Genome-wide identification and expression pattern analysis of the CCT genes of Akebia trifoliata

Genome-wide identification and expression pattern analysis of the CCT genes of Akebia trifoliata | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Genome-wide identification and expression pattern analysis of the CCT genes of Akebia trifoliata Dianxun Liu, Xiaoxiao Yi, Xueying Li, Huai Yang, Jie Li, Feiquan Tan, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9308920/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 6 You are reading this latest preprint version Abstract Flowering is one of the important developmental events for many plants, and the CONSTANS/CONSTANS-LIKE/TIMING OF CAB1 (CCT) genes play key roles in this process. We identified a total of 46 AktCCTs , which are widely distributed across all chromosomes of the Akebia trifoliata genome. Their promoter regions contain hormone- and environment-responsive cis-acting elements. The average length and number of exons were 1,787.2 bp and 5.2, respectively, and the average number of amino acids, molecular weight and isoelectric point of putative protein were 388.4, 43.0 kDa and 6.3, respectively. Evolutionarily, AktCCTs experienced whole or segmental genome duplication and purifying selection and were divided into four classes ( COL , PRR , CMF and GTCC ) on the basis of sequence similarity and conserved domains, among which the CMF had the lowest gene length, number of exons, number of amino acids and molecular weights. Functionally, some AktCCTs expressed a detectable level. Otherwise, the expression of the COL AktCCT17 significantly positively related with that of AktFT2, AktFT4, AktFT5 and AktGI1 , so that AktCCT17 may be the crucial gene regulating the flowering behavior of A. trifoliata by "GI-CO-FT" model. Overall, these results provide valuable insights into the potential roles of these genes in regulating the biological process of flowering. Akebia trifoliata flowering photoperiod CONSTANS/CONSTANS-LIKE/TIMING OF CAB1 (CCT) GI-CO-FT model Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Flowering is one of the most important biological behaviors in the plant kingdom, especially in angiosperms, which comprise approximately 300,000 extant species and dominate the vegetation of most terrestrial ecosystems (Crane et al. 1995 ). It not only has important ecological significance but also has substantial agricultural, horticultural and even ornamental value in practice (Wu et al. 2024 ). Differences in flowering time have an impact on flower morphology in addition to ensuring that plants are evolutionarily adapted to their specific geographical distribution and accomplish their life cycle to reproduce (Shan et al. 2019 ). However, flowering initiation is usually affected by environmental signals such as temperature and sunlight duration and endogenous changes such as developmental status and hormone levels (Liu et al. 2020 ), which are closely related to physiological processes such as photoperiod and vernalization (Liu et al. 2015 ). Therefore, detailing the characteristics of the genes involved in this process is important for understanding both the ecological distribution and flower development of a given plant species. Plant photoperiod that has been discovered by more one hundred years (Garner and Allard 1920 ), refers the behavior that flowering time is influenced by the comparative length of light and dark periods in the diurnal cycle (Sawa et al. 2007 ), and hence flowering plants are classified into three types according to their response to photoperiod: day-neutral, short-day and long-day (Lumsden et al. 2002). Various studies have suggested that CONSTANS/CONSTANS-LIKE/TIMING OF CAB1 ( CCT ) gene families are the core elements affecting the plant photoperiod (Abe et al. 2003 ; Takahashi. 2017; Liu et al. 2020 ; Awal et al. 2022). The proteins encoded by CCTs commonly contain a conserved domain consisting of 43–45 amino acids at the C-terminus (Putterill et al. 1995 ) and are commonly localized in the nucleus (Robson et al. 2001 ). In addition, variations widely exist among intraspecific and interspecific members (Chen et al. 2010 ), which provides important evidence for classifying into subfamilies. According to studies in Arabidopsis thaliana , Oryza sativa , Sorghum bicolor , and Hordeum vulgare (Cockram et al. 2012 ), the CCT family can be structurally divided into three subfamilies: CONSTANS-like (COL), containing one or two B-box domains; Pseudo response regulators (PRR), with a response-regulator (REC) domain beside a CCT domain; and the CCT motif (CMF), with only a CCT domain (Khanna et al. 2009 ; Cockram et al. 2012 ). In fact, a relatively special subclass called the GTCC subfamily containing the GATA domain also exists in some species such as Cajanus cajan and Brassica rapa , which sometimes accompanies a zinc finger protein expressed in the inflorescence meristem (Tribhuvan et al. 2022 ; Li et al. 2024 ). These findings indicate the high genetic diversity in the structure of the CCT family members among various species. Since CONSTANS (CO), a circadian clock regulatory gene belonging to the COL subfamily, was first cloned (Wenkel et al. 2006 ), various studies have functionally demonstrated that the COL subfamily could acts as the central regulatory factor in the photoperiodic pathway, and in the known "GI-CO-FT" model CO is first regulated by the upstream transcription factor named Gigantea (GI) and then it consequently regulates the downstream flowering locus T (FT) (Wu et al. 2014 ; Yang et al. 2014 ; Andrés et al. 2012). At the same time, some studies have also shown that COL genes have important functions in nuclear localization, protein‒protein interactions and transcriptional regulation (Putterill et al. 1995 ; Kaewphalug et al. 2017 ), suggesting that they may be DNA-binding transcription factors, but the CCTs are not listed as a family in the Plant Transcription Factor Database (v5.0) (Jin et al. 2017 ), indicating that at least some CCTs lack the typical DNA-binding motif of classical transcription factors. In addition, some studies in the green algae Chlamydomonas and A. thaliana have shown that the CCT domain, which is homologous to the CCAAT-binding domain, can interact with transcription factors such as NF-YA, NF-YB and NF-YC with the typical DNA-binding motif to form multimers (Gnesutta et al. 2017 ; Lv et al. 2021 ), which further supports the view that COL indirectly regulates the expression of downstream genes, mainly through a complex pathway rather than directly binding the promoter region of the target genes. Ben-Naim et al (Ben-Naim et al. 2006 ) suggested that COL proteins usually lack a known DNA-binding motif and that their success in transcriptional regulation occurs mainly through the mediation of CCAAT binding factors, but some studies have shown that some COL proteins can directly bind DNA in vitro studies (Tiwari et al. 2010 ). Obviously, the details of the regulatory function and corresponding molecular mechanism of COL are not completely understood. In addition, the CMF subfamily evolved from the COL subfamily, and thus, the functions of CMF families are very similar to those of COL subfamilies (Cockram et al. 2012 ). Compared with the COL and CMF subfamilies, the PRR subfamily is functionally more conserved and comprises important components of the circadian clock, usually interacting with circadian clock associated 1 (CCA1) and late elongated hypocotyl (LHY) (Makino et al. 2000 ; David et al. 2002; Nakamichi et al. 2020). Moreover, PRRs such as TOC1 can influence the expression of other transcription factors, such as CO , PIFs and GI , by directly binding to their promoter region and even exerting self-regulatory effects (Hsu et al. 2012 ; Gendron et al. 2012 ). Therefore, more datasets from various species of different evolutionary clades would helpful to completely elucidate the complex regulatory network of the CCT family. Akebia trifoliata (also called augmelon), which originates from the eastern edge of the Qinghai‒Tibet Plateau and is representative of the Lardizabalaceae family, is a climbing perennial woody vine that is narrowly distributed in several regions of East Asia, including subtropical China south of the Yellow River (Chen et al. 2022 ; Dong et al. 2024 ), so that it has largely been ignored by both scientists and entrepreneurs in the past. However, it is becoming a very popular topic in the field of edible wild plant domestication (Yu et al. 2021 ; Yang et al. 2024a ) because of its multiple purposes, such as in traditional medicine (Yang et al. 2024b ), for fresh fruit (Yang et al. 2021 , Yi et al. 2025 ), as a woody oil crop (Guan et al. 2024 ; Han et al. 2025 ) and even as an ornamental plant (Yang et al. 2023 ). In addition, its value for theoretical study is also becoming increasingly appreciated because of its small genome size (Zhong et al. 2022a ), short juvenile stage (Yang et al. 2021 ) and important evolutionary position (Zhong et al. 2022b ), and it has been widely employed in various studies, such as those involving flower development (Zhong et al. 2022c ; Zhong et al. 2024 ), phenylalanine metabolism (Guan et al. 2024 ; Li et al. 2024 ) and flavonoid biosynthesis (Zhong et al. 2022d ; Yang et al. 2024b ). However, to date, there are no reports about its photoperiod characteristics. In fact, the most important reasons for using A. trifoliata to investigate the close involvement of the CCT family with the photoperiod are that we observed that it flowers in both spring and autumn each year in the A. trifoliata Germplasm Nursery of Sichuan Agricultural University at Chongzhou Research Station (Chen et al. 2022 ), and it can even fruit in both March and September each year in the commercially cultivated garden (28° 19' N, 101° 07' E) of Jiulong County, China, which indicates that A. trifoliata might be a day-neutral plant. In addition, abundant available omics data, such as genomic (Zhong et al. 2022a ), transcriptomic (Chen et al. 2022 ) and metabolic (Zhong et al. 2022d ) data, provide a good opportunity to comprehensively characterize the CCT family profile. Therefore, our objectives are to systemically identify the number and components of the CCT family in the A. trifoliata reference genome, to completely detail its structural and evolutionary characteristics and to finally determine their expression levels. These results would be helpful for further understanding the molecular mechanism of flowering in A. trifoliata. Materials and Methods Identification CCT family in A. trifoliata Genome sequence annotation files of A. trifoliata (accession ID: GWHBISH00000000) were downloaded from the National Genomics Data Centre ( https://ngdc.cncb.ac.cn/ ; accessed on 9 September 2024). The hidden Markov model of conserved CCT sequences (PF06203) ( http://pfam-legacy.xfam.org/ ; accessed on 9 November 2024) was employed to identify AktCCTs . In addition, the sequences of all the proteins of A. trifoliata were then scanned by HMMER 3.0 using the HMM with an E value of 1×10 − 5 , in which the amino acid sequences of Arabidopsis CCTs were downloaded from the TAIR website ( https://www.Arabidopsis.org/ ; accessed 9 September 2024) and used to identify AktCCTs . Only those AktCCTs that could be identified by the two methods were treated as the final identified AktCCTs . Moreover, the physicochemical properties of the predicted AktCCTs were determined using the ExPASy ProtParam website ( https://web.expasy.org/protparam/ ; accessed on 9 November 2024). Conserved protein structure and chromosomal localization of AktCCT The MEME website ( https://meme-suite.org/meme/ ; accessed 9 September 2024) was used with the default parameters to analyze the conserved motifs in the putative AktCCTs. Subsequently, TBtools (version 1.0876, CJ Chen, Guangzhou, China) was utilized to map the AktCCTs on the chromosomes, adopting the gff file of A. trifoliata . Additionally, TBtools was used to extract parameters such as gene length and gene location. Calculation of Ka/Ks of homologous AktCCT pairs The nonsynonymous substitution (Ka) to synonymous substitution (Ks) values (Ka/Ks) were calculated using TBtools-II software (v2.012, Chengjie Chen, China). Similarly, we performed intraspecific collinearity analysis using TBtools software to determine the duplication type. Cis-acting elements of the AktCCT gene family To predict putative cis-acting elements in the promoter regions of the AktCCTs , the 2,000 bp upstream sequences of the AktCCT coding regions were analyzed using PlantCARE ( http://bioinformatics.psb.ugent.be/webtools/plantcare/html/ , accessed on 10 September 2024). After deleting unnecessary cis-acting elements, the distribution of the remaining cis-acting elements of the AktCCTs was plotted using Basic Biosequence View in TBtools. In addition, the cis-acting elements were divided into hormone- and environment-responsive types according to their annotation. Phylogenetic tree construction The phylogenetic tree of both all AktCCTs and 37 reference AtCCTs was constructed via the N‒J method with 1,000 bootstrap replicates using MEGA 11 software (v11.0.10, Auckland, New Zealand), in which 37 previously reported AtCCTs were downloaded from the Plant Transcription Factor Database website ( http://planttfdb.cbi.pku.edu.cn/ , accessed on 24 September 2024) (Tamura et al. 2021 ). In addition, the evolutionary relationships were constructed using Gene Structure View, and the online tool iTOL ( https://itol.embl.de/ , accessed on 9 October 2024) was used for subsequent visualization. Subcellular localization of the proteins putatively encoded by AktCCTs To predict the putative subcellular localization of the candidate proteins, in silico analyses were performed using multiple computational tools. Amino acid sequences were submitted to the following online prediction servers: WoLF PSORT ( https://wolfpsort.hgc.jp/ ) for comprehensive prediction based on sorting signals and amino acid composition; Plant-mPLoc ( http://www.csbio.sjtu.edu.cn/bioinf/plant-multi/ ) for plant-specific multi-label localization; and TargetP 2.0 ( http://www.cbs.dtu.dk/services/TargetP/ ) for the identification of N-terminal sequences and prediction of mitochondrial, chloroplast, secretory pathway, or other localizations. Differential expression of CCT in A. trifoliata Transcriptome data for A. trifoliata leaves, sepals and stamens during flowering period were