Identification, evolution and expression analysis of the UDP-glycosyltransferase gene family in grape (Vitis vinifera L.)

doi:10.21203/rs.3.rs-3892939/v1

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Identification, evolution and expression analysis of the UDP-glycosyltransferase gene family in grape (Vitis vinifera L.)

https://doi.org/10.21203/rs.3.rs-3892939/v1

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Uridine diphosphate glycos-yltransferases (UGTs) are involved in the process of glycosylation modification and play an important role in plant growth and development, secondary metabolism and stress responses. Here, a systematic genome-wide analysis of UGT family was conducted to investigate their important functions in grape. In total, 181 VvUGTs were identified and unevenly distributed on 18 chromosomes. These UGT genes were clustered into 17 major groups (A-P, R) based on phylogenetic analysis, supported by conserved gene organization and consensus motifs. Cis-acting elements analysis of the VvUGT genes indicated their potential roles in plant development and responses to various stresses. Collinearity analysis showed that segmental duplication and tandem duplication events had significant effects on the evolution of VvUGT gene family. In addition, genes related to secondary metabolic processes were screened by KEGG and GO enrichment analysis. Expression profiles displayed distinct expression patterns of VvUGT genes in different tissues, fruits development stages, as well as in response to various stress. Overall, these findings lay an important foundation for further functional dissection of VvUGTs and genetic improvement of V. vinifera.

Vitis vinifera

UGT genes

expression profile analysis

Collinear analysis

Glycosylation is an important modification in all living organisms. Glycosyltransferases (GTs) are responsible for modification of glycosylation in various metabolic processes, and ubiquitous present in all organisms. They can transfer sugar moieties from activated donor molecules to specific receptor molecules. According to the most recent update of CAZy (http://www.cazy.org), GTs from various species can be divided into 115 subfamilies (Chen et al. 2022), many of which have been identified in different species (Yin et al. 2010; Kim et al. 2012; Lee et al. 2014). Among them, family-1 glycosyltransferases, also known as UDP glycosyltransferases (UGTs), are the most common glycosyltransferase in plants (Yonekura-Sakakibara and Hanada 2011). UGTs are capable of catalyzing transfer of glycosyl group to specific receptors in plants, including phytohormone, secondary metabolites, and xenobiotics (Caputi et al. 2012). UGTs are often characterized by a conserved motif of 44 amino acids at the C-terminus, called the Plant Secondary Product Glycosyltransferase (PSPG) box (Zhang et al. 2020), which functions for the binding of glycosyl groups. Diametrically, the N-terminal sequence varies considerably and is responsible for recognition of different substrates (Vogt and Jones 2000). To date, with the completion of plant genome sequencing, UGTs have been widely investigated in various plants. UGT has been found to exist as multigene families and expand in many plants. In Arabidopsis, more than 100 UGT genes have been found (Li et al. 2001). Even more UGT members have been identified in other plant species, such as Zea mays (147) (Li et al. 2014), Triticum aestivum (179) (He et al. 2018), Citrus grandis (145) (Wu et al. 2020), Malus x domestica (214) (Caputi et al. 2012), and Prunus persica (168) (Wu et al. 2017).

UGTs play a crucial role in various life activities of plants. Many studies have showed that UGTs are involved in regulation of secondary metabolism and hormone homeostasis in plants. For instance, UGT79B1 and UGT91A1 from Arabidopsis, peach, strawberry, and kiwifruit were found to mediate anthocyanin modification (Montefiori et al. 2011; Yonekura-Sakakibara et al. 2012; Song et al. 2016). In citrus, Cm1_2RhaT, CsUGT76F1, and Cs1,6RhaT were reported to catalyze flavonoid glycosylation (Frydman et al. 2004; Liu et al. 2018; Wu et al. 2020). In Arabidopsis, UGT74E2 gene was shown to participate in plant morphogenesis and stress responses through glycosylation of indole-3-butyric acid (IBA) (Tognetti et al. 2010). In tomato, suppression of SlUGT75C1 significantly accelerated fruit ripening by increasing ABA levels and promoting the early release of ethylene (Sun et al. 2017). Both UGT84B1 and UGT74D1 regulated IAA levels during plant development through dual IAA and oxIAA glycosylation (Mateo‐Bonmatí et al. 2021). The functions of plant UGTs have also been associated with stress responses. In Arabidopsis, overexpression of UGT75B1 gene improved seed germination and seedling greening rate under salt and osmotic stress (Chen et al. 2020). In rice, OsUGT3 was identified and found to act a critical role in drought and salt stress through modulating ABA synthesis and scavenging ROS (Wang et al. 2021). UGT76B1 served as a negative regulator of SA-dependent plant defense in the absence of pathogens, enhanced the JA response, and delayed the onset of senescence (von Saint Paul et al. 2011).

Grapevine (Vitis vinifera L.) is a highly significant fruit crops around the world and the fruit contain special flavors and rich nutrients (Del-Castillo‐Alonso et al. 2021; Rienth et al. 2021). It can be consumed fresh or dried, or can be processed into various products, such as jam, juice, wine, vinegar, and grape seed oil. In this study, we conducted a wide-ranging bioinformatics analysis of UGT gene family in grapevine and explored the potential functions of VvUGTs. In total, 181 members were identified based on grape genome. Phylogenetic analysis, chromosome localization, gene structure, motif and promoter analysis, enrichment analysis, and collinearity analysis were carried out. Expression patterns of VvUGT genes in various tissues and fruits at different development stages were examined. Moreover, stimulus-induced expression patterns of VvUGT genes were also investigated under drought and salt treatment. This work will greatly facilitate further function characterization of UGT genes in grapevine.

