Genome-wide identification reveals the regulatory roles of the TATA-box binding protein associated factor (TAF) gene family in cotton oil accumulation

Genome-wide identification reveals the regulatory roles of the TATA-box binding protein associated factor (TAF) gene family in cotton oil accumulation | 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 reveals the regulatory roles of the TATA-box binding protein associated factor (TAF) gene family in cotton oil accumulation Jie Gao, Li Wang, Pan Feng, Bing Jia, Siqi Chen, Jian Zhang, Weixiao Zhao, and 8 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8328337/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract TBP-associated factors (TAFs), along with the TATA-box binding protein (TBP), com-pose transcription factor IID (TFIID), which is an essential complex for transcription initiation and regulation. TAFs are involved in regulating plant development. However, genome-wide identification and expression pattern analysis of TAFs in cotton remain unreported. In this study, a total of 124 TAF genes were identified in four Gossypium species. TAFs were divided into six subgroups based on the phylogenetic tree analysis. Analysis of collinearity and selection pressure indicated that TAF gene pairs have undergone strong purifying selection during evolution. Gene expression analysis of GhTAFs revealed that TAFs are expressed at all stages of cotton seed and fiber development. Furthermore, a set of key GhTAF genes was identified, which are specifically or constitutively highly expressed across multiple stages of cotton seed and fiber development. Based on WGCNA, we identified Ghir_D11G035840 as a candidate gene involved in lipid accumulation regulation, and further constructed a putative co-expression interaction network for this gene to elucidate its functional connections. Overexpression of Ghir_D11G035840 in Arabidopsis thaliana resulted in a significant increase in seed oil content, thereby confirming its functional role in regulating oil accumulation. Furthermore, a putative interaction network derived from the co-expression network revealed that Ghir_D11G035840 may interact with numerous genes to regulate both ovule development and lipid synthesis. These results provide valuable genetic resources and a theoretical basis for subsequent investigations into the molecular mechanisms underlying cotton seed oil synthesis and fiber quality formation using functional genomics approaches. cotton TAFs genome-wide expression pattern WGCNA Ghir_D11G035840 Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction TAFs, collectively known as TATA-box binding protein associated factors, together with TATA-binding protein (TBP), constitute the general transcription factor complex TFIID (Albright and Tjian 2000; Louder, et al. 2016). TAFs have been demonstrated to perform diverse functions in transcriptional regulation, including the following: (1) enabling broad promoter recognition; (2) acting as coactivators that relay signals from activators to the basal transcription apparatus; (3) mediating communication between TFIID and nucleosomes; (4) supporting activator-independent transcription reinitiation; and (5) interacting with components involved in epigenetic modifications (Luna-Arias and Castro-Muñozledo 2024; Ruppert, et al. 1993). According to their role in the basal transcription machinery, TAFs were expected to be required for accurate transcription of all genes. Studies on TAFs on human, fruit flies and yeast show that TAFs exhibit a high degree of conservation from yeast to humans (Aoyagi and Wassarman 2001; Luna-Arias and Castro-Muñozledo 2024; Uffenbeck and Krebs 2006). A growing body of research has underscored the pivotal functions of TAFs in regulating plant development and stress responses, particularly in the model plant Arabidopsis . The finding of Bertrand et al. (Bertrand, et al. 2005) show that, AtTAF1 gene acts as a coactivator that integrates light signals and promotes histone acetylation, thereby activating light-induced gene transcription. In Arabidopsis , AtTAF5 ( At5g25150 ) is essential for the complete life cycle of plants, affecting processes such as male gametogenesis, inflorescence meristems, and pollen tube growth (Mougiou, et al. 2011). Moreover, AtTAF6 ( At1g04950 ) is also involved in the regulation of pollen tube growth (Lago, et al. 2005). In Arabidopsis , overexpression of AtTAF10 ( At4g31720 ) enhances its tolerance to salt stress (Gao, et al. 2006), and overexpressing Flaveria trinervia FtTAF10 in Arabidopsis caused restricted floral indeterminacy and leaf deformation (Furumoto, et al. 2005). AtTAF12b ( At1g17440 ) contributes to environment-responsive root growth control through an unfolded protein response (Kim, et al. 2022). AtTAF13 ( At1g02680 ) cooperates with the FIS-PRC2 complex to mediate transcriptional regulation during seed development (Lindner, et al. 2013). Furthermore, OsTAF2 regulates grain size in rice via its role in modulating cell division (Jiang, et al. 2023). However, studies on the TAF (Transcription Initiation Factor II D-Associated Factors) family have been rarely reported, especially in cotton ( Gossypium spp.). Cotton ( Gossypium spp.) is a critical economic crop, providing both textile fiber and edible oil. It also represents a premier model system for investigating polyploidy and evolution, making it highly valuable for genetic and evolutionary studies (Senchina, et al. 2003). Evidence from previous studies suggests that all diploid cotton species may evolved from a common ancestor. and subsequently diverged into eight genomic groups A-G and K 3 (Paterson, et al. 2012). Approximately One to two million years ago, hybridization between an African A-genome progenitor and an American D-genome progenitor gave rise to tetraploid cotton (Senchina, et al. 2003). Currently, there are four cultivated cotton species, including the diploids G. arboreum and G. herbaceum , and the allotetraploids G. hirsutum and G. barbadense . Recently, the completion of genome sequencing for these four cotton species has paved the way for studies of gene function and evolution at the whole-genome level (Li, et al. 2014; Wang, et al. 2012; Yuan, et al. 2015; Zhang, et al. 2015). In this study, we identified several TAF genes in four cotton species, including two diploid species ( G. arboreum and G. raimondii ) and two tetraploid species ( G. hirsutum and G. barbadense ). The TAF gene family was analyzed for its protein physicochemical properties, phylogenetic analysis, chromosome location, collinearity analysis, Ka/Ks ratios. The GhTAFs family was also analyzed for its gene structure, conserved motifs, domain alignment. In addition, the functions and evolutionary characteristics of the GhTAF gene family were explored by expression analysis and weighted gene co-expression network analysis (WGCNA) of GhTAFs . Based on WGCNA, we identified Ghir_D11G035840 as a TAF family gene that may be involved in regulating oil accumulation. Its functional role has been validated through overexpression assays in Arabidopsis thaliana . These findings will also provide future studies of the structure and function of the GhTAF gene family. Material and methods Identification of TAF Gene Family Members in Gossypium Species The protein sequences of AtTAFs were obtained from the A. thaliana genome database (http://www.arabidopsis.org). To identify TAF family members, we aligned the protein sequences of four Gossypium genomes, namely, G. arboreum (A2), G. raimondii (D5), G. hirsutum (AD1), and G. barbadense (AD2), against the reference set from Arabidopsis thaliana using a sequence alignment approach with an E-value cutoff of 1e −10 . Subsequently, the presence of conserved TAF protein domains in these genes was examined using the Batch CD search tool (https://www.ncbi.nlm.nih.gov/Structure/bwrpsb/bwrpsb.cgi), and these domains were further validated using the online SMART database (Letunic, et al. 2021). The protein length (pI) and molecular weight (MW) of all candidate TAF genes in Gossypium Species were determined using ExPASy via TBtools-II (Chen, et al. 2023). Phylogenetic Analysis The TAF members in the four Gossypium species were subjected to multiple sequence alignment using (http://www.ebi.ac.uk/Tools/msa/clustalw2) for mutual sequence comparison. To elucidate the evolutionary relationships, a phylogenetic tree was constructed from the multiple sequence alignments of TAF members using the neighbor-joining method in MEGA (v.11.0.13). Branch support evaluated by bootstrapping with 1000 replicates. The final phylogenetic tree was generated and visualized by employing the EvolView online platform (https://www.evolgenius.info/evolview-v3). Chromosomal Distribution, and Gene Duplications The chromosomal distribution of TAF genes was mapped for the four cotton species using MapChart (v.2.2) tool (Voorrips 2002)，based on positional information derived from the corresponding genome GFF3 files. MCScanX was employed to detect gene duplication and synteny, with subsequent visualization performed in TBtools-II. The selective pressure on the TAF gene family was evaluated by calculating the Ka (non-synonymous substitution) and Ks (synonymous substitution) rates. Gene Structure, Motif, and Domain Analysis For the characterization of TAF gene structures, the corresponding gene feature files for the four Gossypium species were retrieved (https://yanglab.hzau.edu.cn/CottonMD) (Yang, et al. 2023). The conserved motifs in TAF proteins were analyzed using the Multiple Em for Motif Elicitation (MEME) (http://memesuite.org/) tool. The exon-intron architecture of TAF genes was analyzed with TBtools-II. Protein domains in TAFs were predicted using the conserved domain database (CDD) resource at NCBI. TBtools-II software was employed for visualization. RNA extraction, library construction and sequencing RNA was extracted from ovule at 10 days post-anthesis (10 DPA), 15 DPA, 20 DPA, 25 DPA, and 30 DPA for RNA sequencing (RNA-seq). The integrity of the isolated total RNA was assessed by electrophoresis on a 1.5% agarose gel, followed by visual inspection of the 18S and 28S rRNA bands. Whereafter, the concentration of RNA was determined using a NanoDrop 2000 spectrophotometer (Thermo Fisher Scientific, Waltham, MA, USA), with the OD260/OD280 absorbance ratio employed for quantification. Altogether, twenty cDNA libraries, comprising ovules and fibers from two genotypes across five developmental stages, were constructed. These libraries were subsequently sequenced on an Illumina NovaSeq 6000 platform (provided by Illumina, located in San Diego, CA, USA) for transcriptomic analysis. Expression Analysis of GhTAF in Different Tissues T111 and T113 are two cotton accessions that exhibit significant differences in fiber quality and cottonseed oil (Song, et al. 2024). To investigate the expression levels of the TAF gene family during the development of fibers and ovules in cotton, we analyzed the expression levels of the GhTAF gene family in both materials across different developmental stages of fiber and ovule. We have submitted all the sequenced raw datasets to the NCBI short read archives (SRA; accession number PRJNA1378545). Weighted gene co-expression network analysis Gene co-expression networks were constructed via Weighted Gene Co-expression Network Analysis (WGCNA) using the corresponding R software package (Zhang and Horvath 2005). Genes with a maximum expression value (among the nineteen samples) of less than five were filtered out. In total, 28,255 genes were selected to constructed the weighted gene co-expression network. A soft-thresholding criterion was selected based on an R 2 value greater than 0.85. To support the classification of modules, the parameter for module merging (denoted as mergeCutHeight) was established at 0.35. For assessing the module-phenotype association, the Pearson correlation coefficient between each module's eigengene and the corresponding phenotype was computed via Python's statistics package. For the construction of the putative interaction network, Pearson’s correlation coefficient was used to quantify the interaction weight between the target gene and each candidate gene. The resulting interaction network was imported into Cytoscape (v.3.7.1) for visualization. Subcellular localization analysis For subcellular localization of Ghir_D11G035840 , the full-length CDS of Ghir_D11G035840 was cloned into the p1300-GFP vector to generate Pro35S:: Ghir_D11G035840 -GFP constructs, which were transformed into Agrobacterium tumefaciens GV3101 for transient expression in tobacco leaves. The GFP fluorescence in leaf epidermal cells was observed using a laser scanning confocal microscope (TCS SP8, Leica, Germany). Isolation of the candidate gene Ghir_D11G035840 and Arabidopsis transformation The Ghir_D11G035840 genes were cloned into the expression vector pEarleyGate 101 using BP and LR reactions. Recombinant vectors were transformed into Agrobacterium GV3101, and transgenic positive Arabidopsis was obtained by floral dip method. Following genetic transformation and three consecutive generations of self-pollination, homozygous T3 transgenic lines were obtained for subsequent experimental analysis. Results Genome-wide identification of TAF members in Gossypium For the identification of all TAF members in G. hirsutum , G. barbadense , G. arboreum , and G. raimondii , we initially obtained the protein sequences of AtTAFs in Arabidopsis and performed sequence alignment between these AtTAF sequences and all protein sequences from the four Gossypium species. Since there is currently no biochemical evidence in Arabidopsis demonstrating that TAF14 and TAF15 are TBP-associated factors, these two proteins are not considered TATA-box binding protein associated factor. Protein sequences sharing similarity with AtTAFs were identified as candidate TAF members. Next, the obtained protein sequences were filtered via CDD and SMART. Ultimately, a total of 22, 19, 39, and 44 TAF genes were identified in G. arboreum (A2), G. raimondii (D5), G. hirsutum (AD1), and G. barbadense (AD2), respectively. The number of TAF genes in tetraploid cotton species is approximately twice that in diploid cottons species, which suggests that TAF genes are relatively evolutionarily conservative. Following the identification of TAF family members, we proceeded to characterize the molecular properties of these TAFs, including gene id, gene location, direction, isoelectric point (PI), molecular weight (Mw) ( Table 1 ). There were significant differences in protein length (from 77 to 1897 aa), molecular weight (from 9.00 to 214.20 kDa), and isoelectric point (from 4.38 to 10.21). Table 1 The basic information and characteristics of the TAF genes in G. arboreum , G. barbadense , G. hirsutum , and G. raimondii Sequence ID Chr Location Strand AA Mw (Da) pI Instability Index Aliphatic Index Garb_01G013900 Chr01 65032255-65037588 + 518 55133.31 9.79 79.29 60.85 Garb_01G015420 Chr01 77907291-77912577 + 890 97131.11 6.37 57.7 64.09 Garb_04G004880 Chr04 8196528-8197651 - 333 36579.24 8.75 31.48 71.74 Garb_05G004260 Chr05 4031203-4036380 - 669 74483.86 6.11 51.61 79.15 Garb_05G004680 Chr05 4313731-4315658 + 136 15411.31 5.78 42.47 73.9 Garb_05G025290 Chr05 25861243-25866683 - 941 102468.6 9 56.28 59.33 Garb_05G041030 Chr05 91920731-91924327 + 526 58458.79 8.32 49.54 97.19 Garb_06G001980 Chr06 1366291-1377539 - 623 67821.48 9.82 78.45 69.9 Garb_07G001070 Chr07 1148179-1150183 - 128 14587.63 5.3 46.16 