downloaded from the NGDC database (login number: PRJCA019245). Additionally, HISAT2 software (v2.1.0, Mihaela Pertea, USA) and DESeq2 (v1.36.0, Michael I. Love, Germany) were used to extract the FPKM values from the transcriptomic data. We determined the expression of AktCCTs in these 3 tissues using HeatMap. Further mapping of the correlations between AktCCTs and both AktGIs and AktFTs was performed using OriginPro (2024 (64-bit) SR1 10.1.0.178). Statistical analysis The significance of the differences in the characteristic parameters among the various groups was determined using SPSS (version 19.0; Armonk, NY: IBM Corp.), and correlation analysis of the expression levels was also performed using the same software. Results Chromosomal distribution and physiochemical characteristics of the CCT family in A. trifoliata A total of 46 CCT genes were identified in the A. trifoliata reference genome, and they were orderly named AktCCT1 to AktCCT46 (Table 1 , Table S1 ). Physically, all 46 AktCCTs were unevenly distributed all 16 chromosomes and most of them were mapped to regions near chromosome ends (Fig. 1 ), in which chromosome 3 had the most 7 AktCCTs while chromosomes 9, 13, 15 and 16 each had only one. According to the definition of gene clusters, the genes were assigned to 41 loci on the chromosomes, including 36 singletons and 5 gene clusters within 250-kb chromosomal regions, in which each cluster consisted of two AktCCTs ( AktCCT3 / AktCCT4 , AktCCT13 / AktCCT14 , AktCCT15 / AktCCT16 , AktCCT19 / AktCCT20 , and AktCCT26 / AktCCT27 ). Note The genes in red are distributed in the cluster, and the purple lines represent collinearity between the AktCCTs. Main evolutionary experience of the AktCCTs We found that 33 (71.7%) AktCCTs were produced by whole-genome duplication (WGD) or segmental duplication while and the remaining 13 (28.3%) AktCCTs including all AktCCTs on chromosomes 9, 15 and 16 were produced by dispersal duplication (Table 1 ) . Interestingly, all 10 AktCCTs in cluster were produced by WGD or segmental duplication. Table 1 CCTs identified in A. trifoliata. Name Genes family Characteristics of putative proteins DT CP Begin End GL(bp) NE NAA MW (kDa) PI SL AktCCT1 1 28523241 28526070 2829 5 436 49.1 8.1 N D AktCCT2 1 31772454 31779638 7184 6 408 44.9 5.8 N W/S AktCCT3 2 5902189 5912295 10106 11 542 40.1 4.9 N W/S AktCCT4 2 5976398 5978442 2044 7 296 31.9 9.1 N W/S AktCCT5 3 1015390 1019400 4010 4 340 38.6 6.0 N W/S AktCCT6 3 2575299 2579508 4209 3 292 33.7 7.1 N W/S AktCCT7 3 11202320 11204761 2441 2 382 43.2 5.8 N D AktCCT8 3 25181097 25189683 8586 4 481 54.8 5.8 N D AktCCT9 3 44086815 44102632 15817 13 422 45.5 5.0 N W/S AktCCT10 3 51615841 51624543 8702 8 712 78.9 6.5 N W/S AktCCT11 3 53009727 53014571 4844 7 282 32.0 4.8 N W/S AktCCT12 4 6504460 6511344 6884 5 400 44.5 4.4 N D AktCCT13 4 10441408 10453360 11952 12 347 38.3 4.9 N W/S AktCCT14 4 10456733 10466278 9545 7 254 27.5 8.2 N W/S AktCCT15 4 40953416 40973777 20361 11 341 37.6 5.1 N W/S AktCCT16 4 41035410 41059349 23939 7 310 33.9 5.0 N W/S AktCCT17 5 955460 960517 5057 4 480 54.1 5.1 N W/S AktCCT18 5 9772961 9774777 1816 2 329 37.3 5.4 N W/S AktCCT19 6 5801167 5811188 10021 9 767 83.6 6.3 N W/S AktCCT20 6 5976398 5978442 2044 3 268 30.6 5.6 N W/S AktCCT21 7 32202888 32210121 7233 8 558 63.2 6.2 N D AktCCT22 7 33382781 33385175 2394 2 362 40.0 5.1 C W/S AktCCT23 8 2406849 2408961 2112 3 226 26.0 7.7 N W/S AktCCT24 8 24582546 24584147 1601 3 217 25.9 5.1 N D AktCCT25 8 49018032 49020644 2612 2 381 42.2 5.5 C W/S AktCCT26 8 49876689 49879917 3228 2 403 45.7 8.6 N W/S AktCCT27 8 49976675 49987459 10784 7 317 35.2 7.2 N W/S AktCCT28 9 18517830 18520944 3114 2 414 45.6 5.4 C D AktCCT29 10 4329166 4330869 1703 2 359 39.6 5.6 C W/S AktCCT30 10 6725436 6742598 17162 8 270 29.3 6.8 N W/S AktCCT31 10 7217500 7220765 3265 3 389 44.5 8.7 N W/S AktCCT32 10 29942599 29944927 2328 4 320 36.5 5.1 N W/S AktCCT33 10 30988975 30995773 6798 4 289 32.6 6.1 N W/S AktCCT34 11 8023901 8027094 3193 4 259 29.6 5.0 N D AktCCT35 11 11833091 11837383 4292 4 282 32.0 6.3 N W/S AktCCT36 11 23047610 23048741 1131 3 224 26.6 4.9 N D AktCCT37 11 30582722 30584224 1502 2 156 17.9 9.6 N D AktCCT38 12 34672988 34674880 1892 4 322 36.9 5.8 N W/S AktCCT39 12 35707409 35711202 3793 4 274 30.9 5.0 N W/S AktCCT40 12 36585577 36592109 6532 7 727 81.0 5.6 N W/S AktCCT41 12 36873128 36878636 5508 8 703 78.6 5.8 N W/S AktCCT42 13 351777 358812 7035 5 414 45.4 5.4 N W/S AktCCT43 14 2520450 2525582 5132 7 722 80.5 8.2 N D AktCCT44 14 4925216 4932511 7295 9 725 80.0 8.7 N W/S AktCCT45 15 7240017 7240459 442 2 61 7.4 11.4 N D AktCCT46 16 29040983 29045522 4539 4 404 45.3 5.3 N D Note : CP: chromosomal position; GL: gene length; NE: number of exons; DT: duplication type; NAA: number of amino acids; MW: molecular weight; PI: isoelectric point; SL: subcellular localization; N: nuclear; C: chloroplast; D: dispersed; W/S: WGD/segmental. In addition, 27 orthologous pairs related with 33 AktCCTs were detected, among which 8 AktCCTs ( AktCCT3 , AktCCT7 , AktCCT11 , AktCCT14 , AktCCT16 , AktCCT29 , AktCCT34 and AktCCT41 ) had 3 orthologous gene pairs while 20 AktCCTs ( AktCCT2 , AktCCT 8 , AktCCT10 , AktCCT12 , AktCCT15 , AktCCT17 , AktCCT18 , AktCCT19 , AktCCT20 , AktCCT23 , AktCCT25 , AktCCT26 , AktCCT27 , AktCCT30 , AktCCT31 , AktCCT32 , AktCCT33 , AktCCT39 , AktCCT42 and AktCCT44 ) only had one orthologous gene pair. The Ka/Ks values varied from 0.15 ( AktCCT30 and AktCCT25 ) to 0.57 ( AktCCT40 and AktCCT38 ), and the average mean Ka/Ks was only 0.33 ( Table S2 ). Conserved structure, class and phylogenetic tree of the AktCCTs We identified a total of 10 conserved motifs (from 1 to 10) with a varying width from 11 amino acids in Motif 6 to 50 amino acids in five motifs (Motif 2, 4, 5, 7 and 9) among 46 putative AktCCTs (Table 2 ), and we also identified five conserved domains (CCT, B-box, GATA, TIFY and REC) (Fig. 2 ; Table 2 ). By sequence comparing, we found that CCT, B-box and REC domain distributed across Motif 1 and Motif 6, Motif 2 and Motif 8, and Motif 5 and Motif 7, respectively, while GATA and TIFY only distributed within Motif 3 and Motif 4, respectively. On one hand, two AktCCTs (AktCCT6 and AktCCT26) had only Motif 1 while 31 AktCCTs had the most 4 motifs ( Table S3 ). On the other hand, Motif 1 distributed all 46 AktCCTs while both Motif 9 and Motif 10 only existed in five AktCCTs. Based on the existence of sequence encoding conserved domain, we classified 46 AktCCTs into four classes: 18 CMFs , 12 COLs , 9 GTCCs , and 7 PRRs . Many characters such as gene whole length and the exon number, and amino acids number and molecular weight of putative protein existed significant difference among various classes at p = 0.05 level. For example, the COL class had the lowest average gene length, number of exons while GTCC class had the lowest number of amino acids and molecular weight of putative protein. On the contrary, PRR had the greatest average gene length, exon number, amino acid number and molecular weights (Table 3 ). In addition, the phylogenetic tree of the 46 AktCCTs was constructed according to the sequence component, in which 37 reference AtCCTs were also employed as the control. According to the phylogenetic tree, 46 AktCCTs were assigned into four distinct branches: Ⅰ, Ⅱ, Ⅲ and Ⅳwith 8, 9, 18 and 11 AktCCTs , respectively (Fig. 3 ), and the clustering results of 37 AtCCTs were well agreed by the previously reported classification (Huang et al. 2024 ). Interestingly, all 7 PRRs and 9 GTCCs were completely assigned intoⅠandⅡbranch, respectively, and similarly, all 18 CMFs except of AktCCT45 and 12 COLs except of AktCCT28 were assigned into Ⅲ and Ⅳ, respectively, which indicated that some DNA sequence beside that encoded conserved domain would be also experienced a fast evolution. Table 2 Sequence annotation of motifs of 46 putative AktCCTs Domain Motif Sequences Width CCT Motif 1 REAR LL RYREKRK ERN FEK K IRY A SRK ALAD RRP 34 B-box Motif 2 C EF C EEAPAVIY C KADAAAL C LT C DRNI H SANPLSRR H ERTPJCPICGSQ 50 GATA Motif 3 CQHCGISEKSTPMMRRGPAGPRT LCNACGL MWANKGT L RD L 41 TIFY Motif 4 GTDZ LT LS F ZGEV Y VFDAV SPEKVQA VL LLLG GYEIPSAVPTIDIPSHNN 50 REC Motif 5 HEICKNI PVIMM SSHDSVGVVFKCMLK GAADFLVKP VRKNELRNLWQHVW 50 CCT Motif 6 RV KG R F VK A KE 11 REC Motif 7 LVESD DS TRQIVSALLRK CGYEVTAVADGLKAWEILKGRSH NIDLV LTEV 50 B-box Motif 8 PASVY C RADSAYL C QB C DWNI H GANTLA 28 Motif 9 IQKSISSGCLSSMDWIHGGAMRPNFLDFQGVDFEAAYGMRRAFSEGDIQT 50 Motif 10 GDLFKAPEPIIEEPTIGLDPMSAAISMISCGEDVISPQTMKV 42 Note : the part in bold for conserved domain. Table 3 Multiple comparative analyses of the protein physicochemical properties of the AktCCTs in different subgroups Group GL NE NAA MW (kDa) PI max min mean max min mean max min mean max min mean max min mean COL 8586 1703 6065.5 c 13 2 5.28 bc 480 320 388.4 bc 54.1 36.5 43.0 ab 8.6 5 6.28 a GTCC 23939 2044 6498.4 ab 12 2 5.54 c 542 296 368.5 c 44.5 31.9 40.4 b 9.05 4.37 6.31 a CMF 6884 442 6227.2 bc 8 2 5.39 ab 727 61 393.9 ab 81 7.4 43.6 ab 11.4 4.9 6.22 a PRR 10021 5132 6626.3 a 9 8 5.60 a 767 274 405.1 a 83.6 30.9 45.3 a 8.71 4.76 6.24 a Note : GL, NE, NAA, MW and PI represent the gene length, number of exons, number of amino acids, molecular weight and isoelectric point, respectively. a, b and c indicate significance at the p = 0.05 level. Cis-acting components of the AktCCTs By examining the 2,000 bp upstream regions of the AktCCTs , we identified a total of 1,220 functional cis-acting elements among all the AktCCTs , among which 530 and 690 cis-acting elements belong to 13 hormone- and 15 environment-responsive types, respectively ( Table S4 ). Further analysis revealed that among the 13 kinds of hormone-responsive cis-acting elements, the number of cis-acting regulatory elements involved in MeJA responsiveness (MeJAR) and the number of cis-acting regulatory elements involved in root specificity (CRERS) were the greatest (193) and the lowest (1), respectively, and the sole CRERS was in the upstream region of AktCCT17 . Similarly, among the 15 kinds of environment-responsive cis-acting elements, the number of light-responsive elements (LRMs) and the number of MYB binding sites involved in drought inducibility (MYBBS(D)) and the number of wound-responsive elements (W-Rs) were the greatest (257) and least (2), respectively. In addition, the number of cis-acting elements in each AktCCT exhibited large variation. For example, there were up to 59 (34 hormone-responsive types and 25 environment-responsive types) cis-acting elements in AktCCT8 , whereas there were only 9 (4 hormone-responsive types and 5 environment-responsive types) cis-acting elements in AktCCT1. Note CEIAAR: cis-acting element involved in abscisic acid responsiveness; CEICCR: cis-acting element involved in cell cycle regulation; CEIDSR: cis-acting element involved in defense and stress responsiveness; CEIGR: cis-acting element involved in gibberellin responsiveness; CEILR: cis-acting element involved in light responsiveness; CEILTR: cis-acting element involved in low-temperature responsiveness; CEISAR: cis-acting element involved in salicylic acid responsiveness; CREEAI: cis-acting regulatory element essential for anaerobic induction; CREIAR: cis-acting regulatory element involved in auxin responsiveness; CREICC: cis-acting regulatory element involved in circadian control; CREIEE: cis-regulatory element involved in endosperm expression; CREILR: cis-acting regulatory element involved in light responsiveness; CRERS: cis-acting regulatory element, root specific; DPMC: element involved in the differentiation of palisade mesophyll cells; EFMEMA: element for maximal elicitor-mediated activation (2 copies); ELASI: enhancer-like element involved in anoxic-specific inducibility; GRE: gibberellin-responsive element; LRM: light-responsive element; MeJAAI: MeJAR essential for anaerobic induction; MeJAR: cis-acting regulatory element involved in MeJA-responsiveness; MYBBS(D): MYB binding site involved in drought-inducibility; MYBBS(FBGR): MYB binding site involved in flavonoid biosynthetic genes regulation; MYBBS(LR): MYB binding site involved in light responsiveness; PCMLR: part of a conserved DNA module involved in light responsiveness; PLRE: part of a light response element; PLRM: part of a light-responsive module; W-R: wound-responsive element; ZMR: cis-acting regulatory element involved in zein metabolism regulation. Expression pattern analysis of the CCT family genes of A. trifoliata The transcript levels of all 46 AkCCTs were analyzed in various tissues (leaves, sepals and stamens) at different developmental stages during flowering. The results showed that although the majority of AktCCTs had relatively low expression, but 19 AktCCTs had a detectable expression level in leaf. AkCCT27 exhibited high expression across all tissues during whole period (Fig. 5 ) . In addition, AkCCT26 was highly expressed at sepals and stamens. Previous studies showed that FT and GI were involved in the CO related pathway (Mizoguchi, 2005), and they should also be used to determine the expression profile of the AktCCTs in the known "GI-CO-FT" model, so that ee further identified two AktGIs ( AktGI1 and AktGI2 ) and seven AktFTs (from AktFT1 to AktFT7 ) ( Table S5 ). In addition, we found that AktFT2 and AktFT4 was highly expressed in leaf and that the expression of both AktGIs was very low in all tissues at all time points, but the pression of AktGI1 was detectable ( Fig. 5 ). Interestingly, AktFT2, AktFT4 and AktFT5 as well as AktGI1 had significantly co-expressed relationship with many AktCCTs such as AktCCT17 (Fig. 6 ). Note Red represents a positive correlation, blue represents a negative correlation, and the darker the colour, the stronger the correlation. * for the significance at the level of p = 0. 