Identification of VvUGT genes in grapes

The grape protein sequences were downloaded from the Phytozome V13 (https://phtozome-next.jgi.doe.gov/). The sequences of Arabidopsis UGT were obtained from TAIR (https://www.arabidopsis.org/). In this study, two methods were used for identifying VvUGT genes. First, a BLASTP search was conducted in a local protein database of grape using Arabidopsis UGT sequences as queries. Second, the hidden Markov model (HMM) profile of the UGT domain (PF00201) was retrieved from Pfam (http://pfam.xfam.org/). The putative VvUGT genes were identified from the grape protein data using HMM file and hmmsearch software with an E-value threshold of 1e-5. Candidate members yielded by the two methods were screened using SMART to remove proteins without a complete PSPG motif. Basic characteristics of VvUGTs were also analyzed using the ExPASy Online program (http://web.expasy.org/protparam/).

Phylogenetic, gene structure and conserved motif analysis

The protein sequences of UGTs from grape, Arabidopsis, tomato, mouse-ear hawkweed and maize were aligned with MUSCLE, and a neighbor-joining (NJ) phylogenetic tree was constructed by MEGA 11 with 1000 bootstrap replicates. Gene structures of VvUGTs were mapped by TBtools software based on the intron-exon data in the grape GFF3 annotation file (Chen et al. 2020a). The conserved motifs of VvUGT proteins were identified using MEME (https://meme-suite.org/meme/tools/meme) with a maximum of 10 motifs per sequence.

Chromosomal locations and collinearity analysis of VvUGTs

A physical chromosomal map of VvUGTs was performed by TBtools according to positional information from grape genome. The tandem repeat and segmental duplication events of UGT genes in grape genome were searched using multiple collinear scanning toolkits (MCScanX) with default parameters (Wang et al. 2012).The syntenic relationships UGTs between grape and A. thaliana were also performed using MCScanX.

Detection of cis-regulatory elements and enrichment analysis

The upstream 2000 bp sequences from the start codon of each VvUGT were extracted from grape genome. Then cis-regulatory elements in the promoter regions of VvUGTs were predicted using the PlantCARE (http://bioinformatics.psb.ugent.be/webtools/plantcare/html). GO and KEGG enrichment analysis were performed using TBtools among VvUGTs.

Expression profile analysis of VvUGT genes

Expression data of VvUGT genes in different tissues and under abiotic stress were obtained from NCBI (GSE36128 and GES31677). Transcriptome data including two grape cultivars, namely, Muscat Blanc (a white variety) and Muscat Rouge (a red variety), was used to investigate the expression profiles of VvUGT genes at different development stages of grape fruit. Each grape cultivar had three fruit developmental stages: young fruit stage, veraison, and maturity. Fragments Per kb per Million reads (FPKM) values were calculated to evaluate the gene expression. The obtained FPKM values of VvUGT genes were log2 transformed, and visualized as a heatmap using TBtools software.

Identification of UGT gene family in grape

A total of 181 UGT genes were identified in the grape genome (Table 1). These UGTs were named VvUGT1 to VvUGT181 according to their chromosomal locations. They were unevenly distributed on 18 grape chromosomes except for 10 UGT members (Fig. 1). Among them, chromosome 5 contained the most UGT genes (26 members), followed chromosome 18 (23 members), and chromosome 12 (19 members). Only one member was mapped to chromosome 9. Notably, many clusters were observed on 18 chromosomes, such as Chr3, Chr5, Chr6, and Chr17 (Fig. 1). The physicochemical properties analysis showed that the length of VvUGTs ranged from 380 aa (VvUGT74) to 1397 aa (VvUGT181), with an average of approximately 486 aa. The predicted molecular weight of UGTs ranged from 42.60 kDa (VvUGT74) to 156.47 kDa (VvUGT181), averaging 54.07 kDa. Isoelectric point (PI) values were between 4.84 (VvUGT144) and 7.36 (VvUGT77), averaging 5.76. The instability index of most VvUGTs were found to be more than 40 and regarded as unstable proteins. Grand Average of Hydropathicity (GRAVY) scores of more than three quarters of VvUGT proteins were negative, suggesting their hydrophilicity (Table 1).

Analysis of phylogenetic, gene structure and conserved motif

To investigate the evolutionary relationship of the VvUGT gene family, a phylogenetic tree was constructed based on the UGT protein sequences from grape, Arabidopsis, tomato, mouse-ear hawkweed and maize. The phylogenetic tree revealed that 181 VvUGTs can be divided into 17 groups (A-P, R) (Fig. 2). Group E contained the largest VvUGT members (56), followed by group P (24). Group O was the smallest (only one VvUGT member). The group Q was observed only in maize, indicating that it may be monocotyledon-specific group.

Analysis of gene structure is helpful to understand the evolution of gene families. Thus, we analyzed genomic makeup of each VvUGT member (Fig. 3). There were 101 VvUGT genes with no introns and 80 with at least one intron. Of the 80 intron-containing VvUGT genes, 70 contained one intron, 6 had two introns, and the remaining genes possessed between three and five introns. Combined with the phylogenetic tree, most VvUGT genes within the same group possessed a comparable gene structure. For instance, there were no introns in R group, and almost members of group L contained one intron.

