and stress responses. Considering that transcription factors are widely involved in plant responses to abiotic stresses [34], these conserved chromosomal regions may provide useful candidate loci for subsequent functional validation and molecular breeding [9,28,29].

3.6. Analysis of Cis-Acting Elements in the Promoter Regions of CsCAMTA Genes in Tieguanyin

As shown in Figure 5, promoter analysis identified four major categories of cis-acting elements in the upstream regions of CsCAMTA genes, including hormone-responsive elements, stress-responsive elements, light-responsive elements, and growth/development-related elements. Hormone-responsive elements, such as ABRE, SARE, MeJA-responsive elements, GBRE, and AuxRE, suggest that CsCAMTA genes may be regulated by multiple hormone-related signals. Stress-related elements, including MYB-1, LTR, WUN, and ARE, were also detected in the promoter regions of several CsCAMTA genes, indicating their potential association with environmental stress responses. In addition, light-responsive and growth/development-related elements were widely distributed among CsCAMTA promoters. These results suggest that CsCAMTA genes may be transcriptionally regulated by diverse environmental and developmental signals, providing clues for further analysis of their potential roles in heat stress responses. These findings systematically revealed the multifunctional nature of the CsCAMTA gene family in plant environmental adaptation and growth/development, providing new directions for molecular mechanism research [35].

3.7. Expression Analysis of 18 CsCAMTA Genes Under Heat Stress

To further investigate the functions and expression patterns of CsCAMTA genes in Tieguanyin, we used Tieguanyin seedlings as experimental materials and subjected them to heat stress treatment at 40 °C. Samples were collected at 0 h, 4 h, 8 h, 12 h, and 24 h post-treatment, and quantitative real-time polymerase chain reaction (qRT-PCR) was employed to detect gene expression levels. A total of 18 representative CsCAMTA genes (CsCAMTA01–CsCAMTA08, CsCAMTA10–CsCAMTA19) were screened from the results.
As shown in Figure 6, the expression levels of these genes exhibited significant differences. Further analysis revealed that their expression patterns were diverse, mainly including rise first then decline, decline first then rise, and fluctuating patterns. For example, CsCAMTA1 and CsCAMTA4 were typical rise first then decline genes: their expression levels were significantly upregulated and reached peaks at 4 h post-treatment, followed by a gradual decrease. In contrast, CsCAMTA17 and CsCAMTA18 belonged to the decline first then rise type, with their expression levels peaking at 24 h post-treatment. Additionally, several special expression patterns were observed: CsCAMTA3 and CsCAMTA10 showed a decline-rise-decline fluctuating pattern, with expression turning points at 8 h and 12 h post-treatment; CsCAMTA7 exhibited a unique expression pattern, with no significant fluctuation in expression levels during 0–8 h post-treatment, followed by rapid downregulation during 12–24 h.
Combined with the significance labels in the bar charts of Figure 6, it can be seen that most CsCAMTA genes showed significant or extremely significant differences compared with the control group (CK) at least one time point after heat treatment. This indicates that these genes exhibit distinct temporal specificity in response to heat stress, and their expression dynamics are directly associated with the perception and transduction of stress signals.

3.8. GO Functional Enrichment and KEGG Pathway Analysis of the CsCAMTA Gene Family

In this study, Gene Ontology (GO) functional enrichment analysis and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway analysis were performed on CsCAMTA genes, and the results are shown in Figure 7. The GO classification system categorizes gene functions into three core domains: Molecular Function (blue), Cellular Component (orange), and Biological Process (yellow), with the −log10(p-value) indicating enrichment significance (higher values denote stronger enrichment). As shown in Figure 7A, in the Molecular Function category, CsCAMTA genes were most significantly enriched in the term DNA-binding transcription activator activity, RNA polymerase II-specific (−log10(p-value) ≈ 14), while terms including calmodulin binding, sequence-specific DNA binding, and DNA-binding transcription factor activity were also highly enriched. In the Cellular Component category, CsCAMTA gene products were primarily localized to terms such as nucleus and intracellular organelle, which aligns with the subcellular localization characteristics of transcription factors. In the Biological Process category, the most prominent term was positive regulation of gene expression, while transcription regulation-related terms such as positive regulation of macromolecule biosynthetic process and positive regulation of transcription by RNA polymerase II were also highly enriched. Additionally, stress response terms including cellular response to cold were significantly enriched, suggesting that this gene family is involved in plant adaptation to low-temperature stress [36].
As shown in Figure 7B, KEGG pathway enrichment analysis showed that Membrane trafficking was the most significantly enriched pathway (−log10(p-value) ≈ 3.0) [37]. In addition, pathways including Steroid biosynthesis, Protein phosphatases and associated proteins, and Protein families: genetic information processing were also relatively enriched.