05. Discussion To adapt to seasonal and environmental changes, plants have specifically evolved a regulatory mechanism that optimizes developmental events via the light response pathway (Takagi et al. 2023 ), in which CCT family members play very important roles in various processes, such as flowering, circadian rhythms and even the abiotic stress response (Liu et al. 2020 ). This suggests that the structure and function of CCTs may have experienced rapid variation and divergence during evolution. In the present study, we found that among the 46 AktCCTs , many physiochemical parameters, such as gene length, number of exons, number of amino acids, and the molecular weight and isoelectric point of the putative proteins, differed substantially (Table 1 ). In fact, structural diversity may be favored by various evolutionary experiences. Previous studies have demonstrated that WGD is an important evolutionary force in almost all organisms, especially plants, and moreover, WGD is usually related to different clades (Jiao et al. 2011 ). For example, a recent report suggested that θ WGD and ω WGD favored the survival of eudicots and the thriving of core eudicots around the Jurassic-Cretaceous boundary and the Middle Cretaceous, respectively (Zhong et al. 2022c ). Here, 33 (71.7%), including all 10 AktCCTs found in gene clusters out of the 46 AktCCTs , were produced by WGD or segmental duplication (Table 1 ), which could have presented an important opportunity for numerous variations in AktCCTs to evolve. In addition, the small Ka/Ks value suggested that AktCCTs might have undergone purifying selection during evolution and maintained a conserved function ( Table S2 ), which helps A. trifoliata quickly eliminate damaged mutants, especially individuals whose light-responsive pathways are disrupted (Wang et al. 2011 ; Hsu et al. 2014). The chloroplast is the most important organ related to the light-responsive process (Schwenkert et al. 2022 ), and interestingly, all 4 AktCCTs ( AktCCT22 , AktCCT25 , AktCCT28 and AktCCT29 ), encoding proteins localized in the chloroplast, only had two exons and they belonged to the COL type. (Fig. 3 ; Table 1 ). Moreover, their promoters contained both MeJAR hormone-responsive and LRM environment-responsive cis-acting elements (Fig. 4 ; Table S5 ). On the other hand, their compartmental distribution and cis-acting element component indicated that they may retain the basic function of responding to changes in light, further suggesting that the AktCCTs might be older. In addition, on the basis of the knowledge of CCTs , the COL subfamily is the oldest, whereas other subfamilies, especially CMF , are usually the evolved offspring of COLs . Therefore, how COLs produced other subfamilies is one of our concerns. In total, the most distinct difference between the COLs and other types was the existence of a B-box (Cockram et al. 2012 ; Tribhuvan et al. 2022 ; Li et al. 2024 ). In this study, we found that among the various subfamilies, the gene length, number of exons, number of amino acids and molecular weight of the PRR members were the largest. In contrast, the number of amino acids and molecular weight of GTCC were the smallest while the gene length and exon number of COL was the smallest (Table 3 ). Based on this information, we propose that the CMFs could have been produced from COLs mainly by losing the DNA regions putatively encoding the B-box conserved domain, whereas the PRRs and GTCCs might have been produced mainly via the insertion of new DNA sequences into the encoding and non-encoding regions of the corresponding B-box sequence, respectively. Various studies have suggested that CO or COL can directly regulate the key downstream flowering gene FT by altering its status and that this process is regulated by the upstream gene GI; thus, a known "GI-CO-FT" regulatory model was proposed (Fowler et al. 1999 ; David et al. 2006 ; Sawa et al. 2007 ; Imaizumi et al. 2010). From the perspective of the system phylogenetic tree, most AktCCTs such as AktCCT17 were distributed into the Ⅳ branch that contained the key Arabidopsis flowering regulatory gene CO ( At5g15840 ) (Figs. 3 ). Otherwise, two AktCCTs ( AktCCT17 and AktCCT27 ) obviously upregulate in leaves (Fig. 5 , Fig. 6 ). Both AktCCT17 and AktCCT27 exhibited a significant co-expression with AktFT2, AktFT4, AktFT5 and AktGI1 involved in "GI-CO-FT" model, but the unique cis-acting elements binding with the sole root-related CRERS transcription factor existed in AktCCT17 ( Table S4 ). Moreover, previous studies showed that flowering is highly related with root behavior (Zou et al. 2020 ), so that AktCCT17 could play a key role in flowering of A. trifoliata . Overall, genome duplication events, especially WGD, afforded an important opportunity for the diversification of the AktCCTs , and at the same time, strong purifying selection facilitated their evolution toward favorable environmental adaptation. In addition, the sequences encoding conserved domains and the other sequences in the AktCCTs may have evolved in parallel to further enrich the structural and functional diversity. Finally, AktCCT17 may play a very important role in regulating flowering. Declarations Conflict of Interest The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as potential conflicts of interest. Supplementary Material The Supplementary Material for this article can be found online. Funding This work was supported by grants from the Natural Science Foundation of Sichuan Province, grant number 2024NSFSC1310; Regional Innovation Cooperation of Sichuan Province, grant number 2024YFHZ0109; and Central guidance for local regional system construction, grant number 2024ZYD0284; Central Guidance for Local Areas - Construction of the Regional System 2024ZYD0284. Author Contribution DL: formal analysis, Software, Writing - Original Draft - Visualization. XY, HY, YX, CC, FT, YH and JL: Obtained the data - Participated in the data analysis, Discussion. XL: Proofreading the language of the manuscript. PL: Conceptualization, Writing - Review and Editing, Project - Administration, Funding Acquisition. All the authors contributed to the discussion of the results, reviewed the manuscript and approved the final article. Details of all funding sources should be provided, including grant numbers if applicable. Please ensure to add all necessary funding information, as after publication this is no longer possible. Data Availability The datasets presented in this study can be found in online repositories. The names of the repositories/repositories and accession number(s) can be found in the article/Supplementary Material. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9308920","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":628305628,"identity":"de306d95-a92d-43f0-9762-307afa3e6951","order_by":0,"name":"Dianxun Liu","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Dianxun","middleName":"","lastName":"Liu","suffix":""},{"id":628305629,"identity":"fb4116fd-1ba2-4d2b-9fdd-fe6c0a614724","order_by":1,"name":"Xiaoxiao Yi","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Xiaoxiao","middleName":"","lastName":"Yi","suffix":""},{"id":628305630,"identity":"de618a0e-5140-48df-9fb1-4b829c4d3239","order_by":2,"name":"Xueying Li","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Xueying","middleName":"","lastName":"Li","suffix":""},{"id":628305631,"identity":"716bc5f7-a196-44e7-ad60-624d641c61d6","order_by":3,"name":"Huai Yang","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Huai","middleName":"","lastName":"Yang","suffix":""},{"id":628305632,"identity":"e55262a2-0a66-4a46-a615-00c60e0780ae","order_by":4,"name":"Jie Li","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Jie","middleName":"","lastName":"Li","suffix":""},{"id":628305633,"identity":"1206676f-0315-4cdd-8ddf-8c4a97749969","order_by":5,"name":"Feiquan Tan","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Feiquan","middleName":"","lastName":"Tan","suffix":""},{"id":628305634,"identity":"531e4ad4-b99a-4dae-90e5-5367260ac13c","order_by":6,"name":"Chen Chen","email":"","orcid":"","institution":"","correspondingAuthor":false,"prefix":"","firstName":"Chen","middleName":"","lastName":"Chen","suffix":""},{"id":628305636,"identity":"ce31fa19-fcb8-4d85-bf7e-4f856b3bfdec","order_by":7,"name":"Peigao Luo","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA10lEQVRIiWNgGAWjYBACPgYGxgOMDTYMbBA+M2EtIJVALWmkazkM4xOjRbr9woGfO87b87EffibBUGGd2MB+9gB+LTJnCg72nrmd2MaTZibBcCY9sYEnLwG/FomchAO8bbcT2CQYzCQY2w4nNkjwGBDUcvBv2zl7Ngn2bxKM/4jSkn7gMG/bAcY2CR6gLQ3EaJE5w3BY9kwy0C85xRYJx9KNgQz8Wvil2x8+fLvDzl6+/fjGGx9qrGX72c/g18KA4owEBgZYnOLTwv6AoJpRMApGwSgY4QAARuBCWkYA0hcAAAAASUVORK5CYII=","orcid":"","institution":"","correspondingAuthor":true,"prefix":"","firstName":"Peigao","middleName":"","lastName":"Luo","suffix":""}],"badges":[],"createdAt":"2026-04-03 05:08:54","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9308920/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9308920/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":107717582,"identity":"209252bc-05b6-4080-a480-45504b972070","added_by":"auto","created_at":"2026-04-24 10:13:01","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":303190,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution and collinearity of the \u003cem\u003eAktCCT\u003c/em\u003egenes on chromosomes in \u003cem\u003eA. trifoliata\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eNote\u003c/strong\u003e: The genes in red are distributed in the cluster, and the purple lines represent collinearity between the \u003cem\u003eAktCCTs.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/1bd1bc877b402aa62001b51e.png"},{"id":107869473,"identity":"694e92bc-5803-4feb-a75c-e7032eef0127","added_by":"auto","created_at":"2026-04-27 07:37:07","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":157889,"visible":true,"origin":"","legend":"\u003cp\u003eGene and protein structure analyses of the AktCCT family. A for phylogenetic analysis of the AktCCTs; B for the distribution of conserved motif in the putative AktCCTs\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/3df45c25e2f42246a77e5cdf.png"},{"id":107717552,"identity":"3d48fdef-2988-4b33-b57a-08ab6d70c13e","added_by":"auto","created_at":"2026-04-24 10:12:56","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":234905,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree consisting of the 46 \u003cem\u003eAktCCTs\u003c/em\u003e and 37 \u003cem\u003eAtCCTs\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/554ede8b93f1de4530cd3cb4.png"},{"id":107717557,"identity":"1a350ebe-2b40-47e6-860c-a5180aed73e5","added_by":"auto","created_at":"2026-04-24 10:12:57","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":267857,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of cis-acting elements within the upstream 2000 bp region of the \u003cem\u003eAktCCTs\u003c/em\u003ein \u003cem\u003eA. trifoliata\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eNote\u003c/strong\u003e: CEIAAR: cis-acting element involved in abscisic acid responsiveness; CEICCR: cis-acting element involved in cell cycle regulation; CEIDSR: cis-acting element involved in defense and stress responsiveness; CEIGR: cis-acting element involved in gibberellin responsiveness; CEILR: cis-acting element involved in light responsiveness; CEILTR: cis-acting element involved in low-temperature responsiveness; CEISAR: cis-acting element involved in salicylic acid responsiveness; CREEAI: cis-acting regulatory element essential for anaerobic induction; CREIAR: cis-acting regulatory element involved in auxin responsiveness; CREICC: cis-acting regulatory element involved in circadian control; CREIEE: cis-regulatory element involved in endosperm expression; CREILR: cis-acting regulatory element involved in light responsiveness; CRERS: cis-acting regulatory element, root specific; DPMC: element involved in the differentiation of palisade mesophyll cells; EFMEMA: element for maximal elicitor-mediated activation (2 copies); ELASI: enhancer-like element involved in anoxic-specific inducibility; GRE: gibberellin-responsive element; LRM: light-responsive element; MeJAAI: MeJAR essential for anaerobic induction; MeJAR: cis-acting regulatory element involved in MeJA-responsiveness; MYBBS(D): MYB binding site involved in drought-inducibility; MYBBS(FBGR): MYB binding site involved in flavonoid biosynthetic genes regulation; MYBBS(LR): MYB binding site involved in light responsiveness; PCMLR: part of a conserved DNA module involved in light responsiveness; PLRE: part of a light response element; PLRM: part of a light-responsive module; W-R: wound-responsive element; ZMR: cis-acting regulatory element involved in zein metabolism regulation.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/d0b6ddd01ca3b210e1625221.png"},{"id":107717550,"identity":"d211d62c-2609-46b2-a248-5cf4dbd50022","added_by":"auto","created_at":"2026-04-24 10:12:56","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":186652,"visible":true,"origin":"","legend":"\u003cp\u003eExpression level of 46 \u003cem\u003eAktCCTs\u003c/em\u003e, 2 \u003cem\u003eAktGIs\u003c/em\u003eand 7 \u003cem\u003eAktFTs\u003c/em\u003e in various tissues of \u003cem\u003eA. trifoliata\u003c/em\u003e at different developmental stages\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/57300920164cc27bd356de8f.png"},{"id":107717554,"identity":"3fcb0f1e-88ba-4ad2-998d-2aa872fb8d0f","added_by":"auto","created_at":"2026-04-24 10:12:56","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":441406,"visible":true,"origin":"","legend":"\u003cp\u003eCo-expression between 46 \u003cem\u003eAktCCTs\u003c/em\u003e and both 2 \u003cem\u003eAktGIs\u003c/em\u003e and 7 \u003cem\u003eAktFTs\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eNote: \u003c/strong\u003eRed represents a positive correlation, blue represents a negative correlation, and the darker the colour, the stronger the correlation. * for the significance at the level of p = 0. 05.\u003c/p\u003e","description":"","filename":"floatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/da613cd9b79e10e6e84e0d14.png"},{"id":108006265,"identity":"e4f6b7ab-9ce6-4630-822f-9f4c42d0b0c7","added_by":"auto","created_at":"2026-04-28 12:54:59","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2273739,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9308920/v1/33ce9e38-11a8-45ee-9bcc-7c67c106d551.