The distribution of conserved motifs of VvUGTs was performed by MEME online software (Fig. 3). A total of 20 conserved motifs were detected. Motif 1, 2, 3, 4, 5, 6, 7, 8, 10, 12 and 13 existed in most VvUGT proteins, but some motifs only were found in specific groups. For example, motif 20 was specific to group P and H. Motif 11, 14 and 18 only existed in group E. Additionally, motifs 5 and 15 located at the beginning and motif 9 was in the end of almost VvUGT sequences. In general, the same group shared analogous motifs, and this supported evolutionary classification of VvUGTs above.

Cis-regulatory elements in grape UGT gene promoters

To investigate the promoter features, 2000 bp upstream of VvUGT genes was analyzed (Fig. 4). Many kinds of cis-elements were identified and mainly divided into four categories, including light response, phytohormone response, stress response, and development (Fig. 4). All VvUGT gene promoters were found to contained the light responsive elements, such as Box 4, G-box, GT-motif, and I-box. This implied that the transcription regulation of VvUGT genes may influenced by light signals. The hormone-responsive elements comprised 11 types of elements, in which ABRE (ABA responsiveness), CGTCA-motif (MeJA-responsiveness), and TCA-element (SA responsiveness) were the most abundant, accounting for 25.9%, 22.5%, and 12.6%, respectively. Besides, stress-responsive and development-related elements were identified, such as low-temperature responsiveness (LTR), drought inducibility responsiveness (MBS), CAT-boxes, and circadian elements. The findings indicated that the VvUGT genes were closely associated with plant development and stress responses.

Interspecific and intraspecific collinearity analysis

In order to explore the expansion and evolution mechanisms of VvUGT gene family, gene duplication events in grape genome were analyzed (Fig. 5). In this study, MCScanX software was used to compare grape genome based on amino acid sequence homology. A total of 15 segmental-duplication gene pairs were identified, including VvUGT85/93, VvUGT85/98, VvUGT149/8, VvUGT1/41, VvUGT89/99, VvUGT191/3, VvUGT117/141, VvUGT65/69, VvUGT90/95, VvUGT109/66, VvUGT82/93, VvUGT82/97, VvUGT104/53, VvUGT85/94, and VvUGT86/94. In addition, analysis of intrachromosomal duplication revealed that 159 VvUGT genes were from tandem repeat events, leading to the paralogous gene clusters in the genome (Table S1). These results indicated that segmental duplication and tandem duplication acted important roles in the expansion of the VvUGT family.

To further explore the evolution of the VvUGT gene family, we performed collinearity analysis between Arabidopsis and grape (Fig. 6). There were a total of 27 collinear UGT gene pairs between Arabidopsis and grape genomes, in which 24 genes from Arabidopsis and 27 genes from grape. These VvUGTs were located on 13 chromosomes (chr2, 3, 4, 5, 6, 8, 13, 14, 15, 16, 17, 18, and 19), and chromosomes 6 and 18 contained more collinear VvUGT genes than any other chromosome. The nonuniformity distribution of VvUGT genes on the chromosome indicated that there were genetic variations in the evolution of grape.

Enrichment analysis of grape

In order to understand the biological processes of VvUGTs, functional enrichment analysis was conducted by TBtools (Fig. 7; 8). GO analysis found that VvUGT family members were mainly concentrated on transferase activity, such as quercetin 3-O-glucosyltransferase activity, quercetin 7-O-glucosyltransferase activity, hexosyltransferase activity, flavonoids and flavonoid metabolic processes and other pathways (Fig. 7). KEGG enrichment analysis identified some relevant metabolic pathways of VvUGT family, including glucosinolate biosynthesis, tryptophan metabolism, metabolism of terpenoids and polyketides, amino acid metabolism, anthocyanin biosynthesis, and other biosynthesis of various secondary metabolites (Fig. 8). Based on KEGG and GO results, we identified 14 VvUGT genes involved flavonoid synthesis pathway. They are VvUGT60, VvUGT61, VvUGT62, VvUGT63, VvUGT79, VvUGT80, VvUGT81, VvUGT124, VvUGT165, VvUGT166, VvUGT167, VvUGT168, VvUGT169, VvUGT170. These candidate VvUGT genes related to flavonoid biosynthesis will provide evidence for the further research on plant secondary metabolites regulation mechanism.

Expression profiles of VvUGTs in different tissues

The expression pattern of VvUGT genes in five tissues (root, stem, leaf, flower and berrypericarp) was analyzed based on chip data (Fig. 9). The results showed that a total of 60 VvUGT genes were expressed in all tissues. The expression level of VvUGT99 in the root was higher than that in other tissues. 8 VvUGTs (VvUGT87, VvUGT97, VvUGT116, VvUGT119, VvUGT120, VvUGT125, VvUGT129 and VvUGT134) exhibited high levels of expression in all tissues of grape, suggesting their crucial roles in the growth and development of grape. Both VvUGT124 and VvUGT79 showed high expression level in berrypericarp and flower, respectively.

Expression profiles of VvUGTs in fruits at different development stages

To understand a potential function for VvUGT genes in fruit development, we investigated transcriptome data from two grape cultivars, including ‘Muscat Blanc’ and ‘Muscat Rouge’, at three developmental stages (Fig. 10). The findings revealed that 32 VvUTGs were not detected in any stages, while 149 VvUGTs were expressed at different stages. Among them, 13 VvUGT genes showed high expression abundance in all stages, which they may function in fruit development. In ‘Muscat Blanc’, VvUGT10 and VvUGT12 showed high expression levels in the young fruit stage, and expression of VvUGT134 and VvUGT145 was high in fruit veraison. VvUGT10, VvUGT51, VvUGT134 and VvUGT145 were highly expressed in maturity. Similar situation was observed at maturity of Muscat Rouge. The expression levels of VvUGT10, VvUGT12 and VvUGT13 genes of Muscat Rouge is lower than that of Muscat Blanc in at young fruit stage, while VvUGT22 and VvUGT124 genes of Muscat Rouge showed higher expression abundance than that of Muscat Blanc at both veraison and maturity.