4. Discussion

Tieguanyin is a representative oolong tea cultivar in China, and its growth, quality formation, and stress resistance are influenced by both genetic background and environmental conditions [12,38,39,40,41]. The CAMTA gene family encodes a group of highly conserved calmodulin-binding transcription factors that are involved in calcium signal transduction, stress responses, plant development, hormone regulation, DNA binding, and transcriptional regulation [42,43,44]. In the present study, 20 CsCAMTA genes were identified from the Tieguanyin genome, and their phylogenetic relationships, physicochemical properties, conserved motifs, chromosomal distribution, collinearity, promoter cis-elements, GO/KEGG annotation, and heat-responsive expression patterns were systematically analyzed.
The 20 identified CsCAMTA genes were unevenly distributed across eight chromosomes, suggesting that this gene family was not uniformly retained in the Tieguanyin genome. The proteins encoded by these genes varied considerably in length, ranging from 164 to 1158 amino acids, and their predicted molecular weights ranged from 17.96 to 128.57 kDa. Such variation in protein length and molecular weight may be related to differences in domain composition among family members. The theoretical isoelectric points of CsCAMTA proteins ranged from 4.54 to 8.73, indicating differences in predicted charge properties. Most CsCAMTA proteins showed negative GRAVY values, suggesting an overall hydrophilic nature, whereas CsCAMTA17 and CsCAMTA18 had GRAVY values close to zero. Protein instability index analysis showed that several CsCAMTA proteins were predicted to be stable, whereas others were predicted to be unstable [45,46]. The relatively high instability index of several CsCAMTA proteins suggests that these proteins may have more dynamic structural properties, which could be associated with regulatory processes such as stress response and signal transduction [47]. Subcellular localization prediction indicated that CsCAMTA proteins may be distributed in the cytoplasm, nucleus, chloroplast, and plasma membrane. Among them, the predicted nuclear-localized members may be associated with transcriptional regulation, which is consistent with the general characteristics of transcription factors.
To further investigate the evolutionary relationships of CAMTA proteins, a phylogenetic tree was constructed using CAMTA members from Tieguanyin, soybean, tobacco, tomato, and Arabidopsis thaliana. Based on the phylogenetic classification, CAMTA proteins from the five species were divided into four subgroups [30,31,32,33]. Several CsCAMTA proteins clustered closely with CAMTA proteins from soybean, tobacco, tomato, or Arabidopsis, suggesting a certain degree of evolutionary conservation among CAMTA members from different plant species. For example, CsCAMTA11 and NtCAMTA16 were located in the same clade, indicating that these proteins may share relatively close evolutionary relationships. In addition, some CsCAMTA members formed relatively independent branches, suggesting possible sequence divergence within the Tieguanyin CAMTA gene family.
Subgroup classification provides useful clues for inferring the potential functions of CsCAMTA genes based on homologous genes in model plants. In Subgroup I, AtCAMTA3 has been extensively studied and has been reported to participate in plant immune responses and abiotic stress adaptation by regulating downstream target genes [12,13]. For example, AtCAMTA3 is involved in the regulation of EDS1, a key gene associated with immune signaling, and also participates in pathways related to reactive oxygen species (ROS) accumulation and salicylic acid (SA) signaling [12,48,49]. Therefore, Tieguanyin CsCAMTA members clustered with AtCAMTA3 in Subgroup I may have potential roles in stress-related regulatory processes. In Subgroup III, CsCAMTA6, CsCAMTA11, and CsCAMTA15 were clustered with AtCAMTA6. Previous studies showed that AtCAMTA6 is associated with salt stress responses and may participate in the regulation of sodium ion homeostasis and stress-related transcriptional networks [50]. Based on these phylogenetic relationships, CsCAMTA members in this subgroup may be associated with calcium- or hormone-mediated regulatory pathways.
Synteny analysis was performed to explore the evolutionary conservation of CAMTA genes between Tieguanyin and other plant species, including wheat, tobacco, and Arabidopsis. The results showed that Tieguanyin had the largest number of CAMTA homologous gene pairs with wheat, followed by tobacco, whereas fewer homologous gene pairs were detected between Tieguanyin and Arabidopsis. The relatively large number of homologous gene pairs between Tieguanyin and wheat may partly be related to the hexaploid nature of the wheat genome and the retention of homologous genes during genome evolution [51]. In addition, the syntenic relationships between Tieguanyin and tobacco suggest that some CAMTA genes may have been conserved during the evolution of dicotyledonous plants. Gene duplication is considered an important mechanism driving gene family expansion and functional diversification in plants [52,53]. In this study, intragenomic synteny analysis identified several segmental duplication relationships among CsCAMTA genes, suggesting that segmental duplication may have contributed to the expansion of the CsCAMTA gene family in Tieguanyin.