73.83 Garb_07G019690 Chr07 91436632-91441502 - 534 59698.04 6.91 47.15 100.97 Garb_07G025290 Chr07 100971229-100991467 + 1336 149629.01 6.45 45.36 82.81 Garb_07G025840 Chr07 101810338-101812488 - 136 15232.07 5.48 45.71 77.5 Garb_07G026090 Chr07 102080637-102086678 + 668 74171.43 6.04 50.98 79.12 Garb_08G031810 Chr08 140970417-140974799 + 200 21661.7 4.38 49.74 72.15 Garb_09G000890 Chr09 1802982-1804789 - 377 41309.39 5.77 34.11 79.84 Garb_09G018840 Chr09 79195739-79197539 + 370 40933.47 7.65 44.93 73.11 Garb_10G005830 Chr10 5627362-5635147 + 634 69185.83 10.04 81.7 64.84 Garb_10G011560 Chr10 22634276-22637193 + 128 14396.38 5.74 45.31 73.83 Garb_10G014090 Chr10 36481373-36500121 + 1896 214021.59 5.66 51.46 73.61 Garb_10G022130 Chr10 108330082-108334424 + 228 25641.72 5.55 69.18 67.54 Garb_13G006940 Chr13 10557079-10561782 - 476 54344.2 9.35 41.31 95.25 Garb_13G028360 Chr13 127863358-127868797 + 228 25759.91 5.73 73.6 67.11 Grai_04G019550 GrChr04 44391805-44392730 + 287 31536.42 6.2 31.1 68.95 Grai_05G006680 GrChr05 4767536-4769357 + 136 15366.21 5.52 40.07 76.76 Grai_05G028080 GrChr05 23423371-23428901 - 940 102311.4 8.93 55.54 58.96 Grai_05G043170 GrChr05 55896839-55900432 + 526 58462.72 8.09 49.74 97 Grai_05G043190 GrChr05 55910612-55914450 + 352 39571.85 6.72 41.58 91.14 Grai_06G026080 GrChr06 59000989-59008266 + 623 67886.68 9.82 79.13 68.65 Grai_07G019370 GrChr07 42580076-42584912 - 534 59564.98 7.7 46.72 100.97 Grai_07G024740 GrChr07 49114000-49134358 + 1354 151944.78 6.49 44.53 82.78 Grai_07G025330 GrChr07 49803372-49805492 - 142 15968.32 5.49 40.06 98.24 Grai_08G001630 GrChr08 1189468-1193912 - 197 21494.55 4.38 48.41 70.76 Grai_09G001030 GrChr09 2156332-2158115 - 377 41299.38 5.77 33.81 79.58 Grai_09G018220 GrChr09 41294141-41295253 + 370 40933.46 6.62 45.82 75.22 Grai_10G005830 GrChr10 4760846-4768756 + 636 69514.16 9.82 81.87 64.94 Grai_10G005870 GrChr10 4786403-4787720 + 175 19787.34 5.07 51.66 79.09 Grai_10G009520 GrChr10 10071354-10074351 - 128 14366.35 5.74 45.97 74.61 Grai_10G014450 GrChr10 21143575-21162309 + 1897 214201.24 6.15 51.01 76.19 Grai_10G021840 GrChr10 46676987-46681820 + 228 25636.66 5.55 66.26 64.56 Grai_11G000600 GrChr11 367221-369228 - 371 41196.57 6.69 54.58 71.51 Grai_13G028300 GrChr13 57183898-57187199 + 228 25668.72 5.41 72.79 64.56 Ghir_A01G012900.1 GhA01 55598182-55603661 - 522 55590.78 9.79 77.69 60.38 Ghir_A04G001090.1 GhA04 1911061-1915067 - 526 58504.88 8.29 49.54 97 Ghir_A05G006100.1 GhA05 5665014-5669831 - 659 73518.01 6.18 49.75 79.76 Ghir_A05G006490.1 GhA05 5953844-5955876 + 136 15384.24 5.52 40.7 73.9 Ghir_A05G026730.1 GhA05 28222884-28228446 - 908 98755.25 8.87 57.75 57.73 Ghir_A05G038480.1 GhA05 101478501-101479659 + 364 40318.54 6.48 30.46 79.81 Ghir_A07G003630.1 GhA07 3972163-3977751 - 627 69704.6 6.52 48.86 82.58 Ghir_A07G003870.1 GhA07 4299614-4302029 + 136 15232.07 5.48 45.71 77.5 Ghir_A07G009370.1 GhA07 14259030-14264060 + 534 59699.08 6.69 47.66 101.14 Ghir_A07G024160.1 GhA07 96611166-96613434 + 91 10413.96 4.64 35.73 96.37 Ghir_A08G001610.1 GhA08 1367945-1372362 - 200 21661.7 4.38 49.74 72.15 Ghir_A09G000890.1 GhA09 2183869-2185807 - 377 41369.44 5.67 34.04 79.58 Ghir_A09G015600.1 GhA09 71751449-71752559 + 312 34911.71 8.68 46.12 73.24 Ghir_A10G005110.1 GhA10 5324308-5332068 + 634 69300.02 10.14 80.57 65.3 Ghir_A10G005150.1 GhA10 5352069-5353455 + 175 19817.43 5.07 50.56 79.09 Ghir_A10G010120.1 GhA10 21088043-21091147 + 128 14426.4 5.74 47.47 73.83 Ghir_A10G014450.1 GhA10 77732138-77749919 - 1892 213731.43 5.75 51.61 74.13 Ghir_A10G017230.1 GhA10 95831078-95837045 + 320 36732.44 7.64 62.28 69.12 Ghir_A11G034980.1 GhA11 122694995-122697264 + 371 41459.88 6.44 53.66 71.24 Ghir_A13G022670.1 GhA13 106481607-106487183 + 228 25703.84 5.92 73.45 66.27 Ghir_D01G011820.1 GhD01 20230112-20236251 + 629 68363.49 8.8 57.43 65.87 Ghir_D01G013920.1 GhD01 32442370-32447828 - 424 46139.82 9.72 64.28 69.93 Ghir_D04G004600.1 GhD04 6713089-6715354 - 392 43226.83 6.34 30.77 79.82 Ghir_D05G006570.1 GhD05 5282671-5284105 + 136 15366.21 5.52 40.07 76.76 Ghir_D05G026740.1 GhD05 25361486-25367058 - 940 102261.42 8.93 56.26 58.77 Ghir_D06G021850.1 GhD06 65353017-65360767 + 570 62066.26 9.87 79.3 69.19 Ghir_D07G003890.1 GhD07 4027085-4029648 + 136 15232.07 5.48 45.71 77.5 Ghir_D07G004430.1 GhD07 4721369-4741789 - 1346 150955.55 6.43 45.24 82.98 Ghir_D07G009490.1 GhD07 11434212-11439121 + 534 59595 7.7 46.56 100.79 Ghir_D08G001640.1 GhD08 1352898-1357344 - 198 21535.64 4.41 48.41 71.87 Ghir_D09G000870.1 GhD09 2162143-2164264 - 377 41394.52 5.78 34.36 79.31 Ghir_D09G015070.1 GhD09 43508727-43509839 + 370 40997.51 6.58 46.88 74.16 Ghir_D10G005890.1 GhD10 5353890-5361763 + 594 65312.88 10.21 79.32 67.86 Ghir_D10G005930.1 GhD10 5392617-5393950 + 175 19777.31 5.07 53.2 79.09 Ghir_D10G009150.1 GhD10 10668699-10671687 - 128 14352.33 5.73 43.29 74.61 Ghir_D10G013080.1 GhD10 21641794-21660446 + 1886 212647.32 5.66 51.1 75.09 Ghir_D10G018680.1 GhD10 51418674-51424453 + 242 27543.89 5.84 66.96 64.83 Ghir_D11G035840.1 GhD11 72459161-72460841 + 366 40620.92 7.01 53.56 70.08 Ghir_D13G023330.1 GhD13 61263758-61267625 + 228 25668.72 5.41 72.79 64.56 Gbar_A01G011200.1 GbA01 25962898-25967770 + 940 102756.42 7.08 56.64 63.79 Gbar_A01G013230.1 GbA01 52224808-52230226 - 522 55590.78 9.79 77.69 60.38 Gbar_A04G000930.1 GbA04 1454496-1458150 - 380 42982.99 8.58 49.21 103.95 Gbar_A05G005640.1 GbA05 5314435-5319582 - 627 69849.55 5.63 52.22 78.69 Gbar_A05G006040.1 GbA05 5584844-5586755 + 136 15384.24 5.52 40.7 73.9 Gbar_A05G025780.1 GbA05 27014699-27020603 - 941 102381.57 9 56.7 59.85 Gbar_A05G037650.1 GbA05 95215368-95216526 + 364 40318.54 6.48 30.46 79.81 Gbar_A06G020740.1 GbA06 113715155-113726674 + 622 67683.37 9.74 79.53 69.86 Gbar_A07G003410.1 GbA07 3999121-4005238 - 654 72390.39 6.01 50.98 78.27 Gbar_A07G003660.1 GbA07 4261018-4263142 + 153 17405.56 5.43 42.99 79.08 Gbar_A07G009140.1 GbA07 14051978-14057020 + 534 59699.08 6.69 47.66 101.14 Gbar_A07G023840.1 GbA07 91647527-91649832 + 91 10487.08 4.75 34.55 92.09 Gbar_A08G001560.1 GbA08 1326516-1330935 - 200 21675.72 4.38 49.74 72.65 Gbar_A09G000960.1 GbA09 2123646-2126079 - 377 41339.42 5.77 34.11 79.58 Gbar_A09G015820.1 GbA09 67951827-67952939 + 370 40845.41 8.11 44.18 73.11 Gbar_A10G005730.1 GbA10 5569764-5582319 + 634 69271.96 10.04 80.68 65.3 Gbar_A10G005770.1 GbA10 5597727-5599006 + 175 19817.43 5.07 50.56 79.09 Gbar_A10G010930.1 GbA10 21075070-21078221 + 128 14426.4 5.74 47.47 73.83 Gbar_A10G015330.1 GbA10 74022401-74041303 - 1896 214124.8 5.68 51.29 73.72 Gbar_A10G018160.1 GbA10 90311619-90317524 + 320 36704.41 7.67 60.64 70.34 Gbar_A11G009500.1 GbA11 8445257-8445618 - 77 9006.33 6.72 51.68 64.68 Gbar_A11G034380.1 GbA11 112664951-112667060 + 373 41697.1 6.74 57.24 70.08 Gbar_A13G022920.1 GbA13 107501051-107506519 + 228 25703.84 5.92 73.45 66.27 Gbar_D01G011760.1 GbD01 20308205-20314078 + 872 95166.93 6.76 54.57 65.31 Gbar_D01G013930.1 GbD01 32262924-32268206 - 516 54984.02 9.87 79.41 59.38 Gbar_D04G004560.1 GbD04 6274837-6277074 - 392 43285.86 6.05 29.71 79.82 Gbar_D05G006050.1 GbD05 4872830-4877678 - 603 66898.59 6.48 46.18 79.72 Gbar_D05G006460.1 GbD05 5148292-5150334 + 136 15414.3 5.78 40.7 76.76 Gbar_D05G026630.1 GbD05 25105293-25111109 - 940 102404.61 9 56.26 58.86 Gbar_D05G038870.1 GbD05 62312756-62316415 + 526 58562.88 8.09 49.4 97.36 Gbar_D06G021530.1 GbD06 61540090-61547588 + 621 67564.31 9.82 79.71 68.24 Gbar_D07G003880.1 GbD07 3975665-3977988 + 186 21300.04 8.4 53.72 76.13 Gbar_D07G004470.1 GbD07 4716453-4722704 - 513 57488.82 6.16 43.6 89.73 Gbar_D07G009550.1 GbD07 11369872-11374664 + 534 59633.09 7.68 46 101.52 Gbar_D08G001560.1 GbD08 1231334-1235798 - 198 21535.64 4.41 48.41 71.87 Gbar_D09G000900.1 GbD09 2004189-2006026 - 377 41394.52 5.78 34.36 79.31 Gbar_D09G015550.1 GbD09 42115829-42116941 + 370 40949.51 6.62 45.76 76.27 Gbar_D10G005610.1 GbD10 4970942-4978790 + 634 69224.87 9.93 81.61 64.53 Gbar_D10G005650.1 GbD10 4996498-4997834 + 175 19787.34 5.07 51.66 79.09 Gbar_D10G008880.1 GbD10 10172290-10174608 - 91 10389.91 4.62 32.14 92.09 Gbar_D10G012840.1 GbD10 21280556-21299560 + 1893 213608.29 5.7 51.5 74.09 Gbar_D10G018280.1 GbD10 49046196-49051900 + 228 25641.72 5.55 65.37 67.54 Gbar_D11G036400.1 Scaffold3329 418050-420228 - 373 41435.76 6.74 56.36 70.86 Gbar_D13G023380.1 GbD13 58256028-58259823 + 228 25668.72 5.41 72.79 64.56 Phylogenetic Analysis To investigate the evolutionary trajectories of TAF family members in cotton, a phylogenetic tree was constructed using the protein sequences of 18 Arabidopsis thaliana TAFs ( AtTAFs ) and 124 TAFs derived from 4 cotton species. In the phylogenetic tree ( Fig. 1 ), we noticed that the TAF proteins were clustered into 6 subfamilies (I-VI) with 32 members in subfamily I, 21 in II, 8 in III, 17 in IV, 26 in V, and 38 in VI. Among them, TAF8 contained the largest number of members, whereas TAF2 had the fewest, indicating that members of different TAFs have undergone different evolutionary events involving duplication and deletion. Chromosome Localization, Collinearity Analysis, and Selection Pressure Analysis To visually illustrate the chromosomal distributions of the TAF gene family, localization analyses were conducted for all TAF genes identified in cotton. The results showed that TAFs were distributed across different chromosomes in the four cotton species ( Fig. 2 ). In G. arboreum , twenty-two GaTAFs were located on nine chromosomes (Chr01, Chr04, Chr05, Chr06, Chr07, Chr8, Chr9, Chr10 and Chr13). In G. raimondii , nineteen GrTAFs were located on seven chromosomes (Chr04, Chr05, Chr06, Chr07, Chr8, Chr9, Chr10, Chr11 and Chr13). In G. hirsutum , 39 GhTAF genes were mapped to 19 chromosomes, comprising nine chromosomes from the A t subgenome and ten chromosomes from the D t subgenome. In Gossypium barbadense , 44 GbTAFs were mapped on 20 chromosomes, including ten chromosomes from the A t subgenome and ten chromosomes from the D t subgenome. Gene duplication events play a pivotal role in plant evolution, among which whole-genome duplication, segmental duplication, and tandem duplication are recognized as the major drivers underlying the expansion of gene families (Cannon, et al. 2004). To detect gene duplication events in the TAF gene family, we analyzed the intra- and interspecific collinearity of TAF genes across the four cotton species. As we all know, Gossypium hirsutum and Gossypium barbadense are allotetraploid species derived from two distinct diploid progenitors, so we generated collinearity plots to compare TAF gene collinearity between diploid and tetraploid cotton species. To identify gene duplication events, we assessed the interspecific collinearity of the 124 previously identified TAF genes across four cotton species, namely Gossypium arboreum , G. hirsutum , G. barbadense , and G. raimondii ( Fig. 3 A, B). A total of 25 and 26 pairs of gene duplication from Ga-Gh and Ga-Gb comparison, whereas 25 and 21 pairs were found in the Gr-Gh and Gr-Gb comparisons. The results show that there were 25 and 21 collinear TAF genes within the A subgenomes and D sub-genomes, respectively. The intraspecific collinearity of 83 TAF genes was also assessed in G. hirsutum and G. barbadense , including collinearity between the A and D subgenomes within each species ( Fig. 3 C, D). Most TAFs exhibit good collinearity within the A t -genome or D t -genome of the two tetraploid cotton species. To estimate the correlation of repeating genes over along evolutionary history, Ka/Ks values in GhTAF gene homologous pairs were calculated based on different selection pressures such as purification, neutral, and active selection. According to the Ka/Ks analysis (Table S1), the Ka/Ks values were below 1.0, indicating that these GhTAF genes underwent strong purification selection during evolution. A total of 48 gene pairs exhibited a Ka/Ks ratio of less than 0.5, and 7 gene pairs were between 0.5 and 1, leading us to hypothesize that these TAF gene pairs have undergone strong purifying selection during evolution. Structure Analysis Structural difference among genes is a key factor shaping the evolution of gene families. To better understand the structural characteristics of TAF genes in G. hirsutum , we analyzed the gene structure, motif, and conserved domains of TAF genes in this species, with the analysis conducted based on the constructed phylogenetic tree ( Fig. 4 ). Online MEME analysis was performed to identify 20 conserved motifs among the 39 GhTAF genes. Most members exhibit motif 6, but different types of GhTAF motifs exhibit distinct distributions. One plausible explanation is that many TAFs appear to have evolved independently. The gene structure of GhTAF showed that the number of exons and introns shows considerable variation across different types of genes, and a few TAFs only contained a single exon. These results are consistent with the clustering pattern of the phylogenetic tree. Analysis of GhTAFs Gene Expression in Different Tissues of upland cotton Although the sequence structure was thoroughly investigated, their potential roles in ovule and fiber development remained unclear. In this study, we integrated published transcriptome data comprising 20 samples (T111 and T113 across five developmental stages). Transcriptome data for fiber development were obtained from previous studies, whereas data for ovule development are reported here for the first time (Table S2). T111 shows high oil content and low lint percentage, whereas T113 displays the opposite traits: low oil content and high lint percentage. The spatiotemporal expression patterns of genes are strongly correlated with their biological functions. Therefore, characterizing the transcription landscape of GhTAFs is essential for illustrating their roles in ovule and fiber development. Our analysis revealed distinct, tissue-specific transcription patterns for several GhTAFs . According to the results of expression clustering, the GhTAF genes were categorized into 5 clusters (I-V) in ovule ( Fig. 5 ). Among these, cluster I and IV showed almost no expression, while cluster II, III, and V exhibited high expression levels. In cluster I, the expression levels of Ghir_A09G015600 and Ghir_D09G015070 were undetectable in terms of expression across all experimental periods in T111 and T113, whereas Ghir_D10G009150 displayed significantly higher expression level in ovules from line T113 relative to line T111. In cluster IV, most GhTAF genes exhibited two key expression patterns: first, their expression levels were higher in T111 than in T113; second, their expression at 10, 15 and 20 DPA was higher than that at 25 and 30 DPA. Ghir_A07G003870 , Ghir_A01G012900 , and Ghir_D10G005890 were typical examples of these patterns. Meanwhile, GhTAFs were clustered into 2 clusters in fiber. Specifically, genes in cluster II exhibited high expression level, while those in cluster I showed low expression level. In cluster II, Ghir_D10G009150 showed significantly higher expression level in the fibers of line T113 than in those of line T111. Nearly half of the GhTAFs , such as Ghir_A05G006100 , Ghir_A10G010120 and Ghir_A07G024160 , exhibited stage-specific expression patterns during ovule and fiber development. In addition, several GhTAF genes exhibited high expression levels in both ovules and fibers; representative examples include Ghir_A10G005150 , Ghir_D07G004430 , and Ghir_D10G005930 . Weighted gene co-expression network analysis (WGCNA) For further investigation of the potential association between GhTAFs and ovule and fiber development, we employed weighted gene co-expression network analysis on the set of retained genes ( Fig. 6 ). The associations between gene regulatory modules (GRMs) and phenotypes traits are multifaceted. In ovule, Purple, turquoise, cyan and green GRMs were highly correlated to ovule at 10 DPA. Brown, magenta and red GRMs were highly correlated to ovule at 15 DPA. Yellow, pink, blue, green and midnightblue GRMs were correlated to ovule at 20 DPA. Meanwhile, cyan and blue GRMs were correlated to ovule at 25 DPA. Black, red, blue and turquoise GRMs were correlated to ovule at 30 DPA (Fig. 6B). Among the retained 28 GhTAFs , we found that these 28 genes were mainly from three GRMs, namely the turquoise, blue and red GRMs, which were involved in ovule development at different stages. A total of 16 GhTAFs were identified to be involved in the turquoise GRM, which is primarily associated with the 10 DPA stage. Similarly, 2 GhTAFs were involved in the red GRM (predominantly associated with 15 DPA). And 9 GhTAFs participated in the blue GRM, which is primarily associated with 20 DPA developmental stage. 