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Genome-wide identification and expression pattern analysis of the CCT genes of Akebia trifoliata","fulltext":[{"header":"Introduction","content":"\u003cp\u003eFlowering is one of the most important biological behaviors in the plant kingdom, especially in angiosperms, which comprise approximately 300,000 extant species and dominate the vegetation of most terrestrial ecosystems (Crane et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). It not only has important ecological significance but also has substantial agricultural, horticultural and even ornamental value in practice (Wu et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Differences in flowering time have an impact on flower morphology in addition to ensuring that plants are evolutionarily adapted to their specific geographical distribution and accomplish their life cycle to reproduce (Shan et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, flowering initiation is usually affected by environmental signals such as temperature and sunlight duration and endogenous changes such as developmental status and hormone levels (Liu et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), which are closely related to physiological processes such as photoperiod and vernalization (Liu et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Therefore, detailing the characteristics of the genes involved in this process is important for understanding both the ecological distribution and flower development of a given plant species.\u003c/p\u003e \u003cp\u003ePlant photoperiod that has been discovered by more one hundred years (Garner and Allard \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1920\u003c/span\u003e), refers the behavior that flowering time is influenced by the comparative length of light and dark periods in the diurnal cycle (Sawa et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), and hence flowering plants are classified into three types according to their response to photoperiod: day-neutral, short-day and long-day (Lumsden et al. 2002). Various studies have suggested that \u003cem\u003eCONSTANS/CONSTANS-LIKE/TIMING OF CAB1\u003c/em\u003e (\u003cem\u003eCCT\u003c/em\u003e) gene families are the core elements affecting the plant photoperiod (Abe et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Takahashi. 2017; Liu et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Awal et al. 2022). The proteins encoded by \u003cem\u003eCCTs\u003c/em\u003e commonly contain a conserved domain consisting of 43\u0026ndash;45 amino acids at the C-terminus (Putterill et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and are commonly localized in the nucleus (Robson et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). In addition, variations widely exist among intraspecific and interspecific members (Chen et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), which provides important evidence for classifying into subfamilies.\u003c/p\u003e \u003cp\u003eAccording to studies in \u003cem\u003eArabidopsis thaliana\u003c/em\u003e, \u003cem\u003eOryza sativa\u003c/em\u003e, \u003cem\u003eSorghum bicolor\u003c/em\u003e, and \u003cem\u003eHordeum vulgare\u003c/em\u003e (Cockram et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), the CCT family can be structurally divided into three subfamilies: CONSTANS-like (COL), containing one or two B-box domains; Pseudo response regulators (PRR), with a response-regulator (REC) domain beside a CCT domain; and the CCT motif (CMF), with only a CCT domain (Khanna et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Cockram et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). In fact, a relatively special subclass called the GTCC subfamily containing the GATA domain also exists in some species such as \u003cem\u003eCajanus cajan\u003c/em\u003e and \u003cem\u003eBrassica rapa\u003c/em\u003e, which sometimes accompanies a zinc finger protein expressed in the inflorescence meristem (Tribhuvan et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These findings indicate the high genetic diversity in the structure of the \u003cem\u003eCCT\u003c/em\u003e family members among various species.\u003c/p\u003e \u003cp\u003eSince CONSTANS (CO), a circadian clock regulatory gene belonging to the \u003cem\u003eCOL\u003c/em\u003e subfamily, was first cloned (Wenkel et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), various studies have functionally demonstrated that the \u003cem\u003eCOL\u003c/em\u003e subfamily could acts as the central regulatory factor in the photoperiodic pathway, and in the known \"GI-CO-FT\" model \u003cem\u003eCO\u003c/em\u003e is first regulated by the upstream transcription factor named Gigantea (GI) and then it consequently regulates the downstream flowering locus T (FT) (Wu et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Andr\u0026eacute;s et al. 2012). At the same time, some studies have also shown that \u003cem\u003eCOL\u003c/em\u003e genes have important functions in nuclear localization, protein‒protein interactions and transcriptional regulation (Putterill et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Kaewphalug et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), suggesting that they may be DNA-binding transcription factors, but the \u003cem\u003eCCTs\u003c/em\u003e are not listed as a family in the Plant Transcription Factor Database (v5.0) (Jin et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), indicating that at least some CCTs lack the typical DNA-binding motif of classical transcription factors. In addition, some studies in the green algae Chlamydomonas and \u003cem\u003eA. thaliana\u003c/em\u003e have shown that the CCT domain, which is homologous to the CCAAT-binding domain, can interact with transcription factors such as NF-YA, NF-YB and NF-YC with the typical DNA-binding motif to form multimers (Gnesutta et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Lv et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), which further supports the view that COL indirectly regulates the expression of downstream genes, mainly through a complex pathway rather than directly binding the promoter region of the target genes. Ben-Naim et al (Ben-Naim et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) suggested that COL proteins usually lack a known DNA-binding motif and that their success in transcriptional regulation occurs mainly through the mediation of CCAAT binding factors, but some studies have shown that some COL proteins can directly bind DNA \u003cem\u003ein vitro\u003c/em\u003e studies (Tiwari et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Obviously, the details of the regulatory function and corresponding molecular mechanism of COL are not completely understood.\u003c/p\u003e \u003cp\u003eIn addition, the \u003cem\u003eCMF\u003c/em\u003e subfamily evolved from the \u003cem\u003eCOL\u003c/em\u003e subfamily, and thus, the functions of \u003cem\u003eCMF\u003c/em\u003e families are very similar to those of \u003cem\u003eCOL\u003c/em\u003e subfamilies (Cockram et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Compared with the \u003cem\u003eCOL\u003c/em\u003e and \u003cem\u003eCMF\u003c/em\u003e subfamilies, the PRR subfamily is functionally more conserved and comprises important components of the circadian clock, usually interacting with circadian clock associated 1 (CCA1) and late elongated hypocotyl (LHY) (Makino et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; David et al. 2002; Nakamichi et al. 2020). Moreover, \u003cem\u003ePRRs\u003c/em\u003e such as TOC1 can influence the expression of other transcription factors, such as \u003cem\u003eCO\u003c/em\u003e, \u003cem\u003ePIFs\u003c/em\u003e and \u003cem\u003eGI\u003c/em\u003e, by directly binding to their promoter region and even exerting self-regulatory effects (Hsu et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Gendron et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Therefore, more datasets from various species of different evolutionary clades would helpful to completely elucidate the complex regulatory network of the \u003cem\u003eCCT\u003c/em\u003e family.\u003c/p\u003e \u003cp\u003e \u003cem\u003eAkebia trifoliata\u003c/em\u003e (also called augmelon), which originates from the eastern edge of the Qinghai‒Tibet Plateau and is representative of the Lardizabalaceae family, is a climbing perennial woody vine that is narrowly distributed in several regions of East Asia, including subtropical China south of the Yellow River (Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Dong et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), so that it has largely been ignored by both scientists and entrepreneurs in the past. However, it is becoming a very popular topic in the field of edible wild plant domestication (Yu et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2024a\u003c/span\u003e) because of its multiple purposes, such as in traditional medicine (Yang et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e), for fresh fruit (Yang et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, Yi et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), as a woody oil crop (Guan et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Han et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) and even as an ornamental plant (Yang et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In addition, its value for theoretical study is also becoming increasingly appreciated because of its small genome size (Zhong et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e), short juvenile stage (Yang et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and important evolutionary position (Zhong et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2022b\u003c/span\u003e), and it has been widely employed in various studies, such as those involving flower development (Zhong et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e; Zhong et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), phenylalanine metabolism (Guan et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and flavonoid biosynthesis (Zhong et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2022d\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2024b\u003c/span\u003e). However, to date, there are no reports about its photoperiod characteristics.\u003c/p\u003e \u003cp\u003eIn fact, the most important reasons for using \u003cem\u003eA. trifoliata\u003c/em\u003e to investigate the close involvement of the \u003cem\u003eCCT\u003c/em\u003e family with the photoperiod are that we observed that it flowers in both spring and autumn each year in the \u003cem\u003eA. trifoliata\u003c/em\u003e Germplasm Nursery of Sichuan Agricultural University at Chongzhou Research Station (Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and it can even fruit in both March and September each year in the commercially cultivated garden (28\u0026deg; 19' N, 101\u0026deg; 07' E) of Jiulong County, China, which indicates that \u003cem\u003eA. trifoliata\u003c/em\u003e might be a day-neutral plant. In addition, abundant available omics data, such as genomic (Zhong et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2022a\u003c/span\u003e), transcriptomic (Chen et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and metabolic (Zhong et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2022d\u003c/span\u003e) data, provide a good opportunity to comprehensively characterize the \u003cem\u003eCCT\u003c/em\u003e family profile. Therefore, our objectives are to systemically identify the number and components of the \u003cem\u003eCCT\u003c/em\u003e family in the \u003cem\u003eA. trifoliata\u003c/em\u003e reference genome, to completely detail its structural and evolutionary characteristics and to finally determine their expression levels. These results would be helpful for further understanding the molecular mechanism of flowering in \u003cem\u003eA. trifoliata.\u003c/em\u003e\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eIdentification \u003cem\u003eCCT\u003c/em\u003e family in \u003cem\u003eA. trifoliata\u003c/em\u003e\u003c/p\u003e \u003cp\u003eGenome sequence annotation files of \u003cem\u003eA. trifoliata\u003c/em\u003e (accession ID: GWHBISH00000000) were downloaded from the National Genomics Data Centre (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://ngdc.cncb.ac.cn/\u003c/span\u003e\u003cspan address=\"https://ngdc.cncb.ac.cn/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e; accessed on 9 September 2024). The hidden Markov model of conserved \u003cem\u003eCCT\u003c/em\u003e sequences (PF06203) (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://pfam-legacy.xfam.org/\u003c/span\u003e\u003cspan address=\"http://pfam-legacy.xfam.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e; accessed on 9 November 2024) was employed to identify \u003cem\u003eAktCCTs\u003c/em\u003e. In addition, the sequences of all the proteins of \u003cem\u003eA. trifoliata\u003c/em\u003e were then scanned by HMMER 3.0 using the HMM with an E value of 1\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;5\u003c/sup\u003e, in which the amino acid sequences of \u003cem\u003eArabidopsis\u003c/em\u003e CCTs were downloaded from the TAIR website (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.Arabidopsis.org/\u003c/span\u003e\u003cspan address=\"https://www.Arabidopsis.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e; accessed 9 September 2024) and used to identify \u003cem\u003eAktCCTs\u003c/em\u003e. Only those \u003cem\u003eAktCCTs\u003c/em\u003e that could be identified by the two methods were treated as the final identified \u003cem\u003eAktCCTs\u003c/em\u003e. Moreover, the physicochemical properties of the predicted AktCCTs were determined using the ExPASy ProtParam website (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://web.expasy.org/protparam/\u003c/span\u003e\u003cspan address=\"https://web.expasy.org/protparam/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e; accessed on 9 November 2024).\u003c/p\u003e \u003cp\u003eConserved protein structure and chromosomal localization of \u003cem\u003eAktCCT\u003c/em\u003e\u003c/p\u003e \u003cp\u003eThe MEME website (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://meme-suite.org/meme/\u003c/span\u003e\u003cspan address=\"https://meme-suite.org/meme/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e; accessed 9 September 2024) was used with the default parameters to analyze the conserved motifs in the putative AktCCTs. Subsequently, TBtools (version 1.0876, CJ Chen, Guangzhou, China) was utilized to map the \u003cem\u003eAktCCTs\u003c/em\u003e on the chromosomes, adopting the gff file of \u003cem\u003eA. trifoliata\u003c/em\u003e. Additionally, TBtools was used to extract parameters such as gene length and gene location.\u003c/p\u003e \u003cp\u003eCalculation of Ka/Ks of homologous \u003cem\u003eAktCCT\u003c/em\u003e pairs\u003c/p\u003e \u003cp\u003eThe nonsynonymous substitution (Ka) to synonymous substitution (Ks) values (Ka/Ks) were calculated using TBtools-II software (v2.012, Chengjie Chen, China). Similarly, we performed intraspecific collinearity analysis using TBtools software to determine the duplication type.