Gene structure and conserved domain analyses showed that most CsCAMTA proteins contained ANKYR domains [54]. The ANKYR domain is generally associated with protein–protein interactions and may contribute to the connection between calcium-related signaling and transcriptional regulation. In addition, members such as CsCAMTA3, CsCAMTA6, and CsCAMTA12 contained CaMBDs, which are calmodulin-binding regions associated with calcium/calmodulin-mediated regulation. The presence of these domains suggests that CsCAMTA proteins may retain the basic structural characteristics required for calcium/calmodulin-associated transcriptional regulation [55]. Notably, CsCAMTA14 was the only member containing a DHHC zinc finger domain, suggesting that this member may have a distinct structural feature compared with other CsCAMTA proteins. This domain composition provides useful information for further analysis of potential functional divergence within the CsCAMTA family [56,57].
Heat stress caused by global warming has become one of the major abiotic stresses limiting plant growth, development, and crop productivity [58]. Under heat stress conditions, plants regulate gene expression through complex molecular networks, in which transcription factors serve as important regulatory components [59]. In this study, qRT-PCR analysis was performed to examine the relative expression levels of 18 CsCAMTA genes after 40 °C heat treatment at 0 h, 4 h, 8 h, 12 h, and 24 h. The results showed that different members of the CsCAMTA gene family exhibited diverse temporal expression patterns, including early-induced, sustained-response, and late-induced patterns. For example, CsCAMTA10 was upregulated at 4 h and maintained relatively high expression levels at 8 h and 12 h, whereas CsCAMTA16 began to increase at 12 h and reached its highest expression level at 24 h. Statistical analysis showed that several CsCAMTA genes, such as CsCAMTA2, CsCAMTA10, and CsCAMTA16, exhibited significant expression changes compared with the control at one or more heat treatment time points. These expression patterns suggest that several CsCAMTA genes may represent candidate heat-responsive genes in Tieguanyin, which is consistent with previous findings that CAMTA genes in maize exhibit transcriptional responses under heat stress [60].
Plants have evolved complex molecular mechanisms to perceive and respond to heat stress, and transcription factors are important components of these regulatory networks [34,61,62]. Previous studies have shown that CAMTA genes participate in plant responses to various stresses, including cold, drought, salt, heat, and pathogen-related stresses. For example, TaCAMTA genes in wheat have been reported to respond to drought, cold, heat, and saline–alkali stresses [10,63]. Previous studies in maize showed that ZmCAMTA genes exhibit transcriptional responses under heat stress, supporting the potential involvement of CAMTA genes in plant heat stress responses [64]. These findings support the possibility that CAMTA genes may participate in stress responses through transcriptional regulation. Under heat stress, Ca2+ channels on the cell membrane can be activated, leading to transient changes in cytoplasmic Ca2+ concentration and the formation of calcium signals [65]. These signals are sensed by calcium sensor proteins, including calmodulin (CaM) [66]. After binding Ca2+, CaM undergoes conformational changes and can interact with downstream target proteins, including CAMTA transcription factors. Based on previous studies, CAMTA proteins may participate in calcium/calmodulin-mediated signaling and regulate downstream stress-related genes. In the proposed model shown in Figure 8, heat stress may activate Ca2+/CaM signaling, which is then connected with CAMTA-mediated transcriptional regulation and downstream responses such as antioxidant defense and heat shock protein-related pathways. This model provides a conceptual framework for understanding the potential involvement of CsCAMTA genes in heat stress responses, although the specific regulatory relationships in Tieguanyin require further functional investigation.
To explore the potential biological functions of the CsCAMTA gene family in Tieguanyin, GO functional enrichment and KEGG pathway analyses were performed. In the molecular function category, CsCAMTA genes were enriched in terms such as “DNA-binding transcription activator activity” and “calmodulin binding”, which is consistent with the typical characteristics of CAMTA transcription factors as calmodulin-binding transcriptional regulators [42,43,58,59]. Terms related to sequence-specific DNA binding and DNA-binding transcription factor activity were also enriched, suggesting that CsCAMTA genes may be associated with transcriptional regulation. In the biological process category, enriched terms included “positive regulation of gene expression” and “cellular response to cold”, indicating that CsCAMTA genes may be broadly involved in stress-related biological processes. Previous studies have shown that CAMTA genes in other plant species, such as rice, grape, and Arabidopsis, participate in stress-related regulatory pathways [19,67,68,69], which provides useful references for interpreting the potential roles of CsCAMTA genes in Tieguanyin.
Through KEGG pathway analysis, CsCAMTA genes were significantly enriched in pathways such as “Transcription”, “Membrane trafficking”, and “Steroid biosynthesis”. Among these, the enriched pathways may reflect potential links between CsCAMTA genes and transcriptional regulation, intracellular transport, and hormone-related processes; the coupling of Ca2+ signals and membrane transport is a crucial link in plant stress signal transduction. The enrichment in the “Steroid biosynthesis” pathway indicates that CsCAMTA may regulate the synthesis of steroid substances (brassinosteroids) in tea plants, and brassinosteroids are key hormones for stress resistance, growth, and development of tea plants [70].