20 DPA is a critical developmental stage during which cottonseed oil accumulates rapidly (Song, et al. 2022). Among the 9 GhTAFs , Ghir_D11G035840 exhibits the strongest correlation with the blue GRM. WGCNA results served as a platform to establish a putative interaction network for Ghir_D11G035840 . As Ghir_D11G035840 is a member of the blue module, we constructed its interaction network with other blue module-associated genes. A total of 133 genes from the blue GRM, which exhibited a Pearson’s correlation coefficient of 0.95 with Ghir_D11G035840 , were identified as candidate interacting genes, including VRN1 and MYB3R4 ( Fig. 6 C and Table S3). VRN1 is a member of the B3 domain transcription factor family. Among this family, ABI3 , FUS3 , LEC2 , etc. are core members that regu-late seed development and lipid synthesis (Moreno 2024; Yang, et al. 2022). They directly activate the expression of key genes involved in lipid biosynthesis (e.g., DGAT , PDAT ) as well as genes encoding seed storage proteins. Although VRN1 , a transcription factor harboring two B3 domains, is primarily associated with vernalization, accumulating evidence supports its definitive role in seed development(Li, et al. 2019; Milec, et al. 2022). The Myb-domain transcription factor MYB3R4 indirectly ensures the normal development of seeds by regulating cell cycle-related genes, indirectly but significantly affects seed size and final yield (Haga, et al. 2011). Additionally, MYB3R4 can directly activate the expression of the cellulose synthase-like gene CSLD5 , thereby facilitating the rapid and efficient assembly of new cell walls at the end of mitosis and providing a structural and cellular foundation for seed development (Lan, et al. 2025). Therefore, Ghir_D11G035840 is likely involved in cottonseed oil accumulation. Ghir_D11G035840 may positively regulates oil content To investigate the in vivo subcellular distribution of Ghir_D11G035840 , we per-formed subcellular localization assays on tobacco leaves. The results demonstrated that Ghir_D11G035840 is localized to the nucleus ( Fig. 7 A). To evaluate the impact of Ghir_D11G035840 on vegetable oil accumulation and ovule development, we produced Ghir_D11G035840 -overexpressing (OE) lines in A. thaliana through Agrobacterium-mediated floral dip transformation with strain GV3101. Subsequently, 8 independent transgenic Arabidopsis thaliana plants were obtained, and their transgenic status was verified by molecular identification and Western blot analysis ( Fig. 7 B-D). Two distinct independent OE lines of Ghir_D11G035840 in A. thaliana were selected from these transformants for phenotypic evaluation. Compared with WT (ecotype Columbia-0), the seed oil content of OE was significantly increased ( Fig. 7 E). Furthermore, the levels of specific fatty acid components, including C18:1 (oleic acid) and C18:2 (linoleic acid), were significantly elevated in the seeds of overexpression lines ( Fig. 7 F). In contrast, no significant difference in thousand-seed weight was detected between the OE lines and WT ( Fig. 7 G). These results suggest that Ghir_D11G035840 may positively regulate seed oil content in plants. Roots have been utilized as an alternative model to study cell elongation in prior cotton genomics studies, owing to their relevance to the cell elongation process shared with fibers. After vernalization and surface sterilization, seeds from both OE lines and WT were plated on 1/2 MS medium and incubated at 22°C with a 16 h light/8 h dark photoperiod for root development monitoring. Following 7 days of cultivation, root length measurements and comparative analysis revealed that OE lines exhibited a no-table reduction in root length compared with the WT control ( Fig. 7 H, I). Root length of the OE lines proved that the expression of Ghir_D11G035840 could repress the fiber elongation. Discussion Nowadays, except for in-depth research on the TAF gene family of Arabidopsis (Lago, et al. 2004) and mungbean (Wu, et al. 2024), research on other plants is relatively shallow. TAFs, as components of the general transcription factor TFIID, with highly conserved sequences from yeast to humans (Hoffman, et al. 1990). Therefore, utilizing bioinformatics tools, such as BLAST, to identify potential homologous proteins is reliable. In our initial screening of the cotton genome, we identified 124 TAFs, distributed across the four cotton species as follows: including 39 genes in G . hirsutum , 44 genes in G. barbadense , 22 in G. arboreum , and 19 in G. raimondii . This distribution suggests that TAF gene family is relatively conserved during cotton genome evolution. These TAF proteins were clustered based on the phylogenetic analysis into six groups ( Fig. 1 ). Typically, the TFIID complex is composed of 13 to 14 distinct TAF subunits. In Arabidopsis thaliana , 18 putative AtTAF proteins were identified (Lago, et al. 2004). TAF proteins are grouped according to their common biochemical feature of being associated in stable complexes with the TATA-binding protein (TBP). So, in order to proof that a putative TAF is a TATA-box binding protein associated factor, a biochemical demonstration of its participation in the TFIID complex is needed. There is currently no exactly biochemical evidence in Arabidopsis demonstrating that AtTAF14 and AtTAF15 are TBP-associated factors, these two proteins are not considered TATA-box binding protein associated factor (Lago, et al. 2004). Furthermore, the Arabidopsis genome lacked an identifiable homolog of TAF3, but now the TAIR database designates protein At5g15570 as TAF3. It is noteworthy that, AtTAF3 and GhTAF3 possesses the BTP and TAF8 domain, instead of TAF3 histone fold domain. Meanwhile, previous researches have proved that there is no homologous TAF3 in the mungbean genome (Wu, et al. 2024) and rice genome (Lago, et al. 2004). In yeast, mutation of TAF3 results in a thermo-sensitive phenotype which can be suppressed by overexpression of TAF10, TAF11, TAF13 or TAF6 (Gangloff, et al. 2001). Therefore, it is hypothesized that TAF3 is absent in plants, while the functional role of TAF3 may be compensated for by other TAF subunits. Gene duplication event is a common phenomenon in plants. Some duplicated genes could be retained in in descendant lineages, which could serve as raw genetic material for adaptive evolution of plants (Flagel and Wendel 2009). The analysis of chromosomal location revealed that TAF family genes from four cotton species were unevenly distributed on their respective chromosomes. In this study, the possible duplication events in TAF gene family were investigated in four cotton species. According to the selection pressure analysis, the Ka/Ks ratio of G. hirsutum and G. barbadense was less than 1 (Table S1), supporting the evolutionary conservation of these genes. The results showed that most TAF genes are slowly evolving. Gene structure analysis showed that exon number of GhTAF genes varied greatly, which might be attributed to the directional evolution of TAF genes in terms of both function and structure over the long course of evolutionary history. Meanwhile the same type of TAFs have similar motifs, structural domains and number of exons, which indicates that the same type of TAFs may be functionally conserved during evolution. Cotton is an economically vital crop that serves as the primary source of natural textile fibers as well as a significant oilseed crop providing edible oil for human consumption. A large number of genes and gene families have been extensively studied in G. hirsutum . Consequently, G. hirsutum was used for gene expression analysis in the present study. Previous studies have demonstrated that TAFs are widely involved in multiple processes of plant development. For example, AtTAF5 (Mougiou, et al. 2011) and AtTAF6 (Lago, et al. 2005) regulates pollen tube growth, AtTAF13 (Lindner, et al. 2013)[13] participate in seed development, and OsTAF2 (Jiang, et al. 2023) positively regulates the grain size by modulating several cell cycle related genes. The pattern of gene expression is directly related to its function. Based on transcriptome data, we analyzed the differential expression of GhTAFs in fiber and ovule ( Fig. 5 ). The expression pattern shows that most half of GhTAFs had stage-specific transcription patterns in ovule and fiber, such as Ghir_A05G006100 , Ghir_A10G010120 , and Ghir_A07G024160 . Several GhTAF genes were found to exhibit high expression levels in both ovules and fibers, with Ghir_A10G005150 , Ghir_D07G004430 , and Ghir_D10G005930 identified as representative examples. Furthermore, the expression level of Ghir_D10G009150 exhibits significant differences between the two lines, T111 and T113. Weighted Gene Co-expression Network Analysis (WGCNA) is a classic systems biology approach that enables the identification of modules of co-expressed genes (Langfelder and Horvath 2008). To systematically identify the transcription factors that play a crucial role in the development of cotton fibers, we performed WGCNA of the TAF family throughout the entire development periods of ovules and fibers, respectively. We found that GhTAF genes participate in the turquoise, blue, and red co-expression modules in the ovule, with these modules linked to the 10, 15, and 20 DPA stages of ovule development. During cotton ovule development, this period falls in the middle stage and is characterized by rapid oil accumulation. At this stage, enzymes such as glycerol-3-phosphoacyltransferase (GPAT) and lysophosphatidylate-acyltransferase (LPAAT) play a crucial role (Cui, et al. 2019; Wang, et al. 2017). They can successively attach fatty acids to glycerol molecules, gradually constructing the molecular structure of triglycerides. During this process, various types of fatty acids are combined in specific proportions and sequences, thereby influencing the fatty acid composition and overall quality of cottonseed oil (Yuan, et al. 2018). At the same time, this stage also corresponds to the early phase of seed size determination. Thus, GhTAFs are hypothesized to regulate plant seed size via several pathways, such as the ubiquitin regulatory pathway, plant hormone pathways (auxin and brassinolide), MAPK signaling pathway, G protein signaling pathway, transcription factor pathway, and IKU signaling pathway (Li and Li 2016). Additionally, we generated transgenic Arabidopsis thaliana lines with overexpression of the target gene Ghir_D11G035840 , followed by phenotypic characterization of seeds. These observations were performed to infer the potential function of the candidate gene in ovule development. Through the integration of transcriptome profiling data and phenotypic analysis of Arabidopsis thaliana , we validated the functional role of the gene Ghir_D11G035840 in ovule development and oil accumulation. Furthermore, we constructed a putative interaction network for Ghir_D11G035840 by leveraging transcriptional abundance information. Notably, B3 domain-containing transcription factors function as central regulators within the regulatory circuitry orchestrating seed development and lipid biosynthesis in this interaction network. Furthermore, MYB3R4 indirectly but markedly impacts seed size and final crop yield by transcriptionally regulating genes involved in cell cycle progression (Haga, et al. 2011). Wei et al. reported that TAF in Drosophila melanogaster may regulate the fatty acid composition of multiple phospholipid classes by modulating the transcription of genes associated with peroxisomal fatty acid β-oxidation (Fan, et al. 2017). Collectively, these characterizations demonstrate the potential roles of TAFs in ovule development and oil accumulation. Conclusion In this study, we identified 124 members of the TAF gene family across four cotton species, which were classified into six subgroups based on phylogenetic analysis. We conducted gene expression analysis, identifying key GhTAF genes that are specifically or persistently highly expressed at multiple cottonseed and fiber developmental stages. Furthermore, based on WGCNA data, we identified Ghir_D11G035840 as a TAF family gene potentially involved in regulating lipid accumulation and ovule development. Further overexpression experiments on Arabidopsis thaliana indicated that Ghir_D11G035840 may positively regulate oil content. These findings will also provide valuable genetic resources and theoretical foundations for future studies on the structure and function of the GhTAF gene family. Declarations Author Contributions YL and JS designed the experiments; JG and LW performed the experiments and collected the data. JG, JS, JZ, PF, BJ, SC, WZ, BZ, JM, WP, MW and KZ analyzed the data. JG wrote the manuscript. JS, LY, YJ, WL and JG revised the manuscript. All authors have read and agreed to the published version of the manuscript. Funding This work was supported by The Key Research and Development Program of Xinjiang (Grant No. 2024B02001-1), the National Natural Science Foundation of China (Grant No. 32501903), the Natural Science Foundation of Henan Province (Grant No. 252300420740), and the Natural Science Foundation of Xinjiang Uygur Autonomous Region (Grant No. 2024D01A132). Data Availability Statement All data generated or analyzed during this study are included in this published article and its additional files. The datasets used and analyzed in the current study are available from the corresponding author upon reasonable request. All the transcriptome raw data we sequenced was deposited in the NCBI short read archives (SRA; accession number PRJNA1378545). Conflicts of Interest The authors declare no conflict of interest. References Albright SR, Tjian R (2000) TAFs revisited: more data reveal new twists and confirm old ideas. 