\u003c/p\u003e \u003cp\u003eCis-acting elements of the \u003cem\u003eAktCCT\u003c/em\u003e gene family\u003c/p\u003e \u003cp\u003eTo predict putative cis-acting elements in the promoter regions of the \u003cem\u003eAktCCTs\u003c/em\u003e, the 2,000 bp upstream sequences of the \u003cem\u003eAktCCT\u003c/em\u003e coding regions were analyzed using PlantCARE (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://bioinformatics.psb.ugent.be/webtools/plantcare/html/\u003c/span\u003e\u003cspan address=\"http://bioinformatics.psb.ugent.be/webtools/plantcare/html/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, accessed on 10 September 2024). After deleting unnecessary cis-acting elements, the distribution of the remaining cis-acting elements of the \u003cem\u003eAktCCTs\u003c/em\u003e was plotted using Basic Biosequence View in TBtools. In addition, the cis-acting elements were divided into hormone- and environment-responsive types according to their annotation.\u003c/p\u003e \u003cp\u003ePhylogenetic tree construction\u003c/p\u003e \u003cp\u003eThe phylogenetic tree of both all \u003cem\u003eAktCCTs\u003c/em\u003e and 37 reference \u003cem\u003eAtCCTs\u003c/em\u003e was constructed via the N‒J method with 1,000 bootstrap replicates using MEGA 11 software (v11.0.10, Auckland, New Zealand), in which 37 previously reported \u003cem\u003eAtCCTs\u003c/em\u003e were downloaded from the Plant Transcription Factor Database website (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://planttfdb.cbi.pku.edu.cn/\u003c/span\u003e\u003cspan address=\"http://planttfdb.cbi.pku.edu.cn/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, accessed on 24 September 2024) (Tamura et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In addition, the evolutionary relationships were constructed using Gene Structure View, and the online tool iTOL (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://itol.embl.de/\u003c/span\u003e\u003cspan address=\"https://itol.embl.de/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, accessed on 9 October 2024) was used for subsequent visualization.\u003c/p\u003e \u003cp\u003eSubcellular localization of the proteins putatively encoded by \u003cem\u003eAktCCTs\u003c/em\u003e\u003c/p\u003e \u003cp\u003eTo predict the putative subcellular localization of the candidate proteins, in silico analyses were performed using multiple computational tools. Amino acid sequences were submitted to the following online prediction servers: WoLF PSORT (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://wolfpsort.hgc.jp/\u003c/span\u003e\u003cspan address=\"https://wolfpsort.hgc.jp/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) for comprehensive prediction based on sorting signals and amino acid composition; Plant-mPLoc (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.csbio.sjtu.edu.cn/bioinf/plant-multi/\u003c/span\u003e\u003cspan address=\"http://www.csbio.sjtu.edu.cn/bioinf/plant-multi/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) for plant-specific multi-label localization; and TargetP 2.0 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.cbs.dtu.dk/services/TargetP/\u003c/span\u003e\u003cspan address=\"http://www.cbs.dtu.dk/services/TargetP/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) for the identification of N-terminal sequences and prediction of mitochondrial, chloroplast, secretory pathway, or other localizations.\u003c/p\u003e \u003cp\u003eDifferential expression of \u003cem\u003eCCT\u003c/em\u003e in \u003cem\u003eA. trifoliata\u003c/em\u003e\u003c/p\u003e \u003cp\u003eTranscriptome data for \u003cem\u003eA. trifoliata\u003c/em\u003e leaves, sepals and stamens during flowering period were downloaded from the NGDC database (login number: PRJCA019245). Additionally, HISAT2 software (v2.1.0, Mihaela Pertea, USA) and DESeq2 (v1.36.0, Michael I. Love, Germany) were used to extract the FPKM values from the transcriptomic data. We determined the expression of \u003cem\u003eAktCCTs\u003c/em\u003e in these 3 tissues using HeatMap. Further mapping of the correlations between \u003cem\u003eAktCCTs\u003c/em\u003e and both \u003cem\u003eAktGIs\u003c/em\u003e and \u003cem\u003eAktFTs\u003c/em\u003e was performed using OriginPro (2024 (64-bit) SR1 10.1.0.178).\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eThe significance of the differences in the characteristic parameters among the various groups was determined using SPSS (version 19.0; Armonk, NY: IBM Corp.), and correlation analysis of the expression levels was also performed using the same software.\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eChromosomal distribution and physiochemical characteristics of the \u003cem\u003eCCT\u003c/em\u003e family in \u003cem\u003eA. trifoliata\u003c/em\u003e\u003c/p\u003e \u003cp\u003eA total of 46 \u003cem\u003eCCT\u003c/em\u003e genes were identified in the \u003cem\u003eA. trifoliata\u003c/em\u003e reference genome, and they were orderly named \u003cem\u003eAktCCT1\u003c/em\u003e to \u003cem\u003eAktCCT46\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, \u003cb\u003eTable S1\u003c/b\u003e). Physically, all 46 \u003cem\u003eAktCCTs\u003c/em\u003e were unevenly distributed all 16 chromosomes and most of them were mapped to regions near chromosome ends (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), in which chromosome 3 had the most 7 \u003cem\u003eAktCCTs\u003c/em\u003e while chromosomes 9, 13, 15 and 16 each had only one. According to the definition of gene clusters, the genes were assigned to 41 loci on the chromosomes, including 36 singletons and 5 gene clusters within 250-kb chromosomal regions, in which each cluster consisted of two \u003cem\u003eAktCCTs\u003c/em\u003e (\u003cem\u003eAktCCT3\u003c/em\u003e/\u003cem\u003eAktCCT4\u003c/em\u003e, \u003cem\u003eAktCCT13\u003c/em\u003e/\u003cem\u003eAktCCT14\u003c/em\u003e, \u003cem\u003eAktCCT15\u003c/em\u003e/\u003cem\u003eAktCCT16\u003c/em\u003e, \u003cem\u003eAktCCT19\u003c/em\u003e/\u003cem\u003eAktCCT20\u003c/em\u003e, and \u003cem\u003eAktCCT26\u003c/em\u003e/\u003cem\u003eAktCCT27\u003c/em\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNote\u003c/strong\u003e \u003cp\u003eThe genes in red are distributed in the cluster, and the purple lines represent collinearity between the \u003cem\u003eAktCCTs.\u003c/em\u003e\u003c/p\u003e \u003c/p\u003e \u003cp\u003eMain evolutionary experience of the \u003cem\u003eAktCCTs\u003c/em\u003e\u003c/p\u003e \u003cp\u003eWe found that 33 (71.7%) \u003cem\u003eAktCCTs\u003c/em\u003e were produced by whole-genome duplication (WGD) or segmental duplication while and the remaining 13 (28.3%) \u003cem\u003eAktCCTs\u003c/em\u003e including all \u003cem\u003eAktCCTs\u003c/em\u003e on chromosomes 9, 15 and 16 were produced by dispersal duplication (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u003cb\u003e)\u003c/b\u003e. Interestingly, all 10 \u003cem\u003eAktCCTs\u003c/em\u003e in cluster were produced by WGD or segmental duplication.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e\u003cem\u003eCCTs\u003c/em\u003e identified in \u003cem\u003eA. trifoliata.\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"11\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eName\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e \u003cp\u003eGenes family\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c10\" namest=\"c7\"\u003e \u003cp\u003eCharacteristics of putative proteins\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eDT\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCP\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBegin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEnd\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGL(bp)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eNE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eNAA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eMW (kDa)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePI\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSL\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT1\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28523241\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e28526070\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2829\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e436\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e49.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT2\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e31772454\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e31779638\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7184\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e408\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e44.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT3\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5902189\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5912295\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10106\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e542\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e40.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT4\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e 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align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT5\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1015390\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1019400\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e340\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e38.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT6\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2575299\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2579508\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4209\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e292\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e33.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e7.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT7\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e11202320\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11204761\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2441\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e382\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e43.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT8\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e25181097\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e25189683\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8586\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e481\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e54.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT9\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e44086815\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e44102632\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e15817\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e422\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e45.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT10\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e51615841\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e51624543\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8702\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e712\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e78.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT11\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e53009727\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e53014571\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4844\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e282\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT12\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e6504460\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6511344\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6884\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e400\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e44.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT13\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10441408\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10453360\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e11952\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e347\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e38.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT14\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10456733\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10466278\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e9545\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e254\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e27.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT15\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e40953416\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e40973777\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e20361\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e341\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e37.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT16\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e41035410\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e41059349\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e23939\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e310\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e33.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT17\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e955460\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e960517\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5057\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e54.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT18\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e9772961\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9774777\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1816\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e329\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e37.