5. Conclusions

In this study, a total of 20 CsCAMTA genes were identified in the Tieguanyin tea genome and systematically analyzed. These genes were unevenly distributed across eight chromosomes and showed differences in protein length, molecular weight, isoelectric point, hydropathicity, instability index, subcellular localization, conserved motifs, and domain composition. Phylogenetic and collinearity analyses suggested that CsCAMTA genes have experienced evolutionary conservation and possible gene duplication events during the expansion of the CAMTA gene family in Tieguanyin. Promoter cis-acting element analysis and GO/KEGG annotation further indicated that CsCAMTA genes may be associated with hormone signaling, transcriptional regulation, and stress-related regulatory processes. In addition, qRT-PCR analysis of 18 representative CsCAMTA genes under heat treatment revealed distinct temporal expression patterns, suggesting that several CsCAMTA members may act as candidate heat-responsive genes in Tieguanyin. Overall, this study provides a systematic overview of the CsCAMTA gene family and offers useful candidate genes for future studies on heat stress responses in Tieguanyin tea plants.

Author Contributions

Conceptualization, Z.C., K.Z. and D.L.; investigation, Z.Y.; methodology, Z.C. and K.Z.; software, Z.C. and B.X.; data curation, Z.C., H.W. and K.Z.; writing—original draft preparation, Z.C., H.W., B.X. and Z.Y.; writing—review and editing, Z.C., H.W., Z.Y., B.X., F.J., K.Z., R.L., L.H., C.L. and D.L.; funding acquisition, D.L. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Central Finance Forestry Science and Technology Extension Demonstration Project (grant No. JXTG [2026] 05). The APC was funded by the Central Finance Forestry Science and Technology Extension Demonstration Project (grant No. JXTG [2026] 05).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The data presented in this study are openly available in publicly accessible repositories. The original genome sequence data of Tieguanyin tea plant (Camellia sinensis cv. Tieguanyin) are available in the National Center for Biotechnology Information (NCBI) under accession number JAFLEL000000000 and in the Genome Warehouse (GWH) at https://bigd.big.ac.cn/gwh/ (accessed on 3 April 2024) under the accession number GWHASIV00000000. The qRT-PCR expression data generated during this study are available from the corresponding author upon reasonable request.