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Nat Genet 44:1098-1103 Wang N, Ma J, Pei W et al (2017) A genome-wide analysis of the lysophosphatidate acyltransferase (LPAAT) gene family in cotton: organization, expression, sequence variation, and association with seed oil content and fiber quality. BMC Genomics 18:218 Wu R, Jia Q, Guo Y et al (2024) Characterization of TBP and TAFs in Mungbean (Vigna radiata L.) and Their Potential Involvement in Abiotic Stress Response. Int J Mol Sci 25:9558 Yang Z, Liu X, Wang K et al (2022) ABA-INSENSITIVE 3 with or without FUSCA3 highly up-regulates lipid droplet proteins and activates oil accumulation. J Exp Bot 73:2077-2092 Yang Z, Wang J, Huang Y et al (2023) CottonMD: a multi-omics database for cotton biological study. Nucleic Acids Res 51:D1446-d1456 Yuan D, Tang Z, Wang M et al (2015) The genome sequence of Sea-Island cotton ( Gossypium barbadense ) provides insights into the allopolyploidization and development of superior spinnable fibres. Sci Rep 5:17662 Yuan Y, Wang X, Wang L et al (2018) Genome-Wide Association Study Identifies Candidate Genes Related to Seed Oil Composition and Protein Content in Gossypium hirsutum L. Front Plant Sci 9:1359 Zhang B, Horvath S (2005) A general framework for weighted gene co-expression network analysis. Stat Appl Genet Mol Biol 4:Article17 Zhang T, Hu Y, Jiang W et al (2015) Sequencing of allotetraploid cotton ( Gossypium hirsutum L. acc. TM-1) provides a resource for fiber improvement. Nat Biotechnol 33:531-537 Additional Declarations No competing interests reported. Supplementary Files SupplementaryMaterials.zip Supplementary Materials: Table S1. The non-synonymous (Ka) to synonymous substitution ratio (Ks) induplicated TAF gene pairs in G. hirsutum and G. barbadense. Table S2. Summary of RNA-seq data quality in 10 libraries. Table S3. Interaction network of Ghir_D11G035840 in the blue module. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-8328337","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":561009984,"identity":"bc16a432-8db3-43ed-85f1-0211a459ad07","order_by":0,"name":"Jie Gao","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Jie","middleName":"","lastName":"Gao","suffix":""},{"id":561009985,"identity":"aaac622e-24a5-4790-847c-c56507504577","order_by":1,"name":"Li Wang","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Li","middleName":"","lastName":"Wang","suffix":""},{"id":561009986,"identity":"45107672-2792-497b-b797-9558b1427602","order_by":2,"name":"Pan Feng","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Pan","middleName":"","lastName":"Feng","suffix":""},{"id":561009987,"identity":"d1543c60-6af6-46ee-92aa-7b867a74c33e","order_by":3,"name":"Bing Jia","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Bing","middleName":"","lastName":"Jia","suffix":""},{"id":561009988,"identity":"9d470b2c-8476-4538-9db7-a75e063150f4","order_by":4,"name":"Siqi Chen","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Siqi","middleName":"","lastName":"Chen","suffix":""},{"id":561009989,"identity":"78b1291b-69a0-4f91-b203-e8a4db4ce6b0","order_by":5,"name":"Jian Zhang","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Jian","middleName":"","lastName":"Zhang","suffix":""},{"id":561009990,"identity":"c3de763b-76b1-4133-8c53-7e78f59fa5c1","order_by":6,"name":"Weixiao Zhao","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Weixiao","middleName":"","lastName":"Zhao","suffix":""},{"id":561009991,"identity":"1694f64f-3cfc-4107-95e7-b29e12244268","order_by":7,"name":"Bingbing Zhang","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Bingbing","middleName":"","lastName":"Zhang","suffix":""},{"id":561009992,"identity":"75e3e20f-559e-4495-bb37-1b3e632200e6","order_by":8,"name":"Jianjiang Ma","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural 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Zheng","email":"","orcid":"","institution":"Engineering Research Centre of Cotton, Ministry of Education / College of Agriculture, Xinjiang Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Kai","middleName":"","lastName":"Zheng","suffix":""},{"id":561009996,"identity":"9287eb0f-3443-4424-a76c-dc4d8037016b","order_by":12,"name":"Jiwen Yu","email":"","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Jiwen","middleName":"","lastName":"Yu","suffix":""},{"id":561009997,"identity":"8c3337a6-834f-4629-9366-4999e551a2c7","order_by":13,"name":"Yajun Liang","email":"","orcid":"","institution":"Cotton Research Institute of Xinjiang Uyghur Autonomous Region Academy of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Yajun","middleName":"","lastName":"Liang","suffix":""},{"id":561009998,"identity":"95b3ccfe-dc93-47dd-8d2e-9b9c4ebc5e7e","order_by":14,"name":"Jikun Song","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAx0lEQVRIiWNgGAWjYFCCYwwMjH8k7PiZmQ8/IEFLg0WyZDtbmgGRWthAWioYN5znUZAgSoN847HEx7w7JJiND/MwGDDU2EQT1GJw4NhhY94zEnxmh3kPPGA4lpbbQFALw/E26Rw2CWazw3wJBowNhwlrkW+AaGHc3MxjIEGUFoYDx45J57ZJMG5gJlYL0C/Jxn/OSCRLHAYGcgIxfpGfcczw4YyKOjv+/sOHH3yosSHCYRIHkDgJBJWDAD9hU0fBKBgFo2CkAwCBmT8Lxo9YggAAAABJRU5ErkJggg==","orcid":"","institution":"National Key Laboratory of Cotton Bio-breeding and lntegrated Utilization, Institute of Cotton Research, Chinese Academy of Agricultural Sciences","correspondingAuthor":true,"prefix":"","firstName":"Jikun","middleName":"","lastName":"Song","suffix":""}],"badges":[],"createdAt":"2025-12-10 14:08:17","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8328337/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8328337/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":98399055,"identity":"805a5bdd-185a-4f5b-bd6f-34aa1e0f83c8","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":371197,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic relationship of TAFs proteins from \u003cem\u003eGossypium hirsutum\u003c/em\u003e, \u003cem\u003eGossypium barbadense\u003c/em\u003e, \u003cem\u003eGossypium arboretum\u003c/em\u003e, \u003cem\u003eGossypium raimondii\u003c/em\u003e, and \u003cem\u003eArabidopsis thaliana\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/1fafd990add1d02fdf8db51f.png"},{"id":98399061,"identity":"700c66b3-fc20-420e-b73c-cac16f5b5dbb","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":63105,"visible":true,"origin":"","legend":"\u003cp\u003eChromosome localization analysis of TAF gene family. (A) \u003cem\u003eG. raimondii\u003c/em\u003e, (B) \u003cem\u003eG. barbadense\u003c/em\u003e, (C) \u003cem\u003eG. hirsutum\u003c/em\u003e (D) \u003cem\u003eG. barbadense\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/e61fab71fad27e270bf7aec2.png"},{"id":98441398,"identity":"e1236aa0-e05b-4557-ab34-3fb36b39a629","added_by":"auto","created_at":"2025-12-17 17:05:19","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":162530,"visible":true,"origin":"","legend":"\u003cp\u003eAnalysis of synteny among multiple \u003cem\u003eGossypium\u003c/em\u003e genomes regarding the TAF genes. (A) Synteny analysis among \u003cem\u003eG. arboreum\u003c/em\u003e, \u003cem\u003eG. hirsutum\u003c/em\u003e (A\u003csub\u003et\u003c/sub\u003e subgenome), and \u003cem\u003eG. barbadense\u003c/em\u003e (A\u003csub\u003et\u003c/sub\u003e subgenome). (B) Synteny analysis among \u003cem\u003eG. raimondii\u003c/em\u003e, \u003cem\u003eG. hirsutum\u003c/em\u003e (D\u003csub\u003et\u003c/sub\u003e subgenome), and \u003cem\u003eG. barbadense\u003c/em\u003e (D\u003csub\u003et\u003c/sub\u003e subgenome). (C) Synteny analysis among \u003cem\u003eG. hirsutum\u003c/em\u003e (A\u003csub\u003et\u003c/sub\u003e subgenome) and \u003cem\u003eG. hirsutum\u003c/em\u003e (D\u003csub\u003et\u003c/sub\u003e subgenome). (D) Synteny analysis among \u003cem\u003eG. barbadense\u003c/em\u003e (A\u003csub\u003et\u003c/sub\u003e subgenome) and \u003cem\u003eG. barbadense\u003c/em\u003e (D\u003csub\u003et\u003c/sub\u003e subgenome).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/207f6e8839d17ea045eaf1db.png"},{"id":98399054,"identity":"cc72176e-c44f-4d46-9e05-8a86a188679a","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":73440,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic relationship, (A)motif, (B)domain, and (C)gene structure of TAF genes from \u003cem\u003eG. hirsutum\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/362e1bb30787977fcbc63fb6.png"},{"id":98441800,"identity":"f9c31f09-2223-473e-9abf-b9a1396682f7","added_by":"auto","created_at":"2025-12-17 17:05:49","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":176784,"visible":true,"origin":"","legend":"\u003cp\u003eTranscriptional expression analysis of \u003cem\u003eGhTAF\u003c/em\u003egene family in (A) ovule and (B) fiber tissues of upland cotton. Purple and yellow represent high and low expression levels respectively.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/47a134470a3445c4f9231805.png"},{"id":98399060,"identity":"e4df99c1-a61d-462b-b3a7-32aa10eefa8a","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":204808,"visible":true,"origin":"","legend":"\u003cp\u003eWGCNA for the ovule transcriptome data in the study. (A)Module assignment and hierarchical clustering of genes based on WGCNA. (B) Heatmap for module-trait relationship. (C) Genome-wide transcriptome data-based construction of the complete interaction network of \u003cem\u003eGhir_D11G035840\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/8b1224d4cab8eda1c6177ba9.png"},{"id":98399058,"identity":"3c4d5e69-eb9f-4a00-a2c9-495fc0a3b8c1","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":464500,"visible":true,"origin":"","legend":"\u003cp\u003eFunctional investigation of the candidate gene \u003cem\u003eGhir_D11G035840\u003c/em\u003e. (A) Subcellular localization observations of 35 S: \u003cem\u003eGhir_D11G035840\u003c/em\u003e-GFP in tobacco. (B) Basta to screen overexpression lines. The seedlings in the red circle are transgenic seedlings. (C) PCR identification of overexpression lines. (D) Detection of \u003cem\u003eGhir_D11G035840\u003c/em\u003e overexpression lines using Western blotting. (E) Oil content of wild type and overexpression lines. (F) FAs composition of wild type and over-expression lines. (G) The root phenotyping of wild type and overexpression lines. (H) Statistics of root length. Asterisks indicate sianicant diterences by two-taled Student's t test (*p \u0026lt; 0.05, **p \u0026lt; 0.01)\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/fc2ec6fe33012693250727c9.png"},{"id":105247284,"identity":"65565a6b-17d5-4543-b9bd-79bc889bdc62","added_by":"auto","created_at":"2026-03-24 01:55:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2892265,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/dd3c7c3f-973f-4af7-b056-f9cb5c22f233.pdf"},{"id":98399056,"identity":"8ec43f9c-2658-443c-bb5d-a00a85ab9c51","added_by":"auto","created_at":"2025-12-17 11:13:06","extension":"zip","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":26301,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSupplementary Materials: Table S1\u003c/strong\u003e. The non-synonymous (Ka) to synonymous substitution ratio (Ks) induplicated TAF gene pairs in G. hirsutum and G. barbadense. \u003cstrong\u003eTable S2\u003c/strong\u003e. Summary of RNA-seq data quality in 10 libraries. \u003cstrong\u003eTable S3\u003c/strong\u003e. Interaction network of \u003cem\u003eGhir_D11G035840\u003c/em\u003e in the blue module.\u003c/p\u003e","description":"","filename":"SupplementaryMaterials.zip","url":"https://assets-eu.researchsquare.com/files/rs-8328337/v1/0f4865dc5fb2f07454b7eb84.zip"}],"financialInterests":"No competing interests reported.","formattedTitle":"Genome-wide identification reveals the regulatory roles of the TATA-box binding protein associated factor (TAF) gene family in cotton oil accumulation","fulltext":[{"header":"Introduction","content":"\u003cp\u003eTAFs, collectively known as TATA-box binding protein associated factors, together with TATA-binding protein (TBP), constitute the general transcription factor complex TFIID (Albright and Tjian 2000; Louder, et al. 2016). TAFs have been demonstrated to perform diverse functions in transcriptional regulation, including the following: (1) enabling broad promoter recognition; (2) acting as coactivators that relay signals from activators to the basal transcription apparatus; (3) mediating communication between TFIID and nucleosomes; (4) supporting activator-independent transcription reinitiation; and (5) interacting with components involved in epigenetic modifications\u0026nbsp;(Luna-Arias and Castro-Muñozledo 2024; Ruppert, et al. 1993).\u003c/p\u003e\n\u003cp\u003eAccording to their role in the basal transcription machinery, TAFs were expected to be required for accurate transcription of all genes. Studies on TAFs on human, fruit flies and yeast show that TAFs exhibit a high degree of conservation from yeast to humans (Aoyagi and Wassarman 2001; Luna-Arias and Castro-Muñozledo 2024; Uffenbeck and Krebs 2006). A growing body of research has underscored the pivotal functions of TAFs in regulating plant development and stress responses, particularly in the model plant \u003cem\u003eArabidopsis\u003c/em\u003e. The finding of Bertrand et al.\u0026nbsp;(Bertrand, et al. 2005)\u0026nbsp;show that, \u003cem\u003eAtTAF1\u003c/em\u003e gene acts as a coactivator that integrates light signals and promotes histone acetylation, thereby activating light-induced gene transcription. In \u003cem\u003eArabidopsis\u003c/em\u003e, \u003cem\u003eAtTAF5\u003c/em\u003e (\u003cem\u003eAt5g25150\u003c/em\u003e) is essential for the complete life cycle of plants, affecting processes such as male gametogenesis, inflorescence meristems, and pollen tube growth\u0026nbsp;(Mougiou, et al. 2011). Moreover, \u003cem\u003eAtTAF6\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAt1g04950\u003c/em\u003e) is also involved in the regulation of pollen tube growth\u0026nbsp;(Lago, et al. 2005). In \u003cem\u003eArabidopsis\u003c/em\u003e, overexpression of \u003cem\u003eAtTAF10\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAt4g31720\u003c/em\u003e) enhances its tolerance to salt stress\u0026nbsp;(Gao, et al. 2006), and overexpressing \u003cem\u003eFlaveria trinervia\u003c/em\u003e \u003cem\u003eFtTAF10\u003c/em\u003e in Arabidopsis caused restricted floral indeterminacy and leaf deformation\u0026nbsp;(Furumoto, et al. 2005). \u003cem\u003eAtTAF12b\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAt1g17440\u003c/em\u003e) contributes to environment-responsive root growth control through an unfolded protein response\u0026nbsp;(Kim, et al. 2022). \u003cem\u003eAtTAF13\u0026nbsp;\u003c/em\u003e(\u003cem\u003eAt1g02680\u003c/em\u003e) cooperates with the FIS-PRC2 complex to mediate transcriptional regulation during seed development\u0026nbsp;(Lindner, et al. 2013). Furthermore, \u003cem\u003eOsTAF2\u003c/em\u003e regulates grain size in rice via its role in modulating cell division\u0026nbsp;(Jiang, et al. 2023). However, studies on the TAF (Transcription Initiation Factor II D-Associated Factors) family have been rarely reported, especially in cotton (\u003cem\u003eGossypium\u003c/em\u003e spp.).\u003c/p\u003e\n\u003cp\u003eCotton (\u003cem\u003eGossypium\u0026nbsp;\u003c/em\u003espp.) is a critical economic crop, providing both textile fiber and edible oil. It also represents a premier model system for investigating polyploidy and evolution, making it highly valuable for genetic and evolutionary studies (Senchina, et al. 2003). Evidence from previous studies suggests that all diploid cotton species may evolved from a common ancestor. and subsequently diverged into eight genomic groups A-G and K\u003csub\u003e3\u0026nbsp;\u003c/sub\u003e(Paterson, et al. 2012). Approximately One to two million years ago, hybridization between an African A-genome progenitor and an American D-genome progenitor gave rise to tetraploid cotton\u0026nbsp;(Senchina, et al. 2003). Currently, there are four cultivated cotton species, including the diploids \u003cem\u003eG. arboreum\u003c/em\u003e and \u003cem\u003eG. herbaceum\u003c/em\u003e, and the allotetraploids \u003cem\u003eG. hirsutum\u003c/em\u003e and \u003cem\u003eG. barbadense\u003c/em\u003e. Recently, the completion of genome sequencing for these four cotton species has paved the way for studies of gene function and evolution at the whole-genome level\u0026nbsp;(Li, et al. 2014; Wang, et al. 2012; Yuan, et al. 2015; Zhang, et al. 2015).