Expression profiles of VvUGTs under abiotic stress

In order to further analyze the expression characteristics of VvUGT genes under abiotic stress, we investigated the response of VvUGT genes to salt and drought stress in this study (Fig. 11). 24 VvUGT genes were expressed under salt and drought stress. Under salt stress, the expression levels of 8 genes (VvUGT12, VvUGT30, VvUGT36, VvUGT59, VvUGT104, VvUGT122, VvUGT124, and VvUGT133) decreased after 16 days under salt stress treatment. 7 genes (VvUGT25, VvUGT40, VvUGT69, VvUGT97, VvUGT107, VvUGT125, VvUGT179) showed an increasing trend under salt stress. Under drought stress, the expression of VvUGT125 was significantly upregulated, while VvUGT122 exhibited a remarkable decrease after 16 days. Overall, the trend of gene expression was similar between salt and drought stress.

The UGTs are involved in the glycosylation and modification of plant secondary metabolites, and play an essential role in plant growth, development and stress response in plants (Zhang et al. 2020). Grapevine is one of important fruit crops cultivated worldwide to produce fresh fruits, juice, and dried raisins. It also serves as a vital raw material for wine industry (Fia et al. 2022). In the present study, a systematic analysis of the UGT gene family was conducted in grape, and a total of 181 VvUGT genes were identified. The number of UGTs are less than in grape compared to Epimedium pubescens (339) (Yao et al. 2022), Medicago sativa (409) (Ao et al. 2022) and Arachis hypogaea L. (267) (Ouyang et al. 2023), but more than Zea mays (147) (Li et al. 2014), Prunus persica L. (168) (Wu et al. 2017), Punica granatum L. (120) (Li et al. 2023), and Camellia sinensis (178) (Cui et al. 2016). Based on phylogenetic analysis, the 181 VvUGTs were divided into 17 groups (A-P, and R), in which group E had the largest number of VvUGT members (56), accounting for 30.9%. Notably, there were no VvUGT members in group Q, but it was found in maize (Li et al. 2014). Similar phenomenon was found in wheat (He et al. 2018), speculating that group Q may only exist in monocotyledons.

Gene duplication promotes the expansion of gene families during plant evolution. In our study, 15 segmental duplications and 103 tandem duplications were identified in grape genome (Fig. 5; Table S1), indicating that tandem duplication was a main driving force for expansion of VvUGT family. This was the reason that VvUGT genes were unevenly distributed on chromosomes as gene clusters (Fig. 1). Introns are crucial structural components of genes. Although they are not involved in the encoding of proteins, deletion or alteration of introns are generally regarded as key clues to understand the evolution or diversification of gene family (Wu et al. 2020; Zhang et al. 2022). Here, intron mapping of 181 VvUGTs revealed that 55.8% VvUGT members lacked introns, which was more than the number (44%) of petunia and the number (48%) of pomelo (Dong et al. 2022; Wu et al. 2020), while less than the number (58%) in Arabidopsis and the number (60%) of maize (Caputi et al. 2012; Li et al. 2014). Additionally, 38.7% of the VvUGTs contained one intron (Fig. 3).

Understanding the characteristics of cis-regulatory elements within promoters is a prerequisite for recognizing characteristics of gene transcriptional regulation and expression patterns. There were many different types of cis-elements in VvUGT promoters (Fig. 4A). Box4 was the most widely distributed light-responsive element, and was present in 91% of VvUGT genes (Fig. 4B). ABA serves as a major signaling molecule involved in various physiological processes in plants. Here, 117 of 181 VvUGT genes were found to contain ABRE in their promoter regions, which indicated that ABA was one of the important hormones affecting UGT genes expression. Moreover, 98% of the VvUGT promoters contained one or more cis-elements related to abiotic stresses (Fig. 4B). Among them, low-temperature responsiveness (LTR) elements were detected in 35% of VvUGT members, among which the number of LTR elements in VvUTG78 members was up to three, indicating that VvUGT genes might be involved in plant cold stress responses. In rice, expression of OsUGT90A1 was initially induced by low temperature, and its overexpression contributed to maintain membrane integrity during cold stress (Shi et al. 2020). In Camellia sinensis, CsUGT78A14 acted a critical role in cold stress by increasing flavonol accumulation and ROS scavenging capacity (Zhao et al. 2019). Altogether, VvUGT genes play a key role in plant growth and development, as well as response to abiotic stress.