Acknowledgments

The authors thank the Forestry Technology Program of Jiangxi Environmental Engineering Vocational College for supporting this work.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Phylogenetic analysis of CAMTA proteins in Tieguanyin (Camellia sinensis cv. Tieguanyin), soybean (Glycine max), tobacco (Nicotiana tabacum), tomato (Solanum lycopersicum), and Arabidopsis thaliana. The inner circle indicates CAMTA proteins from different species using different colors, whereas the outer colored blocks represent the four CAMTA subgroups. Branches indicate phylogenetic relationships among CAMTA proteins. Purple dots at internal nodes indicate bootstrap support values based on 1000 replicates, with larger dots representing higher support values.
Figure 1. Phylogenetic analysis of CAMTA proteins in Tieguanyin (Camellia sinensis cv. Tieguanyin), soybean (Glycine max), tobacco (Nicotiana tabacum), tomato (Solanum lycopersicum), and Arabidopsis thaliana. The inner circle indicates CAMTA proteins from different species using different colors, whereas the outer colored blocks represent the four CAMTA subgroups. Branches indicate phylogenetic relationships among CAMTA proteins. Purple dots at internal nodes indicate bootstrap support values based on 1000 replicates, with larger dots representing higher support values.
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Figure 2. Circos diagram of CsCAMTA genes. The outer ideograms illustrate the tea plant chromosomes, with red peaks indicating the distribution of gene density along each chromosome. Black lines connect collinear CsCAMTA gene pairs, representing segmental duplication events within the Tieguanyin genome.
Figure 2. Circos diagram of CsCAMTA genes. The outer ideograms illustrate the tea plant chromosomes, with red peaks indicating the distribution of gene density along each chromosome. Black lines connect collinear CsCAMTA gene pairs, representing segmental duplication events within the Tieguanyin genome.
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Figure 3. Structural analysis of the CsCAMTA gene family in Tieguanyin. (A) Phylogenetic relationships of CsCAMTA proteins. (B) Conserved motif distribution of CsCAMTA proteins. (C) Conserved domain organization of CsCAMTA proteins. (D) Exon–intron structures of CsCAMTA genes.
Figure 3. Structural analysis of the CsCAMTA gene family in Tieguanyin. (A) Phylogenetic relationships of CsCAMTA proteins. (B) Conserved motif distribution of CsCAMTA proteins. (C) Conserved domain organization of CsCAMTA proteins. (D) Exon–intron structures of CsCAMTA genes.
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Figure 4. Collinearity analysis of CAMTA genes between Camellia sinensis Cv. Tieguanyin and other plant species. (A) Collinearity analysis of CAMTA genes between Camellia sinensis and Arabidopsis thaliana. (B) Collinearity analysis of CAMTA genes between Camellia sinensis and Triticum aestivum. (C) Collinearity analysis of CAMTA genes between Camellia sinensis and Zea mays. (D) Collinearity analysis of CAMTA genes between Camellia sinensis and Nicotiana tabacum. Different species names and chromosomes are represented by different colors. The red lines (in panel (A)) and blue lines (in panels (B–D)) represent the homologous CAMTA gene pairs between the corresponding species and the tea plant CAMTA genes (CsCAMTAs), and the gray lines represent all other homologous gene pairs on the chromosome.
Figure 4. Collinearity analysis of CAMTA genes between Camellia sinensis Cv. Tieguanyin and other plant species. (A) Collinearity analysis of CAMTA genes between Camellia sinensis and Arabidopsis thaliana. (B) Collinearity analysis of CAMTA genes between Camellia sinensis and Triticum aestivum. (C) Collinearity analysis of CAMTA genes between Camellia sinensis and Zea mays. (D) Collinearity analysis of CAMTA genes between Camellia sinensis and Nicotiana tabacum. Different species names and chromosomes are represented by different colors. The red lines (in panel (A)) and blue lines (in panels (B–D)) represent the homologous CAMTA gene pairs between the corresponding species and the tea plant CAMTA genes (CsCAMTAs), and the gray lines represent all other homologous gene pairs on the chromosome.
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Figure 5. Distribution of cis-acting elements in the promoters of CsCAMTA genes in Tieguanyin. The legend on the right corresponds to the element types.
Figure 5. Distribution of cis-acting elements in the promoters of CsCAMTA genes in Tieguanyin. The legend on the right corresponds to the element types.