\u003c/p\u003e\n\u003cp\u003eIn this study, we identified several TAF genes in four cotton species, including two diploid species (\u003cem\u003eG. arboreum\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;G. raimondii\u003c/em\u003e) and two tetraploid species (\u003cem\u003eG. hirsutum\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;G. barbadense\u003c/em\u003e). The \u003cem\u003eTAF\u003c/em\u003e gene family was analyzed for its protein physicochemical properties, phylogenetic analysis, chromosome location, collinearity analysis, Ka/Ks ratios. The \u003cem\u003eGhTAFs\u003c/em\u003e family was also analyzed for its gene structure, conserved motifs, domain alignment. In addition, the functions and evolutionary characteristics of the \u003cem\u003eGhTAF\u003c/em\u003e gene family were explored by expression analysis and weighted gene co-expression network analysis (WGCNA) of \u003cem\u003eGhTAFs\u003c/em\u003e. Based on WGCNA, we identified \u003cem\u003eGhir_D11G035840\u003c/em\u003e as a TAF family gene that may be involved in regulating oil accumulation. Its functional role has been validated through overexpression assays in \u003cem\u003eArabidopsis thaliana\u003c/em\u003e. These findings will also provide future studies of the structure and function of the \u003cem\u003eGhTAF\u003c/em\u003e gene family.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cp\u003e\u003cstrong\u003eIdentification of TAF Gene Family Members in \u003cem\u003eGossypium \u003c/em\u003eSpecies \u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe protein sequences of \u003cem\u003eAtTAFs\u003c/em\u003e were obtained from the \u003cem\u003eA. thaliana\u003c/em\u003e genome database (http://www.arabidopsis.org). To identify TAF family members, we aligned the protein sequences of four \u003cem\u003eGossypium\u003c/em\u003e genomes, namely, \u003cem\u003eG.\u003c/em\u003e\u003cem\u003earboreum\u003c/em\u003e (A2), \u003cem\u003eG. raimondii\u003c/em\u003e (D5), \u003cem\u003eG. hirsutum\u003c/em\u003e (AD1), and \u003cem\u003eG. barbadense\u003c/em\u003e (AD2), against the reference set from \u003cem\u003eArabidopsis thaliana\u003c/em\u003e using a sequence alignment approach with an E-value cutoff of 1e\u003csup\u003e−10\u003c/sup\u003e. Subsequently, the presence of conserved TAF protein domains in these genes was examined using the Batch CD search tool (https://www.ncbi.nlm.nih.gov/Structure/bwrpsb/bwrpsb.cgi), and these domains were further validated using the online SMART database (Letunic, et al. 2021). The protein length (pI) and molecular weight (MW) of all candidate TAF genes in \u003cem\u003eGossypium\u003c/em\u003e Species were determined using ExPASy via TBtools-II (Chen, et al. 2023).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePhylogenetic Analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe TAF members in the four \u003cem\u003eGossypium\u003c/em\u003e species were subjected to multiple sequence alignment using (http://www.ebi.ac.uk/Tools/msa/clustalw2) for mutual sequence comparison. To elucidate the evolutionary relationships, a phylogenetic tree was constructed from the multiple sequence alignments of TAF members using the neighbor-joining method in MEGA (v.11.0.13). Branch support evaluated by bootstrapping with 1000 replicates. The final phylogenetic tree was generated and visualized by employing the EvolView online platform (https://www.evolgenius.info/evolview-v3).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eChromosomal Distribution, and Gene Duplications\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe chromosomal distribution of TAF genes was mapped for the four cotton species using MapChart (v.2.2) tool (Voorrips 2002)，based on positional information derived from the corresponding genome GFF3 files. MCScanX was employed to detect gene duplication and synteny, with subsequent visualization performed in TBtools-II. The selective pressure on the TAF gene family was evaluated by calculating the Ka (non-synonymous substitution) and Ks (synonymous substitution) rates.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eGene Structure, Motif, and Domain Analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor the characterization of TAF gene structures, the corresponding gene feature files for the four \u003cem\u003eGossypium\u003c/em\u003e species were retrieved (https://yanglab.hzau.edu.cn/CottonMD) (Yang, et al. 2023). The conserved motifs in TAF proteins were analyzed using the Multiple Em for Motif Elicitation (MEME) (http://memesuite.org/) tool. The exon-intron architecture of TAF genes was analyzed with TBtools-II. Protein domains in TAFs were predicted using the conserved domain database (CDD) resource at NCBI. TBtools-II software was employed for visualization.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eRNA extraction, library construction and sequencing\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eRNA was extracted from ovule at 10 days post-anthesis (10 DPA), 15 DPA, 20 DPA, 25 DPA, and 30 DPA for RNA sequencing (RNA-seq). The integrity of the isolated total RNA was assessed by electrophoresis on a 1.5% agarose gel, followed by visual inspection of the 18S and 28S rRNA bands. Whereafter, the concentration of RNA was determined using a NanoDrop 2000 spectrophotometer (Thermo Fisher Scientific, Waltham, MA, USA), with the OD260/OD280 absorbance ratio employed for quantification. Altogether, twenty cDNA libraries, comprising ovules and fibers from two genotypes across five developmental stages, were constructed. These libraries were subsequently sequenced on an Illumina NovaSeq 6000 platform (provided by Illumina, located in San Diego, CA, USA) for transcriptomic analysis.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eExpression Analysis of \u003cem\u003eGhTAF\u003c/em\u003e in Different Tissues\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eT111 and T113 are two cotton accessions that exhibit significant differences in fiber quality and cottonseed oil (Song, et al. 2024). To investigate the expression levels of the TAF gene family during the development of fibers and ovules in cotton, we analyzed the expression levels of the \u003cem\u003eGhTAF\u003c/em\u003e gene family in both materials across different developmental stages of fiber and ovule. We have submitted all the sequenced raw datasets to the NCBI short read archives (SRA; accession number PRJNA1378545).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eWeighted gene co-expression network analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eGene co-expression networks were constructed via Weighted Gene Co-expression Network Analysis (WGCNA) using the corresponding R software package (Zhang and Horvath 2005). Genes with a maximum expression value (among the nineteen samples) of less than five were filtered out. In total, 28,255 genes were selected to constructed the weighted gene co-expression network. A soft-thresholding criterion was selected based on an R\u003csup\u003e2\u003c/sup\u003e value greater than 0.85. To support the classification of modules, the parameter for module merging (denoted as mergeCutHeight) was established at 0.35. For assessing the module-phenotype association, the Pearson correlation coefficient between each module's eigengene and the corresponding phenotype was computed via Python's statistics package.\u003c/p\u003e\n\u003cp\u003eFor the construction of the putative interaction network, Pearson’s correlation coefficient was used to quantify the interaction weight between the target gene and each candidate gene. The resulting interaction network was imported into Cytoscape (v.3.7.1) for visualization.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSubcellular localization analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor subcellular localization of \u003cem\u003eGhir_D11G035840\u003c/em\u003e, the full-length CDS of \u003cem\u003eGhir_D11G035840\u003c/em\u003e was cloned into the p1300-GFP vector to generate Pro35S::\u003cem\u003eGhir_D11G035840\u003c/em\u003e-GFP constructs, which were transformed into Agrobacterium tumefaciens GV3101 for transient expression in tobacco leaves. The GFP fluorescence in leaf epidermal cells was observed using a laser scanning confocal microscope (TCS SP8, Leica, Germany).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eIsolation of the candidate gene \u003cem\u003eGhir_D11G035840\u003c/em\u003e and \u003cem\u003eArabidopsis \u003c/em\u003etransformation\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe \u003cem\u003eGhir_D11G035840\u003c/em\u003e genes were cloned into the expression vector pEarleyGate 101 using BP and LR reactions. Recombinant vectors were transformed into \u003cem\u003eAgrobacterium\u003c/em\u003e GV3101, and transgenic positive \u003cem\u003eArabidopsis\u003c/em\u003e was obtained by floral dip method. Following genetic transformation and three consecutive generations of self-pollination, homozygous T3 transgenic lines were obtained for subsequent experimental analysis.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eGenome-wide identification of TAF members in\u003cem\u003e\u0026nbsp;Gossypium\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor the identification of all TAF members in \u003cem\u003eG. hirsutum\u003c/em\u003e, \u003cem\u003eG. barbadense\u003c/em\u003e, \u003cem\u003eG. arboreum\u003c/em\u003e, and \u003cem\u003eG. raimondii\u003c/em\u003e, we initially obtained the protein sequences of \u003cem\u003eAtTAFs\u003c/em\u003e in \u003cem\u003eArabidopsis\u003c/em\u003e and performed sequence alignment between these \u003cem\u003eAtTAF\u003c/em\u003e sequences and all protein sequences from the four \u003cem\u003eGossypium\u003c/em\u003e species. Since there is currently no biochemical evidence in \u003cem\u003eArabidopsis\u003c/em\u003e demonstrating that TAF14 and TAF15 are TBP-associated factors, these two proteins are not considered TATA-box binding protein associated factor. Protein sequences sharing similarity with \u003cem\u003eAtTAFs\u003c/em\u003e were identified as candidate TAF members. Next, the obtained protein sequences were filtered via CDD and SMART. Ultimately, a total of 22, 19, 39, and 44 TAF genes were identified in \u003cem\u003eG. arboreum\u003c/em\u003e (A2), \u003cem\u003eG. raimondii\u003c/em\u003e (D5), \u003cem\u003eG. hirsutum\u003c/em\u003e (AD1), and \u003cem\u003eG. barbadense\u003c/em\u003e (AD2), respectively. The number of TAF genes in tetraploid cotton species is approximately twice that in diploid cottons species, which suggests that TAF genes are relatively evolutionarily conservative.\u003c/p\u003e\n\u003cp\u003eFollowing the identification of TAF family members, we proceeded to characterize the molecular properties of these TAFs, including gene id, gene location, direction, isoelectric point (PI), molecular weight (Mw) (\u003cstrong\u003eTable 1\u003c/strong\u003e). There were significant differences in protein length (from 77 to 1897 aa), molecular weight (from 9.00 to 214.20 kDa), and isoelectric point (from 4.38 to 10.21).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e1\u003c/strong\u003e The basic information and characteristics of the TAF genes in \u003cem\u003eG. arboreum\u003c/em\u003e, \u003cem\u003eG. barbadense\u003c/em\u003e, \u003cem\u003eG. hirsutum\u003c/em\u003e, and \u003cem\u003eG. raimondii\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSequence ID\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eChr\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLocation\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eStrand\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAA\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMw (Da)\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e\u003cstrong\u003epI\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eInstability Index\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAliphatic Index\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_01G013900\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e65032255-65037588\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e518\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e55133.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e9.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e79.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e60.85\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_01G015420\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e77907291-77912577\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e890\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e97131.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e57.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e64.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_04G004880\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e8196528-8197651\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e333\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e36579.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e31.48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e71.74\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_05G004260\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e4031203-4036380\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e669\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e74483.86\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e51.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.15\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_05G004680\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e4313731-4315658\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e136\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e15411.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.78\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e42.47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e73.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_05G025290\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e25861243-25866683\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n 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style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.91\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e47.15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e100.97\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_07G025290\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e100971229-100991467\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e1336\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e149629.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.45\u003c/p\u003e\n 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style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.12\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_08G031810\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr08\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e140970417-140974799\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e21661.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e4.