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT19\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5801167\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e 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\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5976398\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5978442\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2044\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e268\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e30.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT21\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e32202888\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e32210121\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7233\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e558\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e63.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT22\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e33382781\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e33385175\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2394\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e362\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e40.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT23\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2406849\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2408961\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2112\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e226\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e26.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e7.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT24\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e24582546\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e24584147\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1601\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e217\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e25.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT25\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e49018032\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e49020644\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2612\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e381\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e42.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT26\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e49876689\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e49879917\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3228\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e403\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e45.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e 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\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e7.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT28\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e18517830\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e18520944\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3114\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e414\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e45.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT29\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4329166\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4330869\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1703\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e359\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e39.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT30\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e6725436\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6742598\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e17162\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e270\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e29.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT31\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7217500\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7220765\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3265\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e389\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e44.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT32\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29942599\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e29944927\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2328\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e320\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e36.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT33\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e30988975\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30995773\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6798\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e289\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT34\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8023901\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e8027094\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3193\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e259\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e29.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT35\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e11833091\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11837383\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4292\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e282\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e6.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT36\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e23047610\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e23048741\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1131\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e224\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e26.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e4.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT37\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e30582722\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30584224\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1502\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e156\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e17.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e9.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT38\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e34672988\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e34674880\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1892\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e322\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e36.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT39\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e35707409\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e35711202\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3793\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e274\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e30.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT40\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e36585577\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e36592109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e6532\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e727\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e81.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT41\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e36873128\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e36878636\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5508\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e703\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e78.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT42\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e351777\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e358812\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7035\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e414\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e45.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT43\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2520450\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2525582\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5132\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e722\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e80.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT44\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4925216\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e4932511\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e7295\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e725\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e80.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eW/S\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT45\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7240017\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e7240459\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e442\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e7.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e11.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAktCCT46\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e29040983\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e29045522\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4539\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e404\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e45.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e5.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cb\u003eNote\u003c/b\u003e: CP: chromosomal position; GL: gene length; NE: number of exons; DT: duplication type; NAA: number of amino acids; MW: molecular weight; PI: isoelectric point; SL: subcellular localization; N: nuclear; C: chloroplast; D: dispersed; W/S: WGD/segmental.\u003c/p\u003e \u003cp\u003eIn addition, 27 orthologous pairs related with 33 \u003cem\u003eAktCCTs\u003c/em\u003e were detected, among which 8 \u003cem\u003eAktCCTs\u003c/em\u003e (\u003cem\u003eAktCCT3\u003c/em\u003e, \u003cem\u003eAktCCT7\u003c/em\u003e, \u003cem\u003eAktCCT11\u003c/em\u003e, \u003cem\u003eAktCCT14\u003c/em\u003e, \u003cem\u003eAktCCT16\u003c/em\u003e, \u003cem\u003eAktCCT29\u003c/em\u003e, \u003cem\u003eAktCCT34\u003c/em\u003e and \u003cem\u003eAktCCT41\u003c/em\u003e) had 3 orthologous gene pairs while 20 \u003cem\u003eAktCCTs\u003c/em\u003e (\u003cem\u003eAktCCT2\u003c/em\u003e, \u003cem\u003eAktCCT 8\u003c/em\u003e, \u003cem\u003eAktCCT10\u003c/em\u003e, \u003cem\u003eAktCCT12\u003c/em\u003e, \u003cem\u003eAktCCT15\u003c/em\u003e, \u003cem\u003eAktCCT17\u003c/em\u003e, \u003cem\u003eAktCCT18\u003c/em\u003e, \u003cem\u003eAktCCT19\u003c/em\u003e, \u003cem\u003eAktCCT20\u003c/em\u003e, \u003cem\u003eAktCCT23\u003c/em\u003e, \u003cem\u003eAktCCT25\u003c/em\u003e, \u003cem\u003eAktCCT26\u003c/em\u003e, \u003cem\u003eAktCCT27\u003c/em\u003e, \u003cem\u003eAktCCT30\u003c/em\u003e, \u003cem\u003eAktCCT31\u003c/em\u003e, \u003cem\u003eAktCCT32\u003c/em\u003e, \u003cem\u003eAktCCT33\u003c/em\u003e, \u003cem\u003eAktCCT39\u003c/em\u003e, \u003cem\u003eAktCCT42\u003c/em\u003e and \u003cem\u003eAktCCT44\u003c/em\u003e) only had one orthologous gene pair. The Ka/Ks values varied from 0.15 (\u003cem\u003eAktCCT30\u003c/em\u003e and \u003cem\u003eAktCCT25\u003c/em\u003e) to 0.57 (\u003cem\u003eAktCCT40\u003c/em\u003e and \u003cem\u003eAktCCT38\u003c/em\u003e), and the average mean Ka/Ks was only 0.33 (\u003cb\u003eTable S2\u003c/b\u003e).\u003c/p\u003e \u003cp\u003eConserved structure, class and phylogenetic tree of the \u003cem\u003eAktCCTs\u003c/em\u003e\u003c/p\u003e \u003cp\u003eWe identified a total of 10 conserved motifs (from 1 to 10) with a varying width from 11 amino acids in Motif 6 to 50 amino acids in five motifs (Motif 2, 4, 5, 7 and 9) among 46 putative AktCCTs (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), and we also identified five conserved domains (CCT, B-box, GATA, TIFY and REC) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). By sequence comparing, we found that CCT, B-box and REC domain distributed across Motif 1 and Motif 6, Motif 2 and Motif 8, and Motif 5 and Motif 7, respectively, while GATA and TIFY only distributed within Motif 3 and Motif 4, respectively. On one hand, two AktCCTs (AktCCT6 and AktCCT26) had only Motif 1 while 31 AktCCTs had the most 4 motifs (\u003cb\u003eTable S3\u003c/b\u003e). On the other hand, Motif 1 distributed all 46 AktCCTs while both Motif 9 and Motif 10 only existed in five AktCCTs. Based on the existence of sequence encoding conserved domain, we classified 46 \u003cem\u003eAktCCTs\u003c/em\u003e into four classes: 18 \u003cem\u003eCMFs\u003c/em\u003e, 12 \u003cem\u003eCOLs\u003c/em\u003e, 9 \u003cem\u003eGTCCs\u003c/em\u003e, and 7 \u003cem\u003ePRRs\u003c/em\u003e. Many characters such as gene whole length and the exon number, and amino acids number and molecular weight of putative protein existed significant difference among various classes at p\u0026thinsp;=\u0026thinsp;0.05 level. For example, the \u003cem\u003eCOL\u003c/em\u003e class had the lowest average gene length, number of exons while \u003cem\u003eGTCC\u003c/em\u003e class had the lowest number of amino acids and molecular weight of putative protein. On the contrary, \u003cem\u003ePRR\u003c/em\u003e had the greatest average gene length, exon number, amino acid number and molecular weights (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn addition, the phylogenetic tree of the 46 \u003cem\u003eAktCCTs\u003c/em\u003e was constructed according to the sequence component, in which 37 reference \u003cem\u003eAtCCTs\u003c/em\u003e were also employed as the control. According to the phylogenetic tree, 46 \u003cem\u003eAktCCTs\u003c/em\u003e were assigned into four distinct branches: Ⅰ, Ⅱ, Ⅲ and Ⅳwith 8, 9, 18 and 11 \u003cem\u003eAktCCTs\u003c/em\u003e, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), and the clustering results of 37 \u003cem\u003eAtCCTs\u003c/em\u003e were well agreed by the previously reported classification (Huang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Interestingly, all 7 \u003cem\u003ePRRs\u003c/em\u003e and 9 \u003cem\u003eGTCCs\u003c/em\u003e were completely assigned intoⅠandⅡbranch, respectively, and similarly, all 18 \u003cem\u003eCMFs\u003c/em\u003e except of \u003cem\u003eAktCCT45\u003c/em\u003e and 12 \u003cem\u003eCOLs\u003c/em\u003e except of \u003cem\u003eAktCCT28\u003c/em\u003e were assigned into Ⅲ and Ⅳ, respectively, which indicated that some DNA sequence beside that encoded conserved domain would be also experienced a fast evolution.