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Figure 6. Expression patterns of 18 Tieguanyin CsCAMTA genes under 40 °C heat treatment. qRT-PCR was used to analyze the relative expression levels of CsCAMTA genes at 0, 4, 8, 12, and 24 h. Statistical significance was determined by one-way ANOVA. Asterisks indicate significant differences compared with the control: * p ≤ 0.05; ** p ≤ 0.01; *** p ≤ 0.001; **** p ≤ 0.0001.
Figure 6. Expression patterns of 18 Tieguanyin CsCAMTA genes under 40 °C heat treatment. qRT-PCR was used to analyze the relative expression levels of CsCAMTA genes at 0, 4, 8, 12, and 24 h. Statistical significance was determined by one-way ANOVA. Asterisks indicate significant differences compared with the control: * p ≤ 0.05; ** p ≤ 0.01; *** p ≤ 0.001; **** p ≤ 0.0001.
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Figure 7. GO enrichment (A) and KEGG analysis of CsCAMTA genes (B).
Figure 7. GO enrichment (A) and KEGG analysis of CsCAMTA genes (B).
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Figure 8. Proposed model of CsCAMTA-mediated responses to heat stress in Tieguanyin. The dashed lines divide the diagram into different cellular or functional regions, including cytoplasmic signaling, nuclear transcriptional regulation, and downstream stress-response processes. I–IV indicate the major steps of the proposed model: (I) heat stress-induced Ca2+/CaM signaling activation; (II) interaction between CAMTA proteins and DNA-related regulatory regions; (III) CAMTA-associated regulation of heat stress-responsive genes; and (IV) downstream responses involving antioxidant defense and heat shock protein-related pathways. Arrows indicate the putative direction of signal transduction or regulatory flow.
Figure 8. Proposed model of CsCAMTA-mediated responses to heat stress in Tieguanyin. The dashed lines divide the diagram into different cellular or functional regions, including cytoplasmic signaling, nuclear transcriptional regulation, and downstream stress-response processes. I–IV indicate the major steps of the proposed model: (I) heat stress-induced Ca2+/CaM signaling activation; (II) interaction between CAMTA proteins and DNA-related regulatory regions; (III) CAMTA-associated regulation of heat stress-responsive genes; and (IV) downstream responses involving antioxidant defense and heat shock protein-related pathways. Arrows indicate the putative direction of signal transduction or regulatory flow.
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Table 1. Identified CsCAMTA genes and their characteristics in Tieguanyin.
Table 1. Identified CsCAMTA genes and their characteristics in Tieguanyin.
Gene NameGene IDSize/aaMolecular Weight/kDaTheoretical PIGrand Average of HydropathicityInstability IndexSubcellular Localization
CsCAMTA1CsTGY01G000083947452.146.1−0.27344.15cytoplasm
CsCAMTA2CsTGY01G000146735237.944.54−0.53732.62nucleus
CsCAMTA3CsTGY01G000021591097123.045.29−0.65548.66nucleus
CsCAMTA4CsTGY01G0000592330033.474.95−0.59940.67chloroplast
CsCAMTA5CsTGY02G0000902044350.118.19−0.53245.91chloroplast
CsCAMTA6CsTGY02G000111251108124.535.9−0.56545.33nucleus
CsCAMTA7CsTGY02G0002116932737.316.72−0.35851.98cytoplasm
CsCAMTA8CsTGY02G0002117224927.867.62−0.05951.45chloroplast
CsCAMTA9CsTGY05G0000898548654.768.73−0.28735.25chloroplast
CsCAMTA10CsTGY05G00023727896101.386.66−0.47240.1nucleus
CsCAMTA11CsTGY05G00023086956106.347.37−0.49237.96nucleus
CsCAMTA12CsTGY06G000001131008112.715.88−0.54448.15nucleus
CsCAMTA13CsTGY06G0000604040046.828.27−0.56854.58chloroplast
CsCAMTA14CsTGY06G0001155364069.626.27−0.14834.2Plasma Membrane
CsCAMTA15CsTGY09G00000185939106.056.77−0.45842.82cytoplasm
CsCAMTA16CsTGY10G000018471158128.465.62−0.50547.35nucleus
CsCAMTA17CsTGY11G000178447050.896.390.0333.04cytoplasm
CsCAMTA18CsTGY11G000184731233.986.220.05525.75chloroplast
CsCAMTA19CsTGY11G000185016417.967.79−0.19120.35cytoplasm
CsCAMTA20CsTGY13G0000072054158.326.54−0.21842.42chloroplast
Table 2. Three segmental duplication groups of the CsCAMTA gene family in Tieguanyin.
Table 2. Three segmental duplication groups of the CsCAMTA gene family in Tieguanyin.
GroupGene ID
Group 1CsTGYCAMTA03
CsTGYCAMTA05
CsTGYCAMTA06
Group 2CsTGYCAMTA10
CsTGYCAMTA11
CsTGYCAMTA15
Group 3CsTGYCAMTA12
CsTGYCAMTA16
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Cui, Z.; Wu, H.; Yang, Z.; Xu, B.; Jiang, F.; Lai, R.; Han, L.; Lu, C.; Li, D.; Zheng, K. Genome-Wide Identification and Expression Analysis of the CsCAMTA Gene Family in Tieguanyin Tea Plants Under Heat Stress. Curr. Issues Mol. Biol. 2026, 48, 597. https://doi.org/10.3390/cimb48060597