To examine the function of VvUGT genes, expression analysis was performed in grape. Many studies has been showed that UGT genes function in plant growth and development (Mateo-Bonmatí et al. 2021; Yang et al. 2023). As shown in Fig. 9, 60 VvUGT genes were identified to express in all tissues. Among them, VvUGT97 and VvUGT119 genes showed higher levels in leaves and roots than in other tissues. VvUGT99 was only highly expressed in root, indicating that VvUGT99 might play a specific role in root development. VvUGT124 showed a significantly higher expression abundance in berrypericarp than other tissues, implying that this gene may be associated with fruit development (Fig. 9). Abiotic stresses seriously affects plant growth and productivity, resulting in economic losses (Jalil and Ansari 2020; Kumar 2020). Here, the expression patterns of VvUGT genes were explored under salt and drought (Fig. 11). We found that the expression level of VvUGT25 was increasingly upregulated during drought stress, but downregulated first, then significantly increased and reached the highest at 16 days under salt stress. This indicated that VvUGT25 served important roles in mediating plant response to drought and salt stress. In contrast, VvUGT122 and VvUGT124 always kept downward trend in response to drought and salt stress, indicating that the expression of VvUGT122 and VvUGT124 was inhibited by abiotic stress (Fig. 11). In rice, UGT85E1 gene has been reported to participate in drought stress tolerance through enhancing abscisic acid response (Liu et al. 2021). CrUGT87A1 from Carex rigescens can increase salt tolerance by accumulating flavonoids for antioxidation in Arabidopsis thaliana (Zhang et al. 2021). These results show that UGT family may play an important role in abiotic stress response of plants.

Secondary metabolites are closely related to fruit flavor, contribute to the improvement of fruit color, and their content has an important impact on the characteristics of grape and wine quality (Yilmaz and Toledo 2004; Georgiev et al. 2014). In recent years, several genes related to flavonoid biosynthesis have been identified, including UGT genes (Castellarin et al. 2006; Jeong et al. 2006; Falginella et al. 2012). In this study, KEGG and GO enrichment analysis revealed that VvUGT60, VvUGT61, VvUGT62, VvUGT63, VvUGT79, VvUGT80, VvUGT81, VvUGT124 were enriched in the flavonoid synthesis pathway (Fig. 7 ;8). Among them, we found that VvUGT124 was only expressed in the ‘Muscat Rouge’ (a red variety), while it was hardly expressed in ‘Muscat Blanc’ (a white variety) after fruit veraison. Specifically, the expression of VvUGT124 was significantly up-regulated in the veraison, and its expression level increased with the maturity of fruits. Similar expression pattern was found in UFGT gene in grapes (Boss et al. 1996). Previous study showed UGT78D2, the homolog gene of VvUGT124, participated in the convertion from flavonols to flavonol 3-O-glucosides in Arabidopsis pollens (Yonekura-Sakakibara et al. 2014). This result implied that VvUGT124 might be involved in regulation of secondary metabolism during grape veraison, which still need further experimental exploration.

In this study, we identified 181 VvUGT genes in the grape genome. The VvUGT genes were unevenly located at the 18 chromosomes, and clustered into 17 phylogenetic groups. Tandem duplication was the major driver for VvUGT gene family expansion. Promoter analysis found various cis-regulatory elements related to development, light response, phytohormone response, and stress response. KEGG and GO enrichment analysis identified 14 VvUGT genes involved flavonoid synthesis pathway. Expression profiles analysis indicated VvUGTs were involved in grape growth and development, as well as abiotic stress response. To sum up, this work would facilitate future functional investigations of VvUGT genes in grape.

Author Contributions

ZL and DZ conceptualized and designed the experiments; WH, WL, and CC analyzed the data; WH and XY wrote the paper; NH, MT and JH performed the experiments. All authors have read and approved the final manuscript.

Funding

This work was supported by Shanxi Agricultural University Science and Technology Innovation Improvement Project (CXGC2023086), China Agriculture Research System of Ministry of Finance (MOF) and Ministry of Agriculture and Rural Affairs (MARA) of the People’s Republic of China (CARS-29-yc-5), and Safe Conservation and Collection of Germplasm Resources of Jujubes and Grapes, Identification of Resources, Reproduction Cataloging and Entry into the Nursery (19221876).

Conflict of Interest

All authors declare no conflict of interest.

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Table1: Information of the VvUGTs identified in this study.