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSequence annotation of motifs of 46 putative AktCCTs\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDomain\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSequences\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eWidth\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCCT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eREAR\u003c/b\u003eLL\u003cb\u003eRYREKRK\u003c/b\u003eERN\u003cb\u003eFEK\u003c/b\u003eK\u003cb\u003eIRY\u003c/b\u003eA\u003cb\u003eSRK\u003c/b\u003eALAD\u003cb\u003eRRP\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eB-box\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eC\u003c/b\u003eEF\u003cb\u003eC\u003c/b\u003eEEAPAVIY\u003cb\u003eC\u003c/b\u003eKADAAAL\u003cb\u003eC\u003c/b\u003eLT\u003cb\u003eC\u003c/b\u003eDRNI\u003cb\u003eH\u003c/b\u003eSANPLSRR\u003cb\u003eH\u003c/b\u003eERTPJCPICGSQ\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGATA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCQHCGISEKSTPMMRRGPAGPRT\u003cb\u003eLCNACGL\u003c/b\u003eMWANKGT\u003cb\u003eL\u003c/b\u003eRD\u003cb\u003eL\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e41\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTIFY\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGTDZ\u003cb\u003eLT\u003c/b\u003eLS\u003cb\u003eF\u003c/b\u003eZGEV\u003cb\u003eY\u003c/b\u003eVFDAV\u003cb\u003eSPEKVQA\u003c/b\u003eVL\u003cb\u003eLLLG\u003c/b\u003eGYEIPSAVPTIDIPSHNN\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eREC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHEICKNI\u003cb\u003ePVIMM\u003c/b\u003eSSHDSVGVVFKCMLK\u003cb\u003eGAADFLVKP\u003c/b\u003eVRKNELRNLWQHVW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCCT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eRV\u003c/b\u003eKG\u003cb\u003eR\u003c/b\u003eF\u003cb\u003eVK\u003c/b\u003eA\u003cb\u003eKE\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eREC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eLVESD\u003c/b\u003eDS\u003cb\u003eTRQIVSALLRK\u003c/b\u003eCGYEVTAVADGLKAWEILKGRSH\u003cb\u003eNIDLV\u003c/b\u003eLTEV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eB-box\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePASVY\u003cb\u003eC\u003c/b\u003eRADSAYL\u003cb\u003eC\u003c/b\u003eQB\u003cb\u003eC\u003c/b\u003eDWNI\u003cb\u003eH\u003c/b\u003eGANTLA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eIQKSISSGCLSSMDWIHGGAMRPNFLDFQGVDFEAAYGMRRAFSEGDIQT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMotif 10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGDLFKAPEPIIEEPTIGLDPMSAAISMISCGEDVISPQTMKV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cb\u003eNote\u003c/b\u003e: the part in bold for conserved domain.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eMultiple comparative analyses of the protein physicochemical properties of the \u003cem\u003eAktCCTs\u003c/em\u003e in different subgroups\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"20\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c16\" colnum=\"16\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c17\" colnum=\"17\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c18\" colnum=\"18\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c19\" colnum=\"19\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c20\" colnum=\"20\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eGroup\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e \u003cp\u003eGL\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e \u003cp\u003eNE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c12\" namest=\"c10\"\u003e \u003cp\u003eNAA\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c16\" namest=\"c14\"\u003e \u003cp\u003eMW (kDa)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c17\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c20\" namest=\"c18\"\u003e \u003cp\u003ePI\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003emax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003emin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003emean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003emax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003emin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003emean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003emax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003emin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003emean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c14\"\u003e \u003cp\u003emax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c15\"\u003e \u003cp\u003emin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c16\"\u003e \u003cp\u003emean\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c18\"\u003e \u003cp\u003emax\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c19\"\u003e \u003cp\u003emin\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c20\"\u003e \u003cp\u003emean\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCOL\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8586\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1703\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6065.5\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.28\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e320\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e388.4\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e54.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e36.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e43.0\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e8.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e6.28\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eGTCC\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e23939\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2044\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6498.4\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.54\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e542\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e296\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e368.5\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e44.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e31.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e40.4\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e9.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e4.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e6.31\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eCMF\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e6884\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e442\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6227.2\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.39\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e727\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e393.9\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e7.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e43.6\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e11.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e4.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e6.22\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003ePRR\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e10021\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5132\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6626.3\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.60\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e767\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e274\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e405.1\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c13\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c14\"\u003e \u003cp\u003e83.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e30.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e45.3\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e8.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e4.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e6.24\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"20\"\u003e\u003cb\u003eNote\u003c/b\u003e: GL, NE, NAA, MW and PI represent the gene length, number of exons, number of amino acids, molecular weight and isoelectric point, respectively. a, b and c indicate significance at the p\u0026thinsp;=\u0026thinsp;0.05 level.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eCis-acting components of the \u003cem\u003eAktCCTs\u003c/em\u003e\u003c/p\u003e \u003cp\u003eBy examining the 2,000 bp upstream regions of the \u003cem\u003eAktCCTs\u003c/em\u003e, we identified a total of 1,220 functional cis-acting elements among all the \u003cem\u003eAktCCTs\u003c/em\u003e, among which 530 and 690 cis-acting elements belong to 13 hormone- and 15 environment-responsive types, respectively (\u003cb\u003eTable S4\u003c/b\u003e). Further analysis revealed that among the 13 kinds of hormone-responsive cis-acting elements, the number of cis-acting regulatory elements involved in MeJA responsiveness (MeJAR) and the number of cis-acting regulatory elements involved in root specificity (CRERS) were the greatest (193) and the lowest (1), respectively, and the sole CRERS was in the upstream region of \u003cem\u003eAktCCT17\u003c/em\u003e. Similarly, among the 15 kinds of environment-responsive cis-acting elements, the number of light-responsive elements (LRMs) and the number of MYB binding sites involved in drought inducibility (MYBBS(D)) and the number of wound-responsive elements (W-Rs) were the greatest (257) and least (2), respectively. In addition, the number of cis-acting elements in each \u003cem\u003eAktCCT\u003c/em\u003e exhibited large variation. For example, there were up to 59 (34 hormone-responsive types and 25 environment-responsive types) cis-acting elements in \u003cem\u003eAktCCT8\u003c/em\u003e, whereas there were only 9 (4 hormone-responsive types and 5 environment-responsive types) cis-acting elements in \u003cem\u003eAktCCT1.\u003c/em\u003e\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNote\u003c/strong\u003e \u003cp\u003eCEIAAR: cis-acting element involved in abscisic acid responsiveness; CEICCR: cis-acting element involved in cell cycle regulation; CEIDSR: cis-acting element involved in defense and stress responsiveness; CEIGR: cis-acting element involved in gibberellin responsiveness; CEILR: cis-acting element involved in light responsiveness; CEILTR: cis-acting element involved in low-temperature responsiveness; CEISAR: cis-acting element involved in salicylic acid responsiveness; CREEAI: cis-acting regulatory element essential for anaerobic induction; CREIAR: cis-acting regulatory element involved in auxin responsiveness; CREICC: cis-acting regulatory element involved in circadian control; CREIEE: cis-regulatory element involved in endosperm expression; CREILR: cis-acting regulatory element involved in light responsiveness; CRERS: cis-acting regulatory element, root specific; DPMC: element involved in the differentiation of palisade mesophyll cells; EFMEMA: element for maximal elicitor-mediated activation (2 copies); ELASI: enhancer-like element involved in anoxic-specific inducibility; GRE: gibberellin-responsive element; LRM: light-responsive element; MeJAAI: MeJAR essential for anaerobic induction; MeJAR: cis-acting regulatory element involved in MeJA-responsiveness; MYBBS(D): MYB binding site involved in drought-inducibility; MYBBS(FBGR): MYB binding site involved in flavonoid biosynthetic genes regulation; MYBBS(LR): MYB binding site involved in light responsiveness; PCMLR: part of a conserved DNA module involved in light responsiveness; PLRE: part of a light response element; PLRM: part of a light-responsive module; W-R: wound-responsive element; ZMR: cis-acting regulatory element involved in zein metabolism regulation.\u003c/p\u003e \u003c/p\u003e \u003cp\u003eExpression pattern analysis of the \u003cem\u003eCCT\u003c/em\u003e family genes of \u003cem\u003eA. trifoliata\u003c/em\u003e\u003c/p\u003e \u003cp\u003eThe transcript levels of all 46 \u003cem\u003eAkCCTs\u003c/em\u003e were analyzed in various tissues (leaves, sepals and stamens) at different developmental stages during flowering. The results showed that although the majority of \u003cem\u003eAktCCTs\u003c/em\u003e had relatively low expression, but 19 \u003cem\u003eAktCCTs\u003c/em\u003e had a detectable expression level in leaf. \u003cem\u003eAkCCT27\u003c/em\u003e exhibited high expression across all tissues during whole period (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e\u003cb\u003e)\u003c/b\u003e. In addition, \u003cem\u003eAkCCT26\u003c/em\u003e was highly expressed at sepals and stamens. Previous studies showed that \u003cem\u003eFT\u003c/em\u003e and \u003cem\u003eGI\u003c/em\u003e were involved in the CO related pathway (Mizoguchi, 2005), and they should also be used to determine the expression profile of the \u003cem\u003eAktCCTs\u003c/em\u003e in the known \"GI-CO-FT\" model, so that ee further identified two \u003cem\u003eAktGIs\u003c/em\u003e (\u003cem\u003eAktGI1\u003c/em\u003e and \u003cem\u003eAktGI2\u003c/em\u003e) and seven \u003cem\u003eAktFTs\u003c/em\u003e (from \u003cem\u003eAktFT1\u003c/em\u003e to \u003cem\u003eAktFT7\u003c/em\u003e) (\u003cb\u003eTable S5\u003c/b\u003e). In addition, we found that \u003cem\u003eAktFT2\u003c/em\u003e and \u003cem\u003eAktFT4\u003c/em\u003e was highly expressed in leaf and that the expression of both \u003cem\u003eAktGIs\u003c/em\u003e was very low in all tissues at all time points, but the pression of \u003cem\u003eAktGI1\u003c/em\u003e was detectable \u003cb\u003e(\u003c/b\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Interestingly, \u003cem\u003eAktFT2, AktFT4\u003c/em\u003e and \u003cem\u003eAktFT5\u003c/em\u003e as well as \u003cem\u003eAktGI1\u003c/em\u003e had significantly co-expressed relationship with many \u003cem\u003eAktCCTs\u003c/em\u003e such as \u003cem\u003eAktCCT17\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNote\u003c/strong\u003e \u003cp\u003eRed represents a positive correlation, blue represents a negative correlation, and the darker the colour, the stronger the correlation. * for the significance at the level of p\u0026thinsp;=\u0026thinsp;0. 05.