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e49.74\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e72.15\u003c/p\u003e\n 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17.5258%;\"\u003e\n \u003cp\u003eGarb_09G018840\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e79195739-79197539\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e370\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e40933.47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e7.65\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e44.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e73.11\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_10G005830\u003c/p\u003e\n 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18.5567%;\"\u003e\n \u003cp\u003e36481373-36500121\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e1896\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e214021.59\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e51.46\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e73.61\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGarb_10G022130\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eChr10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e108330082-108334424\u003c/p\u003e\n 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style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e55.54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e58.96\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_05G043170\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e55896839-55900432\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e526\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e58462.72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.09\u003c/p\u003e\n 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style=\"width: 14.433%;\"\u003e\n \u003cp\u003e68.65\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_07G019370\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e42580076-42584912\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e534\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e59564.98\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e7.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e46.72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e100.97\u003c/p\u003e\n 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style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_07G025330\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e49803372-49805492\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e142\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e15968.32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e40.06\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e98.24\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n 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style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e2156332-2158115\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e377\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41299.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e33.81\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.58\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_09G018220\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr09\u003c/p\u003e\n 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style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e175\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e19787.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e51.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_10G009520\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e10071354-10074351\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n 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style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e25636.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.55\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e66.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e64.56\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGrai_11G000600\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGrChr11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e367221-369228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e371\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41196.57\u003c/p\u003e\n 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style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e77.69\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e60.38\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_A04G001090.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhA04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e1911061-1915067\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e526\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e58504.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e49.54\u003c/p\u003e\n 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18.5567%;\"\u003e\n \u003cp\u003e14259030-14264060\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e534\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e59699.08\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.69\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e47.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e101.14\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_A07G024160.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhA07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e96611166-96613434\u003c/p\u003e\n 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style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e50.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_A10G010120.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhA10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e21088043-21091147\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e128\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e14426.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.74\u003c/p\u003e\n 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style=\"width: 14.433%;\"\u003e\n \u003cp\u003e69.12\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_A11G034980.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhA11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e122694995-122697264\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e371\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41459.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e53.66\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e71.24\u003c/p\u003e\n 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style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_D01G011820.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e20230112-20236251\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e629\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e68363.49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e57.43\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e65.87\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n 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style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e6713089-6715354\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e392\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e43226.83\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.34\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e30.77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.82\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_D05G006570.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD05\u003c/p\u003e\n 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style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e59595\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e7.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e46.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e100.79\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_D08G001640.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD08\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e1352898-1357344\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e198\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e21535.64\u003c/p\u003e\n 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style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e46.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e74.16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_D10G005890.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e5353890-5361763\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e594\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e65312.88\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e10.21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e79.32\u003c/p\u003e\n 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style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e72459161-72460841\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e366\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e40620.92\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e7.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e53.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e70.08\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGhir_D13G023330.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGhD13\u003c/p\u003e\n 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style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e79.53\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e69.86\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_A07G003410.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbA07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e3999121-4005238\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e654\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e72390.39\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e50.98\u003c/p\u003e\n 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style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbA09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e2123646-2126079\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e377\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41339.42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e34.11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e79.58\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_A09G015820.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbA09\u003c/p\u003e\n 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style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e36704.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e7.67\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e60.64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e70.34\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_A11G009500.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbA11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e8445257-8445618\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e77\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e9006.33\u003c/p\u003e\n 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style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e73.45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e66.27\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D01G011760.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD01\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e20308205-20314078\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e872\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e95166.93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e54.57\u003c/p\u003e\n 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style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e25105293-25111109\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e940\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e102404.61\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e56.26\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e58.86\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D05G038870.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD05\u003c/p\u003e\n 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style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e186\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e21300.04\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e8.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e53.72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e76.13\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D07G004470.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD07\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e4716453-4722704\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n 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8.24742%;\"\u003e\n \u003cp\u003e21535.64\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e4.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e48.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e71.87\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D09G000900.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD09\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e2004189-2006026\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e377\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41394.52\u003c/p\u003e\n \u003c/td\u003e\n 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\u003cp\u003e92.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D10G012840.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e21280556-21299560\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e1893\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e213608.29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e51.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e74.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D10G018280.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e49046196-49051900\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e25641.72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.55\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e65.37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e67.54\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D11G036400.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eScaffold3329\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e418050-420228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e373\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e41435.76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e6.74\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e56.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e70.86\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 17.5258%;\"\u003e\n \u003cp\u003eGbar_D13G023380.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11.3402%;\"\u003e\n \u003cp\u003eGbD13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 18.5567%;\"\u003e\n \u003cp\u003e58256028-58259823\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 6.18557%;\"\u003e\n \u003cp\u003e+\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e228\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 8.24742%;\"\u003e\n \u003cp\u003e25668.72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 4.12371%;\"\u003e\n \u003cp\u003e5.