Name	Sequence ID	Chr ID	Start position	End position	Direction	Number of Amino Acid	Molecular Weight	Isoelectric point	Instability Index	Aliphatic Index	Grand Average of Hydropathicity
VvUGT1	VIT_201s0026g00630.1	1	9395296	9396993	+	506	56440.82	6.23	38.67	87.27	-0.185
VvUGT2	VIT_201s0010g00530.1	1	15343929	15346265	+	451	51250.83	6.13	43.96	94.66	-0.042
VvUGT3	VIT_202s0025g01240.1	2	1208153	1209559	+	468	51467.11	5.87	43.63	93.5	-0.043
VvUGT4	VIT_202s0012g02830.1	2	10849905	10851759	+	453	50576.65	6.68	30.23	98.37	-0.022
VvUGT5	VIT_202s0033g00130.1	2	13674127	13675656	+	509	54978.65	6.38	55.82	75.32	-0.261
VvUGT6	VIT_202s0033g00180.1	2	13768305	13769834	+	509	55226.02	6.38	52.47	75.32	-0.243
VvUGT7	VIT_202s0033g00240.1	2	13919731	13921116	+	461	50670.15	6.73	56.08	81.48	-0.193
VvUGT8	VIT_203s0063g00040.1	3	3587099	3588547	-	482	53841.81	5.61	42.84	87.66	-0.156
VvUGT9	VIT_203s0063g00050.1	3	3591619	3593043	-	474	52745.57	5.9	50.01	85.13	-0.203
VvUGT10	VIT_203s0180g00200.1	3	6124675	6126114	-	479	53765.18	5.24	51.31	87.66	-0.186
VvUGT11	VIT_203s0180g00280.1	3	6255590	6257062	+	490	54466.76	5.44	43.57	92.24	-0.114
VvUGT12	VIT_203s0180g00320.1	3	6282074	6283513	+	479	53645.99	5.27	50.08	86.03	-0.173
VvUGT13	VIT_203s0091g00040.1	3	6338114	6339550	+	478	53841.2	5.58	53.97	87.66	-0.209
VvUGT14	VIT_203s0017g01040.1	3	16533806	16536517	-	489	55138.82	5.51	40.73	92.25	-0.073
VvUGT15	VIT_203s0017g01130.1	3	16716769	16718607	-	475	53068.29	5.35	43.43	88.23	-0.062
VvUGT16	VIT_203s0017g01140.1	3	16773485	16775172	-	457	51157.64	5.98	42.25	85.71	-0.214
VvUGT17	VIT_203s0017g01360.1	3	17373374	17375164	+	420	46603.47	5.75	40.75	87.05	-0.166
VvUGT18	VIT_203s0017g01990.1	3	18463726	18465183	+	485	53399.45	5.48	42.97	98.8	0.071
VvUGT19	VIT_203s0017g01996.1	3	18509235	18510662	+	475	52104.61	5.47	35.33	96.57	-0.018
VvUGT20	VIT_203s0017g02140.1	3	18846514	18863031	+	475	52013.37	5.54	42.73	96.06	0.051
VvUGT21	VIT_204s0023g01240.1	4	17638953	17640582	-	501	55679.65	5.78	43.71	101.16	0.032
VvUGT22	VIT_204s0023g01290.1	4	17705304	17706812	+	502	55116.92	5.61	40.64	94.54	0.031
VvUGT23	VIT_204s0044g01530.1	4	23119212	23120549	+	445	50008.79	7.23	53.22	96.63	-0.186
VvUGT24	VIT_204s0044g01540.1	4	23134466	23135818	-	450	50052.57	5.27	57.42	98.51	-0.137
VvUGT25	VIT_205s0062g00270.1	5	18421988	18423385	+	465	51658.99	5.32	48.81	87.42	-0.152
VvUGT26	VIT_205s0062g00300.1	5	18450755	18457884	+	530	59071.48	5.79	47.83	84.96	-0.171
VvUGT27	VIT_205s0062g00310.1	5	18459997	18461394	+	430	47659.55	5.46	43.89	87.77	-0.156
VvUGT28	VIT_205s0062g00350.1	5	18473416	18474813	+	465	51488.78	5.52	49.73	87.01	-0.165
VvUGT29	VIT_205s0062g00400.1	5	18544519	18546423	-	634	70563.62	4.88	38.58	82.11	-0.414
VvUGT30	VIT_205s0062g00430.1	5	18570221	18571621	+	466	51380.78	5.21	54.68	93.93	-0.074
VvUGT31	VIT_205s0062g00460.1	5	18605640	18607037	+	465	51097.32	5.25	50.61	92.88	-0.077
VvUGT32	VIT_205s0062g00470.1	5	18614167	18615558	-	463	52028.64	5.04	41.41	87.58	-0.186
VvUGT33	VIT_205s0062g00540.1	5	18717384	18718780	+	392	43470.49	5.15	50.14	97.73	-0.102
VvUGT34	VIT_205s0062g00570.1	5	18749370	18750740	+	456	50330.4	5.43	52.03	93.84	-0.104
VvUGT35	VIT_205s0062g00590.1	5	18760946	18762337	-	463	52021.55	5.09	41.03	87.37	-0.192
VvUGT36	VIT_205s0062g00630.1	5	18804776	18806185	-	469	52071.18	4.98	47.84	87.27	-0.215
VvUGT37	VIT_205s0062g00640.1	5	18853664	18855073	-	469	51993.95	5.04	48.51	83.71	-0.227
VvUGT38	VIT_205s0062g00700.1	5	18934062	18935456	+	464	51218.29	5.19	43.94	90.97	-0.158
VvUGT39	VIT_205s0062g00710.1	5	18956099	18957490	+	463	50884.87	5.34	46.87	91.36	-0.16
VvUGT40	VIT_205s0062g00740.1	5	18999810	19001153	+	447	49642.69	5.23	46.86	93.31	-0.075
VvUGT41	VIT_205s0062g01270.1	5	20096453	20097943	-	496	55058.89	5.94	50.09	84.17	-0.183