\u003c/p\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eTo adapt to seasonal and environmental changes, plants have specifically evolved a regulatory mechanism that optimizes developmental events via the light response pathway (Takagi et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), in which CCT family members play very important roles in various processes, such as flowering, circadian rhythms and even the abiotic stress response (Liu et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This suggests that the structure and function of CCTs may have experienced rapid variation and divergence during evolution. In the present study, we found that among the 46 \u003cem\u003eAktCCTs\u003c/em\u003e, many physiochemical parameters, such as gene length, number of exons, number of amino acids, and the molecular weight and isoelectric point of the putative proteins, differed substantially (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). In fact, structural diversity may be favored by various evolutionary experiences. Previous studies have demonstrated that WGD is an important evolutionary force in almost all organisms, especially plants, and moreover, WGD is usually related to different clades (Jiao et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). For example, a recent report suggested that θ WGD and ω WGD favored the survival of eudicots and the thriving of core eudicots around the Jurassic-Cretaceous boundary and the Middle Cretaceous, respectively (Zhong et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022c\u003c/span\u003e). Here, 33 (71.7%), including all 10 \u003cem\u003eAktCCTs\u003c/em\u003e found in gene clusters out of the 46 \u003cem\u003eAktCCTs\u003c/em\u003e, were produced by WGD or segmental duplication (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), which could have presented an important opportunity for numerous variations in \u003cem\u003eAktCCTs\u003c/em\u003e to evolve. In addition, the small Ka/Ks value suggested that \u003cem\u003eAktCCTs\u003c/em\u003e might have undergone purifying selection during evolution and maintained a conserved function (\u003cb\u003eTable S2\u003c/b\u003e), which helps \u003cem\u003eA. trifoliata\u003c/em\u003e quickly eliminate damaged mutants, especially individuals whose light-responsive pathways are disrupted (Wang et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Hsu et al. 2014).\u003c/p\u003e \u003cp\u003eThe chloroplast is the most important organ related to the light-responsive process (Schwenkert et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), and interestingly, all 4 \u003cem\u003eAktCCTs\u003c/em\u003e (\u003cem\u003eAktCCT22\u003c/em\u003e, \u003cem\u003eAktCCT25\u003c/em\u003e, \u003cem\u003eAktCCT28\u003c/em\u003e and \u003cem\u003eAktCCT29\u003c/em\u003e), encoding proteins localized in the chloroplast, only had two exons and they belonged to the \u003cem\u003eCOL\u003c/em\u003e type. (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Moreover, their promoters contained both MeJAR hormone-responsive and LRM environment-responsive cis-acting elements (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e; \u003cb\u003eTable S5\u003c/b\u003e). On the other hand, their compartmental distribution and cis-acting element component indicated that they may retain the basic function of responding to changes in light, further suggesting that the \u003cem\u003eAktCCTs\u003c/em\u003e might be older. In addition, on the basis of the knowledge of \u003cem\u003eCCTs\u003c/em\u003e, the \u003cem\u003eCOL\u003c/em\u003e subfamily is the oldest, whereas other subfamilies, especially \u003cem\u003eCMF\u003c/em\u003e, are usually the evolved offspring of \u003cem\u003eCOLs\u003c/em\u003e. Therefore, how \u003cem\u003eCOLs\u003c/em\u003e produced other subfamilies is one of our concerns.\u003c/p\u003e \u003cp\u003eIn total, the most distinct difference between the \u003cem\u003eCOLs\u003c/em\u003e and other types was the existence of a B-box (Cockram et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Tribhuvan et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In this study, we found that among the various subfamilies, the gene length, number of exons, number of amino acids and molecular weight of the \u003cem\u003ePRR\u003c/em\u003e members were the largest. In contrast, the number of amino acids and molecular weight of \u003cem\u003eGTCC\u003c/em\u003e were the smallest while the gene length and exon number of \u003cem\u003eCOL\u003c/em\u003e was the smallest (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Based on this information, we propose that the \u003cem\u003eCMFs\u003c/em\u003e could have been produced from \u003cem\u003eCOLs\u003c/em\u003e mainly by losing the DNA regions putatively encoding the B-box conserved domain, whereas the \u003cem\u003ePRRs\u003c/em\u003e and \u003cem\u003eGTCCs\u003c/em\u003e might have been produced mainly via the insertion of new DNA sequences into the encoding and non-encoding regions of the corresponding B-box sequence, respectively.\u003c/p\u003e \u003cp\u003eVarious studies have suggested that \u003cem\u003eCO\u003c/em\u003e or \u003cem\u003eCOL\u003c/em\u003e can directly regulate the key downstream flowering gene \u003cem\u003eFT\u003c/em\u003e by altering its status and that this process is regulated by the upstream gene GI; thus, a known \"GI-CO-FT\" regulatory model was proposed (Fowler et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; David et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Sawa et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Imaizumi et al. 2010). From the perspective of the system phylogenetic tree, most \u003cem\u003eAktCCTs\u003c/em\u003e such as \u003cem\u003eAktCCT17\u003c/em\u003e were distributed into the Ⅳ branch that contained the key \u003cem\u003eArabidopsis\u003c/em\u003e flowering regulatory gene \u003cem\u003eCO\u003c/em\u003e (\u003cem\u003eAt5g15840\u003c/em\u003e) (Figs.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Otherwise, two \u003cem\u003eAktCCTs\u003c/em\u003e (\u003cem\u003eAktCCT17\u003c/em\u003e and \u003cem\u003eAktCCT27\u003c/em\u003e) obviously upregulate in leaves (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). Both \u003cem\u003eAktCCT17\u003c/em\u003e and \u003cem\u003eAktCCT27\u003c/em\u003e exhibited a significant co-expression with \u003cem\u003eAktFT2, AktFT4, AktFT5\u003c/em\u003e and \u003cem\u003eAktGI1\u003c/em\u003e involved in \"GI-CO-FT\" model, but the unique cis-acting elements binding with the sole root-related CRERS transcription factor existed in \u003cem\u003eAktCCT17\u003c/em\u003e (\u003cb\u003eTable S4\u003c/b\u003e). Moreover, previous studies showed that flowering is highly related with root behavior (Zou et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), so that \u003cem\u003eAktCCT17\u003c/em\u003e could play a key role in flowering of \u003cem\u003eA. trifoliata\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eOverall, genome duplication events, especially WGD, afforded an important opportunity for the diversification of the \u003cem\u003eAktCCTs\u003c/em\u003e, and at the same time, strong purifying selection facilitated their evolution toward favorable environmental adaptation. In addition, the sequences encoding conserved domains and the other sequences in the \u003cem\u003eAktCCTs\u003c/em\u003e may have evolved in parallel to further enrich the structural and functional diversity. Finally, \u003cem\u003eAktCCT17\u003c/em\u003e may play a very important role in regulating flowering.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of Interest\u003c/h2\u003e \u003cp\u003eThe authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as potential conflicts of interest.\u003c/p\u003e \u003c/p\u003e\u003cp\u003e \u003ch2\u003eSupplementary Material\u003c/h2\u003e \u003cp\u003eThe Supplementary Material for this article can be found online.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work was supported by grants from the Natural Science Foundation of Sichuan Province, grant number 2024NSFSC1310; Regional Innovation Cooperation of Sichuan Province, grant number 2024YFHZ0109; and Central guidance for local regional system construction, grant number 2024ZYD0284; Central Guidance for Local Areas - Construction of the Regional System 2024ZYD0284.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eDL: formal analysis, Software, Writing - Original Draft - Visualization. XY, HY, YX, CC, FT, YH and JL: Obtained the data - Participated in the data analysis, Discussion. XL: Proofreading the language of the manuscript. PL: Conceptualization, Writing - Review and Editing, Project - Administration, Funding Acquisition. All the authors contributed to the discussion of the results, reviewed the manuscript and approved the final article. Details of all funding sources should be provided, including grant numbers if applicable. Please ensure to add all necessary funding information, as after publication this is no longer possible.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets presented in this study can be found in online repositories. The names of the repositories/repositories and accession number(s) can be found in the article/Supplementary Material.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbe J, Xu D, Miyano A, Komatsu K, Kanazawa A, Shimamoto Y (2003) Photoperiod-insensitive Japanese soybean landraces differ at two maturity loci. Crop Sci 43:1300\u0026ndash;1304. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/jhered/esm114\u003c/span\u003e\u003cspan address=\"10.1093/jhered/esm114\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAlabad\u0026iacute; D, Yanovsky MJ, M\u0026aacute;s P, Harmer SL, Kay SA (2002) Critical role for CCA1 and LHY in maintaining circadian rhythmicity in \u003cem\u003eArabidopsis\u003c/em\u003e. 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Hereditas (Beijing) 42:739\u0026ndash;751. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.16288/j.yczz.20-014\u003c/span\u003e\u003cspan address=\"10.16288/j.yczz.20-014\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"genetic-resources-and-crop-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"gres","sideBox":"Learn more about [Genetic Resources and Crop Evolution](https://www.springer.com/journal/10722)","snPcode":"10722","submissionUrl":"https://submission.nature.com/new-submission/10722/3","title":"Genetic Resources and Crop Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Akebia trifoliata, flowering, photoperiod, CONSTANS/CONSTANS-LIKE/TIMING OF CAB1 (CCT), GI-CO-FT model","lastPublishedDoi":"10.21203/rs.3.rs-9308920/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9308920/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eFlowering is one of the important developmental events for many plants, and the CONSTANS/CONSTANS-LIKE/TIMING OF CAB1 (CCT) genes play key roles in this process. We identified a total of 46 \u003cem\u003eAktCCTs\u003c/em\u003e, which are widely distributed across all chromosomes of the \u003cem\u003eAkebia trifoliata\u003c/em\u003e genome. Their promoter regions contain hormone- and environment-responsive cis-acting elements. The average length and number of exons were 1,787.2 bp and 5.2, respectively, and the average number of amino acids, molecular weight and isoelectric point of putative protein were 388.4, 43.0 kDa and 6.3, respectively. Evolutionarily, \u003cem\u003eAktCCTs\u003c/em\u003e experienced whole or segmental genome duplication and purifying selection and were divided into four classes (\u003cem\u003eCOL\u003c/em\u003e, \u003cem\u003ePRR\u003c/em\u003e, \u003cem\u003eCMF\u003c/em\u003e and \u003cem\u003eGTCC\u003c/em\u003e) on the basis of sequence similarity and conserved domains, among which the \u003cem\u003eCMF\u003c/em\u003e had the lowest gene length, number of exons, number of amino acids and molecular weights. Functionally, some \u003cem\u003eAktCCTs\u003c/em\u003e expressed a detectable level. Otherwise, the expression of the \u003cem\u003eCOL AktCCT17\u003c/em\u003e significantly positively related with that of \u003cem\u003eAktFT2, AktFT4, AktFT5\u003c/em\u003e and \u003cem\u003eAktGI1\u003c/em\u003e, so that \u003cem\u003eAktCCT17\u003c/em\u003e may be the crucial gene regulating the flowering behavior of \u003cem\u003eA. trifoliata\u003c/em\u003e by \"GI-CO-FT\" model. Overall, these results provide valuable insights into the potential roles of these genes in regulating the biological process of flowering.\u003c/p\u003e","manuscriptTitle":"Genome-wide identification and expression pattern analysis of the CCT genes of Akebia trifoliata","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-24 10:12:28","doi":"10.21203/rs.3.rs-9308920/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"40278129410724395113485488617938862873","date":"2026-05-16T03:13:29+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"217934478511274887845882710077480812645","date":"2026-05-07T12:38:02+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-16T18:51:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-07T12:05:24+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-07T12:04:54+00:00","index":"","fulltext":""},{"type":"submitted","content":"Genetic Resources and Crop Evolution","date":"2026-04-03T04:50:46+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"genetic-resources-and-crop-evolution","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"gres","sideBox":"Learn more about [Genetic Resources and Crop Evolution](https://www.springer.com/journal/10722)","snPcode":"10722","submissionUrl":"https://submission.nature.com/new-submission/10722/3","title":"Genetic Resources and Crop Evolution","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6c2ba7bf-084d-4a91-98a9-f01c32244d90","owner":[],"postedDate":"April 24th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewerAgreed","content":"40278129410724395113485488617938862873","date":"2026-05-16T03:13:29+00:00","index":105,"fulltext":""},{"type":"reviewerAgreed","content":"217934478511274887845882710077480812645","date":"2026-05-07T12:38:02+00:00","index":80,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-24T10:12:29+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-24 10:12:28","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9308920","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9308920","identity":"rs-9308920","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}