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.4639%;\"\u003e\n \u003cp\u003e72.79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14.433%;\"\u003e\n \u003cp\u003e64.56\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003ePhylogenetic Analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo investigate the evolutionary trajectories of TAF family members in cotton, a phylogenetic tree was constructed using the protein sequences of 18 \u003cem\u003eArabidopsis thaliana\u003c/em\u003e TAFs (\u003cem\u003eAtTAFs\u003c/em\u003e) and 124 TAFs derived from 4 cotton species. In the phylogenetic tree (\u003cstrong\u003eFig. 1\u003c/strong\u003e), we noticed that the TAF proteins were clustered into 6 subfamilies (I-VI) with 32 members in subfamily I, 21 in II, 8 in III, 17 in IV, 26 in V, and 38 in VI. Among them, TAF8 contained the largest number of members, whereas TAF2 had the fewest, indicating that members of different TAFs have undergone different evolutionary events involving duplication and deletion.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eChromosome Localization, Collinearity Analysis, and Selection Pressure Analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo visually illustrate the chromosomal distributions of the \u003cem\u003eTAF\u0026nbsp;\u003c/em\u003egene family, localization analyses were conducted for all TAF genes identified in cotton. The results showed that TAFs were distributed across different chromosomes in the four cotton species (\u003cstrong\u003eFig. 2\u003c/strong\u003e). In \u003cem\u003eG. arboreum\u003c/em\u003e, twenty-two \u003cem\u003eGaTAFs\u003c/em\u003e were located on nine chromosomes (Chr01, Chr04, Chr05, Chr06, Chr07, Chr8, Chr9, Chr10 and Chr13). In \u003cem\u003eG. raimondii\u003c/em\u003e, nineteen \u003cem\u003eGrTAFs\u003c/em\u003e were located on seven chromosomes (Chr04, Chr05, Chr06, Chr07, Chr8, Chr9, Chr10, Chr11 and Chr13). In \u003cem\u003eG. hirsutum\u003c/em\u003e, 39 \u003cem\u003eGhTAF\u003c/em\u003e genes were mapped to 19 chromosomes, comprising nine chromosomes from the A\u003csub\u003et\u0026nbsp;\u003c/sub\u003esubgenome and ten chromosomes from the D\u003csub\u003et\u003c/sub\u003e subgenome. In \u003cem\u003eGossypium barbadense\u003c/em\u003e, 44 \u003cem\u003eGbTAFs\u003c/em\u003e were mapped on 20 chromosomes, including ten chromosomes from the A\u003csub\u003et\u003c/sub\u003e subgenome and ten chromosomes from the D\u003csub\u003et\u003c/sub\u003e subgenome.\u003c/p\u003e\n\u003cp\u003eGene duplication events play a pivotal role in plant evolution, among which whole-genome duplication, segmental duplication, and tandem duplication are recognized as the major drivers underlying the expansion of gene families (Cannon, et al. 2004). To detect gene duplication events in the TAF gene family, we analyzed the intra- and interspecific collinearity of TAF genes across the four cotton species. As we all know, \u003cem\u003eGossypium hirsutum\u003c/em\u003e and \u003cem\u003eGossypium barbadense\u003c/em\u003e are allotetraploid species derived from two distinct diploid progenitors, so we generated collinearity plots to compare TAF gene collinearity between diploid and tetraploid cotton species. To identify gene duplication events, we assessed the interspecific collinearity of the 124 previously identified TAF genes across four cotton species, namely \u003cem\u003eGossypium arboreum\u003c/em\u003e, \u003cem\u003eG. hirsutum\u003c/em\u003e, \u003cem\u003eG. barbadense\u003c/em\u003e, and \u003cem\u003eG. raimondii\u003c/em\u003e (\u003cstrong\u003eFig. 3\u003c/strong\u003eA, B). A total of 25 and 26 pairs of gene duplication from Ga-Gh and Ga-Gb comparison, whereas 25 and 21 pairs were found in the Gr-Gh and Gr-Gb comparisons. The results show that there were 25 and 21 collinear TAF genes within the A subgenomes and D sub-genomes, respectively. The intraspecific collinearity of 83 TAF genes was also assessed in \u003cem\u003eG. hirsutum\u003c/em\u003e and \u003cem\u003eG. barbadense\u003c/em\u003e, including collinearity between the A and D subgenomes within each species (\u003cstrong\u003eFig. 3\u003c/strong\u003eC, D). Most TAFs exhibit good collinearity within the A\u003csub\u003et\u003c/sub\u003e-genome or D\u003csub\u003et\u003c/sub\u003e-genome of the two tetraploid cotton species.\u003c/p\u003e\n\u003cp\u003eTo estimate the correlation of repeating genes over along evolutionary history, Ka/Ks values in \u003cem\u003eGhTAF\u003c/em\u003e gene homologous pairs were calculated based on different selection pressures such as purification, neutral, and active selection. According to the Ka/Ks analysis (Table S1), the Ka/Ks values were below 1.0, indicating that these \u003cem\u003eGhTAF\u003c/em\u003e genes underwent strong purification selection during evolution. A total of 48 gene pairs exhibited a Ka/Ks ratio of less than 0.5, and 7 gene pairs were between 0.5 and 1, leading us to hypothesize that these TAF gene pairs have undergone strong purifying selection during evolution.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStructure Analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eStructural difference among genes is a key factor shaping the evolution of gene families. To better understand the structural characteristics of TAF genes in \u003cem\u003eG. hirsutum\u003c/em\u003e, we analyzed the gene structure, motif, and conserved domains of TAF genes in this species, with the analysis conducted based on the constructed phylogenetic tree (\u003cstrong\u003eFig. 4\u003c/strong\u003e). Online MEME analysis was performed to identify 20 conserved motifs among the 39 \u003cem\u003eGhTAF\u003c/em\u003e genes. Most members exhibit motif 6, but different types of \u003cem\u003eGhTAF\u003c/em\u003e motifs exhibit distinct distributions. One plausible explanation is that many TAFs appear to have evolved independently. The gene structure of \u003cem\u003eGhTAF\u003c/em\u003e showed that the number of exons and introns shows considerable variation across different types of genes, and a few TAFs only contained a single exon. These results are consistent with the clustering pattern of the phylogenetic tree.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAnalysis of GhTAFs Gene Expression in Different Tissues of upland cotton\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAlthough the sequence structure was thoroughly investigated, their potential roles in ovule and fiber development remained unclear. In this study, we integrated published transcriptome data comprising 20 samples (T111 and T113 across five developmental stages). Transcriptome data for fiber development were obtained from previous studies, whereas data for ovule development are reported here for the first time (Table S2). T111 shows high oil content and low lint percentage, whereas T113 displays the opposite traits: low oil content and high lint percentage. The spatiotemporal expression patterns of genes are strongly correlated with their biological functions. Therefore, characterizing the transcription landscape of \u003cem\u003eGhTAFs\u003c/em\u003e is essential for illustrating their roles in ovule and fiber development. Our analysis revealed distinct, tissue-specific transcription patterns for several \u003cem\u003eGhTAFs\u003c/em\u003e. According to the results of expression clustering, the \u003cem\u003eGhTAF\u003c/em\u003e genes were categorized into 5 clusters (I-V) in ovule (\u003cstrong\u003eFig. 5\u003c/strong\u003e). Among these, cluster I and IV showed almost no expression, while cluster II, III, and V exhibited high expression levels. In cluster I,\u003cem\u003e\u0026nbsp;\u003c/em\u003ethe expression levels of \u003cem\u003eGhir_A09G015600\u003c/em\u003e and \u003cem\u003eGhir_D09G015070\u003c/em\u003e were undetectable in terms of expression across all experimental periods in T111 and T113, whereas\u003cem\u003e\u0026nbsp;Ghir_D10G009150\u003c/em\u003e displayed significantly higher expression level in ovules from line T113 relative to line T111. In cluster IV, most \u003cem\u003eGhTAF\u003c/em\u003e genes exhibited two key expression patterns: first, their expression levels were higher in T111 than in T113; second, their expression at 10, 15 and 20 DPA was higher than that at 25 and 30 DPA. \u003cem\u003eGhir_A07G003870\u003c/em\u003e, \u003cem\u003eGhir_A01G012900\u003c/em\u003e, and \u003cem\u003eGhir_D10G005890\u003c/em\u003e were typical examples of these patterns.\u003c/p\u003e\n\u003cp\u003eMeanwhile, \u003cem\u003eGhTAFs\u003c/em\u003e were clustered into 2 clusters in fiber. Specifically, genes in cluster II exhibited high expression level, while those in cluster I showed low expression level. In cluster II, \u003cem\u003eGhir_D10G009150\u003c/em\u003e showed significantly higher expression level in the fibers of line T113 than in those of line T111. Nearly half of the \u003cem\u003eGhTAFs\u003c/em\u003e, such as \u003cem\u003eGhir_A05G006100\u003c/em\u003e, \u003cem\u003eGhir_A10G010120\u003c/em\u003e and \u003cem\u003eGhir_A07G024160\u003c/em\u003e, exhibited stage-specific expression patterns during ovule and fiber development. In addition, several \u003cem\u003eGhTAF\u003c/em\u003e genes exhibited high expression levels in both ovules and fibers; representative examples include \u003cem\u003eGhir_A10G005150\u003c/em\u003e, \u003cem\u003eGhir_D07G004430\u003c/em\u003e, and \u003cem\u003eGhir_D10G005930\u003c/em\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eWeighted gene co-expression network analysis (WGCNA)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor further investigation of the potential association between \u003cem\u003eGhTAFs\u003c/em\u003e and ovule and fiber development, we employed weighted gene co-expression network analysis on the set of retained genes (\u003cstrong\u003eFig. 6\u003c/strong\u003e). The associations between gene regulatory modules (GRMs) and phenotypes traits are multifaceted. In ovule, Purple, turquoise, cyan and green GRMs were highly correlated to ovule at 10 DPA. Brown, magenta and red GRMs were highly correlated to ovule at 15 DPA. Yellow, pink, blue, green and midnightblue GRMs were correlated to ovule at 20 DPA. Meanwhile, cyan and blue GRMs were correlated to ovule at 25 DPA. Black, red, blue and turquoise GRMs were correlated to ovule at 30 DPA (Fig. 6B). Among the retained 28 \u003cem\u003eGhTAFs\u003c/em\u003e, we found that these 28 genes were mainly from three GRMs, namely the turquoise, blue and red GRMs, which were involved in ovule development at different stages. A total of 16 \u003cem\u003eGhTAFs\u003c/em\u003e were identified to be involved in the turquoise GRM, which is primarily associated with the 10 DPA stage. Similarly, 2 \u003cem\u003eGhTAFs\u003c/em\u003e were involved in the red GRM (predominantly associated with 15 DPA). And 9 \u003cem\u003eGhTAFs\u003c/em\u003e participated in the blue GRM, which is primarily associated with 20 DPA developmental stage. 20 DPA is a critical developmental stage during which cottonseed oil accumulates rapidly (Song, et al. 2022). Among the 9 \u003cem\u003eGhTAFs\u003c/em\u003e, \u003cem\u003eGhir_D11G035840\u003c/em\u003e exhibits the strongest correlation with the blue GRM. WGCNA results served as a platform to establish a putative interaction network for \u003cem\u003eGhir_D11G035840\u003c/em\u003e. As \u003cem\u003eGhir_D11G035840\u003c/em\u003e is a member of the blue module, we constructed its interaction network with other blue module-associated genes. A total of 133 genes from the blue GRM, which exhibited a Pearson\u0026rsquo;s correlation coefficient of 0.95 with \u003cem\u003eGhir_D11G035840\u003c/em\u003e, were identified as candidate interacting genes, including \u003cem\u003eVRN1\u003c/em\u003e and \u003cem\u003eMYB3R4\u003c/em\u003e (\u003cstrong\u003eFig. 6\u003c/strong\u003eC and Table S3). \u003cem\u003eVRN1\u003c/em\u003e is a member of the B3 domain transcription factor family. Among this family, \u003cem\u003eABI3\u003c/em\u003e, \u003cem\u003eFUS3\u003c/em\u003e, \u003cem\u003eLEC2\u003c/em\u003e, etc. are core members that regu-late seed development and lipid synthesis (Moreno 2024; Yang, et al. 2022). They directly activate the expression of key genes involved in lipid biosynthesis (e.g., \u003cem\u003eDGAT\u003c/em\u003e, \u003cem\u003ePDAT\u003c/em\u003e) as well as genes encoding seed storage proteins. Although \u003cem\u003eVRN1\u003c/em\u003e, a transcription factor harboring two B3 domains, is primarily associated with vernalization, accumulating evidence supports its definitive role in seed development(Li, et al. 2019; Milec, et al. 2022). The Myb-domain transcription factor \u003cem\u003eMYB3R4\u003c/em\u003e indirectly ensures the normal development of seeds by regulating cell cycle-related genes, indirectly but significantly affects seed size and final yield (Haga, et al. 2011). Additionally, \u003cem\u003eMYB3R4\u003c/em\u003e can directly activate the expression of the cellulose synthase-like gene \u003cem\u003eCSLD5\u003c/em\u003e, thereby facilitating the rapid and efficient assembly of new cell walls at the end of mitosis and providing a structural and cellular foundation for seed development (Lan, et al. 2025). Therefore, \u003cem\u003eGhir_D11G035840\u003c/em\u003e is likely involved in cottonseed oil accumulation.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eGhir_D11G035840\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;may positively regulates oil content\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo investigate the in vivo subcellular distribution of \u003cem\u003eGhir_D11G035840\u003c/em\u003e, we per-formed subcellular localization assays on tobacco leaves. The results demonstrated that \u003cem\u003eGhir_D11G035840\u003c/em\u003e is localized to the nucleus (\u003cstrong\u003eFig. 7\u003c/strong\u003eA). To evaluate the impact of \u003cem\u003eGhir_D11G035840\u003c/em\u003e on vegetable oil accumulation and ovule development, we produced \u003cem\u003eGhir_D11G035840\u003c/em\u003e-overexpressing (OE) lines in A. thaliana through Agrobacterium-mediated floral dip transformation with strain GV3101. Subsequently, 8 independent transgenic \u003cem\u003eArabidopsis thaliana\u003c/em\u003e plants were obtained, and their transgenic status was verified by molecular identification and Western blot analysis (\u003cstrong\u003eFig. 7\u003c/strong\u003eB-D). Two distinct independent OE lines of \u003cem\u003eGhir_D11G035840\u003c/em\u003e in A. thaliana were selected from these transformants for phenotypic evaluation. Compared with WT (ecotype Columbia-0), the seed oil content of OE was significantly increased (\u003cstrong\u003eFig. 7\u003c/strong\u003eE). Furthermore, the levels of specific fatty acid components, including C18:1 (oleic acid) and C18:2 (linoleic acid), were significantly elevated in the seeds of overexpression lines (\u003cstrong\u003eFig. 7\u003c/strong\u003eF). In contrast, no significant difference in thousand-seed weight was detected between the OE lines and WT (\u003cstrong\u003eFig. 7\u003c/strong\u003eG). These results suggest that \u003cem\u003eGhir_D11G035840\u003c/em\u003e may positively regulate seed oil content in plants.\u003c/p\u003e\n\u003cp\u003eRoots have been utilized as an alternative model to study cell elongation in prior cotton genomics studies, owing to their relevance to the cell elongation process shared with fibers. After vernalization and surface sterilization, seeds from both OE lines and WT were plated on 1/2 MS medium and incubated at 22\u0026deg;C with a 16 h light/8 h dark photoperiod for root development monitoring. Following 7 days of cultivation, root length measurements and comparative analysis revealed that OE lines exhibited a no-table reduction in root length compared with the WT control (\u003cstrong\u003eFig. 7\u003c/strong\u003eH, I). Root length of the OE lines proved that the expression of \u003cem\u003eGhir_D11G035840\u003c/em\u003e could repress the fiber elongation.