AMA Style

Cui Z, Wu H, Yang Z, Xu B, Jiang F, Lai R, Han L, Lu C, Li D, Zheng K. Genome-Wide Identification and Expression Analysis of the CsCAMTA Gene Family in Tieguanyin Tea Plants Under Heat Stress. Current Issues in Molecular Biology. 2026; 48(6):597. https://doi.org/10.3390/cimb48060597

Chicago/Turabian Style

Cui, Zijia, Hua Wu, Zhicheng Yang, Bohao Xu, Fan Jiang, Rien Lai, Lu Han, Ciding Lu, Dandan Li, and Kehui Zheng. 2026. "Genome-Wide Identification and Expression Analysis of the CsCAMTA Gene Family in Tieguanyin Tea Plants Under Heat Stress" Current Issues in Molecular Biology 48, no. 6: 597. https://doi.org/10.3390/cimb48060597

APA Style

Cui, Z., Wu, H., Yang, Z., Xu, B., Jiang, F., Lai, R., Han, L., Lu, C., Li, D., & Zheng, K. (2026). Genome-Wide Identification and Expression Analysis of the CsCAMTA Gene Family in Tieguanyin Tea Plants Under Heat Stress. Current Issues in Molecular Biology, 48(6), 597. https://doi.org/10.3390/cimb48060597

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