VvUGT42	VIT_205s0062g01280.1	5	20108240	20109943	-	479	53241.86	6.04	48.47	81.84	-0.195
VvUGT43	VIT_205s0102g00930.1	5	22907018	22908525	+	476	54212.1	6.03	41.26	83.76	-0.187
VvUGT44	VIT_205s0102g00990.1	5	22987434	22989011	+	487	55432.62	6.08	42.79	84.87	-0.172
VvUGT45	VIT_205s0094g00960.1	5	24298325	24300200	+	457	50672.13	5.37	40	84.68	-0.058
VvUGT46	VIT_205s0094g00990.1	5	24324856	24326531	+	458	51309.81	5.68	36.17	87.03	-0.106
VvUGT47	VIT_205s0094g01000.2	5	24330004	24332084	+	453	50826.58	5.29	55.95	86.71	-0.178
VvUGT48	VIT_205s0094g01030.1	5	24336681	24348881	+	834	94596.51	5.19	42.59	84.65	-0.224
VvUGT49	VIT_205s0094g01040.1	5	24357327	24358983	-	454	51445.77	5.7	52.86	86.3	-0.241
VvUGT50	VIT_205s0094g01050.1	5	24370871	24372633	-	459	52018.59	5.32	52.22	86.84	-0.232
VvUGT51	VIT_206s0004g01640.1	6	2038603	2040081	-	492	54944.2	5.85	48.45	82.44	-0.158
VvUGT52	VIT_206s0004g01670.1	6	2069738	2071222	-	494	55598.79	6.26	39.42	83.54	-0.196
VvUGT53	VIT_206s0004g05780.1	6	6525838	6527623	+	442	49510.59	5.72	54.06	89.95	-0.187
VvUGT54	VIT_206s0004g06370.1	6	7130000	7131613	+	537	59860.66	5.6	41.48	88.03	-0.104
VvUGT55	VIT_206s0004g06390.1	6	7141267	7142703	+	478	52420.24	5.34	47.34	97.32	0.029
VvUGT56	VIT_206s0004g06400.1	6	7152176	7153585	+	469	51846.78	5.85	40.15	93.11	-0.012
VvUGT57	VIT_206s0004g07240.1	6	7986284	7988366	+	460	51274.8	6.57	43.45	89.78	-0.015
VvUGT58	VIT_206s0004g07250.1	6	7994721	7996737	+	449	50216.5	5.5	42.69	93.56	-0.022
VvUGT59	VIT_206s0004g07310.1	6	8036833	8038426	+	452	50698.89	6.04	39.46	100.07	0.109
VvUGT60	VIT_206s0009g01960.1	6	14190688	14192343	+	451	48512.79	6.02	33.94	89.73	0.036
VvUGT61	VIT_206s0009g01990.1	6	14213942	14215406	+	445	47598.88	6.64	35.64	93.82	0.071
VvUGT62	VIT_206s0009g02000.1	6	14217726	14226707	+	457	49492.77	6.39	34.2	86.19	-0.114
VvUGT63	VIT_206s0009g02010.1	6	14238458	14239922	+	445	47505.69	6.64	33.9	92.52	0.065
VvUGT64	VIT_206s0009g02450.1	6	14920222	14921780	+	424	47769.97	5.48	39.88	91.06	-0.042
VvUGT65	VIT_206s0061g00360.1	6	17756872	17758382	-	474	52642.36	5.8	29.89	96.48	-0.004
VvUGT66	VIT_206s0061g01430.1	6	19307651	19309012	+	453	50882.95	6	56.15	98.76	-0.042
VvUGT67	VIT_207s0289g00050.1	7	13938451	13940132	-	456	50613.39	5.54	43.55	88.88	-0.059
VvUGT68	VIT_207s0289g00060.2	7	13969701	13971495	-	456	50629.22	5.43	41.75	89.3	-0.054
VvUGT69	VIT_208s0040g01470.1	8	12639957	12641499	+	481	54222.83	6.18	41.29	90.33	-0.151
VvUGT70	VIT_208s0040g01520.1	8	12677222	12678706	-	494	55717	6.3	41.86	90.32	-0.161
VvUGT71	VIT_208s0007g04570.3	8	18481568	18483046	-	492	54996.69	6.26	47.63	94.86	-0.126
VvUGT72	VIT_208s0007g04580.1	8	18495213	18496436	+	407	45556.65	5.51	49.12	92.43	-0.141
VvUGT73	VIT_208s0007g04590.1	8	18500479	18501891	-	470	52583.95	5.73	41.36	98.89	-0.092
VvUGT74	VIT_208s0007g04600.1	8	18503840	18505072	-	380	42606.63	6.16	52.95	87.45	-0.329
VvUGT75	VIT_208s0007g04610.1	8	18507500	18508987	-	464	51534.79	5.41	49.41	102.31	0.006
VvUGT76	VIT_208s0007g05130.1	8	19062236	19063704	-	458	51095.49	6.73	48.43	88.54	-0.156
VvUGT77	VIT_208s0007g08910.1	8	22239284	22243588	-	540	60562.23	7.36	42.46	85.91	-0.247
VvUGT78	VIT_209s0002g06590.1	9	6522570	6523811	-	413	45673.81	4.9	49.32	86.44	-0.212
VvUGT79	VIT_211s0052g01580.1	11	19310722	19313103	-	521	56883.47	6.95	37.32	92.55	0.042
VvUGT80	VIT_211s0052g01600.1	11	19328609	19331579	-	459	49684.44	6.2	40.84	97.73	0.117
VvUGT81	VIT_211s0052g01630.1	11	19367554	19373012	-	480	52418.65	6.54	44.21	99.96	0.114
VvUGT82	VIT_212s0055g00020.1	12	12595769	12597199	-	476	52930.59	5.14	51.69	85.97	-0.032
VvUGT83	VIT_212s0055g00030.1	12	12614143	12616908	-	477	53001.85	6.44	49.92	82.91	-0.065
VvUGT84	VIT_212s0055g00050.1	12	12637374	12638774	-	466	51697.44	6.41	50.09	84.25	-0.072
VvUGT85	VIT_212s0055g00160.1	12	12742572	12743882	+	436	48498.2	5.3	52.66	85.83	-0.061