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eNowadays, except for in-depth research on the TAF gene family of \u003cem\u003eArabidopsis\u0026nbsp;\u003c/em\u003e(Lago, et al. 2004) and mungbean (Wu, et al. 2024), research on other plants is relatively shallow. TAFs, as components of the general transcription factor TFIID, with highly conserved sequences from yeast to humans (Hoffman, et al. 1990). Therefore, utilizing bioinformatics tools, such as BLAST, to identify potential homologous proteins is reliable. In our initial screening of the cotton genome, we identified 124 TAFs, distributed across the four cotton species as follows: including 39 genes in G\u003cem\u003e. hirsutum\u003c/em\u003e, 44 genes in \u003cem\u003eG. barbadense\u003c/em\u003e, 22 in \u003cem\u003eG. arboreum\u003c/em\u003e, and 19 in \u003cem\u003eG. raimondii\u003c/em\u003e. This distribution suggests that \u003cem\u003eTAF\u003c/em\u003e gene family is relatively conserved during cotton genome evolution. These TAF proteins were clustered based on the phylogenetic analysis into six groups (\u003cstrong\u003eFig. 1\u003c/strong\u003e). Typically, the TFIID complex is composed of 13 to 14 distinct TAF subunits. In \u003cem\u003eArabidopsis thaliana\u003c/em\u003e, 18 putative \u003cem\u003eAtTAF\u003c/em\u003e proteins were identified\u0026nbsp;(Lago, et al. 2004). TAF proteins are grouped according to their common biochemical feature of being associated in stable complexes with the TATA-binding protein (TBP). So, in order to proof that a putative TAF is a TATA-box binding protein associated factor, a biochemical demonstration of its participation in the TFIID complex is needed. There is currently no exactly biochemical evidence in \u003cem\u003eArabidopsis\u003c/em\u003e demonstrating that \u003cem\u003eAtTAF14\u003c/em\u003e and \u003cem\u003eAtTAF15\u003c/em\u003e are TBP-associated factors, these two proteins are not considered TATA-box binding protein associated factor\u0026nbsp;(Lago, et al. 2004). Furthermore, the \u003cem\u003eArabidopsis\u003c/em\u003e genome lacked an identifiable homolog of TAF3, but now the TAIR database designates protein \u003cem\u003eAt5g15570\u003c/em\u003e as TAF3. It is noteworthy that, \u003cem\u003eAtTAF3\u003c/em\u003e and \u003cem\u003eGhTAF3\u003c/em\u003e possesses the BTP and TAF8 domain, instead of TAF3 histone fold domain. Meanwhile, previous researches have proved that there is no homologous TAF3 in the mungbean genome\u0026nbsp;(Wu, et al. 2024)\u0026nbsp;and rice genome\u0026nbsp;(Lago, et al. 2004). In yeast, mutation of TAF3 results in a thermo-sensitive phenotype which can be suppressed by overexpression of TAF10, TAF11, TAF13 or TAF6\u0026nbsp;(Gangloff, et al. 2001). Therefore, it is hypothesized that TAF3 is absent in plants, while the functional role of TAF3 may be compensated for by other TAF subunits.\u003c/p\u003e\n\u003cp\u003eGene duplication event is a common phenomenon in plants. Some duplicated genes could be retained in in descendant lineages, which could serve as raw genetic material for adaptive evolution of plants\u0026nbsp;(Flagel and Wendel 2009). The analysis of chromosomal location revealed that TAF family genes from four cotton species were unevenly distributed on their respective chromosomes. In this study, the possible duplication events in TAF gene family were investigated in four cotton species. According to the selection pressure analysis, the Ka/Ks ratio of \u003cem\u003eG. hirsutum\u003c/em\u003e and \u003cem\u003eG. barbadense\u003c/em\u003e was less than 1 (Table S1), supporting the evolutionary conservation of these genes. The results showed that most TAF genes are slowly evolving. Gene structure analysis showed that exon number of \u003cem\u003eGhTAF\u003c/em\u003e genes varied greatly, which might be attributed to the directional evolution of TAF genes in terms of both function and structure over the long course of evolutionary history. Meanwhile the same type of TAFs have similar motifs, structural domains and number of exons, which indicates that the same type of TAFs may be functionally conserved during evolution.\u003c/p\u003e\n\u003cp\u003eCotton is an economically vital crop that serves as the primary source of natural textile fibers as well as a significant oilseed crop providing edible oil for human consumption. A large number of genes and gene families have been extensively studied in \u003cem\u003eG. hirsutum\u003c/em\u003e. Consequently, \u003cem\u003eG. hirsutum\u003c/em\u003e was used for gene expression analysis in the present study. Previous studies have demonstrated that TAFs are widely involved in multiple processes of plant development. For example, \u003cem\u003eAtTAF5\u0026nbsp;\u003c/em\u003e(Mougiou, et al. 2011) and \u003cem\u003eAtTAF6\u0026nbsp;\u003c/em\u003e(Lago, et al. 2005) regulates pollen tube growth, \u003cem\u003eAtTAF13\u0026nbsp;\u003c/em\u003e(Lindner, et al. 2013)[13] participate in seed development, and \u003cem\u003eOsTAF2\u0026nbsp;\u003c/em\u003e(Jiang, et al. 2023)\u0026nbsp;positively regulates the grain size by modulating several cell cycle related genes. The pattern of gene expression is directly related to its function. Based on transcriptome data, we analyzed the differential expression of \u003cem\u003eGhTAFs\u003c/em\u003e in fiber and ovule (\u003cstrong\u003eFig. 5\u003c/strong\u003e). The expression pattern shows that most half of \u003cem\u003eGhTAFs\u003c/em\u003e had stage-specific transcription patterns in ovule and fiber, such as \u003cem\u003eGhir_A05G006100\u003c/em\u003e, \u003cem\u003eGhir_A10G010120\u003c/em\u003e, and \u003cem\u003eGhir_A07G024160\u003c/em\u003e. Several \u003cem\u003eGhTAF\u003c/em\u003e genes were found to exhibit high expression levels in both ovules and fibers, with \u003cem\u003eGhir_A10G005150\u003c/em\u003e, \u003cem\u003eGhir_D07G004430\u003c/em\u003e, and \u003cem\u003eGhir_D10G005930\u003c/em\u003e identified as representative examples. Furthermore, the expression level of \u003cem\u003eGhir_D10G009150\u003c/em\u003e exhibits significant differences between the two lines, T111 and T113.\u003c/p\u003e\n\u003cp\u003eWeighted Gene Co-expression Network Analysis (WGCNA) is a classic systems biology approach that enables the identification of modules of co-expressed genes (Langfelder and Horvath 2008). To systematically identify the transcription factors that play a crucial role in the development of cotton fibers, we performed WGCNA of the TAF family throughout the entire development periods of ovules and fibers, respectively. We found that \u003cem\u003eGhTAF\u003c/em\u003e genes participate in the turquoise, blue, and red co-expression modules in the ovule, with these modules linked to the 10, 15, and 20 DPA stages of ovule development. During cotton ovule development, this period falls in the middle stage and is characterized by rapid oil accumulation. At this stage, enzymes such as glycerol-3-phosphoacyltransferase (GPAT) and lysophosphatidylate-acyltransferase (LPAAT) play a crucial role\u0026nbsp;(Cui, et al. 2019; Wang, et al. 2017). They can successively attach fatty acids to glycerol molecules, gradually constructing the molecular structure of triglycerides. During this process, various types of fatty acids are combined in specific proportions and sequences, thereby influencing the fatty acid composition and overall quality of cottonseed oil\u0026nbsp;(Yuan, et al. 2018). At the same time, this stage also corresponds to the early phase of seed size determination. Thus, \u003cem\u003eGhTAFs\u003c/em\u003e are hypothesized to regulate plant seed size via several pathways, such as the ubiquitin regulatory pathway, plant hormone pathways (auxin and brassinolide), MAPK signaling pathway, G protein signaling pathway, transcription factor pathway, and IKU signaling pathway\u0026nbsp;(Li and Li 2016).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAdditionally, we generated transgenic \u003cem\u003eArabidopsis thaliana\u003c/em\u003e lines with overexpression of the target gene \u003cem\u003eGhir_D11G035840\u003c/em\u003e, followed by phenotypic characterization of seeds. These observations were performed to infer the potential function of the candidate gene in ovule development. Through the integration of transcriptome profiling data and phenotypic analysis of \u003cem\u003eArabidopsis thaliana\u003c/em\u003e, we validated the functional role of the gene \u003cem\u003eGhir_D11G035840\u003c/em\u003e in ovule development and oil accumulation. Furthermore, we constructed a putative interaction network for \u003cem\u003eGhir_D11G035840\u003c/em\u003e by leveraging transcriptional abundance information. Notably, B3 domain-containing transcription factors function as central regulators within the regulatory circuitry orchestrating seed development and lipid biosynthesis in this interaction network. Furthermore, \u003cem\u003eMYB3R4\u003c/em\u003e indirectly but markedly impacts seed size and final crop yield by transcriptionally regulating genes involved in cell cycle progression (Haga, et al. 2011). Wei et al. reported that TAF in Drosophila melanogaster may regulate the fatty acid composition of multiple phospholipid classes by modulating the transcription of genes associated with peroxisomal fatty acid β-oxidation (Fan, et al. 2017). Collectively, these characterizations demonstrate the potential roles of TAFs in ovule development and oil accumulation.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn this study, we identified 124 members of the TAF gene family across four cotton species, which were classified into six subgroups based on phylogenetic analysis. We conducted gene expression analysis, identifying key \u003cem\u003eGhTAF\u003c/em\u003e genes that are specifically or persistently highly expressed at multiple cottonseed and fiber developmental stages. Furthermore, based on WGCNA data, we identified \u003cem\u003eGhir_D11G035840\u003c/em\u003e as a TAF family gene potentially involved in regulating lipid accumulation and ovule development. Further overexpression experiments on \u003cem\u003eArabidopsis thaliana\u003c/em\u003e indicated that \u003cem\u003eGhir_D11G035840\u003c/em\u003e may positively regulate oil content. These findings will also provide valuable genetic resources and theoretical foundations for future studies on the structure and function of the\u003cem\u003e\u0026nbsp;GhTAF\u003c/em\u003e gene family.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAuthor Contributions\u003c/p\u003e\n\u003cp\u003eYL and JS designed the experiments; JG and LW performed the experiments and collected the data. JG, JS, JZ, PF, BJ, SC, WZ, BZ, JM, WP, MW and KZ analyzed the data. JG wrote the manuscript. JS, LY, YJ, WL and JG revised the manuscript. All authors have read and agreed to the published version of the manuscript.\u003c/p\u003e\n\u003cp\u003eFunding\u003c/p\u003e\n\u003cp\u003eThis work was supported by The Key Research and Development Program of Xinjiang (Grant No. 2024B02001-1), the National Natural Science Foundation of China (Grant No. 32501903), the Natural Science Foundation of Henan Province (Grant No. 252300420740), and the Natural Science Foundation of Xinjiang Uygur Autonomous Region (Grant No. 2024D01A132).\u003c/p\u003e\n\u003cp\u003eData Availability Statement\u003c/p\u003e\n\u003cp\u003eAll data generated or analyzed during this study are included in this published article and its additional files. The datasets used and analyzed in the current study are available from the corresponding author upon reasonable request.\u0026nbsp;All the transcriptome raw data we sequenced was deposited in the NCBI short read archives (SRA; accession number PRJNA1378545).\u003c/p\u003e\n\u003cp\u003eConflicts of Interest\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAlbright SR, Tjian R (2000) TAFs revisited: more data reveal new twists and confirm old ideas. Gene 242:1-13\u003c/li\u003e\n \u003cli\u003eAoyagi N, Wassarman DA (2001) Developmental and transcriptional consequences of mutations in Drosophila TAF(II)60. Mol Cell Biol 21:6808-6819\u003c/li\u003e\n \u003cli\u003eBertrand C, Benhamed M, Li YF et al (2005) Arabidopsis HAF2 gene encoding TATA-binding protein (TBP)-associated factor TAF1, is required to integrate light signals to regulate gene expression and growth. 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Nat Biotechnol 33:531-537\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":true,"hideJournal":true,"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":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"cotton, TAFs, genome-wide, expression pattern, WGCNA, Ghir_D11G035840","lastPublishedDoi":"10.21203/rs.3.rs-8328337/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8328337/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eTBP-associated factors (TAFs), along with the TATA-box binding protein (TBP), com-pose transcription factor IID (TFIID), which is an essential complex for transcription initiation and regulation. TAFs are involved in regulating plant development. However, genome-wide identification and expression pattern analysis of TAFs in cotton remain unreported. In this study, a total of 124 TAF genes were identified in four \u003cem\u003eGossypium \u003c/em\u003especies. TAFs were divided into six subgroups based on the phylogenetic tree analysis. Analysis of collinearity and selection pressure indicated that TAF gene pairs have undergone strong purifying selection during evolution. Gene expression analysis of \u003cem\u003eGhTAFs\u003c/em\u003e revealed that TAFs are expressed at all stages of cotton seed and fiber development. Furthermore, a set of key \u003cem\u003eGhTAF\u003c/em\u003e genes was identified, which are specifically or constitutively highly expressed across multiple stages of cotton seed and fiber development. Based on WGCNA, we identified \u003cem\u003eGhir_D11G035840 \u003c/em\u003eas a candidate gene involved in lipid accumulation regulation, and further constructed a putative co-expression interaction network for this gene to elucidate its functional connections. Overexpression of \u003cem\u003eGhir_D11G035840\u003c/em\u003e in \u003cem\u003eArabidopsis thaliana\u003c/em\u003e resulted in a significant increase in seed oil content, thereby confirming its functional role in regulating oil accumulation. Furthermore, a putative interaction network derived from the co-expression network revealed that \u003cem\u003eGhir_D11G035840\u003c/em\u003e may interact with numerous genes to regulate both ovule development and lipid synthesis. These results provide valuable genetic resources and a theoretical basis for subsequent investigations into the molecular mechanisms underlying cotton seed oil synthesis and fiber quality formation using functional genomics approaches.\u003c/p\u003e","manuscriptTitle":"Genome-wide identification reveals the regulatory roles of the TATA-box binding protein associated factor (TAF) gene family in cotton oil accumulation","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-17 11:13:01","doi":"10.21203/rs.3.rs-8328337/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"9bdb2261-84e6-4cae-abbd-ac73317c7be6","owner":[],"postedDate":"December 17th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-03-24T01:54:46+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-17 11:13:01","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8328337","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8328337","identity":"rs-8328337","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}