VvUGT86	VIT_212s0055g00180.1	12	12758874	12760304	+	476	53372.94	5.4	51.26	86.81	-0.07
VvUGT87	VIT_212s0055g00200.1	12	12790596	12792026	+	476	53257.96	5.41	47.89	90.48	-0.011
VvUGT88	VIT_212s0055g00250.1	12	12882390	12883814	+	474	52801.49	5.45	46.68	89.85	-0.018
VvUGT89	VIT_212s0055g00280.1	12	12922979	12924397	+	472	52814.76	5.39	49.06	88.39	0.01
VvUGT90	VIT_212s0055g00290.1	12	12949533	12964018	+	932	103449.59	5.73	49.19	90.26	-0.007
VvUGT91	VIT_212s0055g00310.1	12	12989378	12990784	+	468	51817.5	5.47	50.62	87.07	-0.005
VvUGT92	VIT_212s0055g00320.1	12	12999329	13000735	+	468	51892.61	5.61	45.41	91.22	-0.006
VvUGT93	VIT_212s0034g00030.1	12	15391502	15393291	+	426	47585.14	5.69	54.17	77.3	-0.179
VvUGT94	VIT_212s0034g00040.1	12	15417193	15418602	-	469	52379.89	5.46	45.09	86.84	-0.024
VvUGT95	VIT_212s0034g00060.1	12	15441562	15442983	-	473	52573.19	5.58	43.52	89.2	-0.031
VvUGT96	VIT_212s0034g00080.1	12	15462117	15463526	-	469	52339.95	5.53	48.03	89.53	-0.01
VvUGT97	VIT_212s0034g00130.1	12	15534618	15536048	-	476	52706.46	5.74	48.29	87.18	-0.011
VvUGT98	VIT_212s0034g00140.1	12	15584311	15590823	-	491	54575.24	6.4	41.79	86.9	0.062
VvUGT99	VIT_212s0034g00160.1	12	15623174	15624592	-	472	52676.12	5.17	49.33	84.22	-0.126
VvUGT100	VIT_212s0034g01120.1	12	17015928	17017355	+	475	53025.68	5.45	45.65	90.25	-0.041
VvUGT101	VIT_213s0019g02810.1	13	4023902	4025493	+	462	51709.26	5.94	52.71	96.23	-0.085
VvUGT102	VIT_213s0019g02860.1	13	4055461	4063630	+	815	90995.67	6.04	46.33	100.63	0.046
VvUGT103	VIT_213s0019g02890.1	13	4089631	4091178	+	480	53637.42	5.83	35.04	91.6	-0.14
VvUGT104	VIT_213s0019g02940.1	13	4114395	4115920	+	482	54573.68	6.16	47.81	82.51	-0.161
VvUGT105	VIT_213s0019g02990.1	13	4145646	4147193	-	480	53860.68	5.8	38.97	92.63	-0.147
VvUGT106	VIT_213s0019g03020.1	13	4159206	4160748	-	478	53629.32	5.63	38.16	87.49	-0.163
VvUGT107	VIT_213s0019g03040.1	13	4177370	4179239	-	534	59669.69	5.94	44.49	87.81	-0.05
VvUGT108	VIT_213s0019g03060.1	13	4183650	4185218	-	479	53714.73	6.18	45.74	88.33	-0.118
VvUGT109	VIT_213s0074g00610.1	13	8644831	8646180	+	449	50552.47	6.88	53.66	89.4	-0.194
VvUGT110	VIT_213s0047g01230.1	13	18377964	18379304	-	446	50490.22	6.2	49.26	90.92	-0.265
VvUGT111	VIT_214s0006g02140.1	14	19247015	19248472	-	485	52650.27	6.41	33.28	91.07	-0.025
VvUGT112	VIT_214s0006g02145.1	14	19302993	19304495	-	500	54568.8	6.72	38.36	95.54	0.029
VvUGT113	VIT_214s0068g00440.1	14	24220065	24221586	+	453	50467.53	5.11	34.27	82.63	-0.14
VvUGT114	VIT_214s0068g00450.1	14	24222580	24224424	+	454	50831.4	4.97	38.43	91.06	-0.035
VvUGT115	VIT_214s0068g00460.1	14	24225718	24227483	+	454	50728.61	5.61	33.51	95.15	-0.045
VvUGT116	VIT_214s0068g00500.1	14	24240476	24246177	+	454	50760.09	5.17	33.47	89.56	-0.13
VvUGT117	VIT_214s0066g01480.1	14	27896211	27897647	+	478	53749.7	5.23	41.69	84.58	-0.184
VvUGT118	VIT_215s0021g02060.1	15	12835467	12836873	+	468	51794.92	6.1	48.19	97.03	-0.029
VvUGT119	VIT_215s0046g01190.1	15	18217502	18218911	-	469	51528.52	6.03	48.86	94.37	-0.028
VvUGT120	VIT_215s0046g01210.1	15	18227103	18228512	-	469	51339.38	5.93	50.46	95.63	0.022
VvUGT121	VIT_215s0046g01950.1	15	18779360	18786610	+	489	54395.91	5.71	55.39	94.91	0.024
VvUGT122	VIT_215s0046g01960.1	15	18789471	18790868	-	465	51665.85	5.75	41.79	93.78	-0.009
VvUGT123	VIT_215s0046g01970.1	15	18796883	18804535	-	424	47040.98	5.78	39.69	91.3	0.008
VvUGT124	VIT_216s0039g02230.1	16	2333232	2334679	-	456	50030.41	5.86	40.8	86.18	0.003
VvUGT125	VIT_216s0115g00340.1	16	4082672	4084093	+	473	51903.41	5.63	43.09	88.65	0.041
VvUGT126	VIT_216s0115g00360.1	16	4251316	4252758	-	480	53314.72	5.87	47.65	97.48	-0.004
VvUGT127	VIT_216s0022g01970.1	16	14269762	14271192	-	476	52376.39	5.9	46.76	93.38	-0.047
VvUGT128	VIT_216s0050g00240.1	16	17046250	17047656	+	468	51734.37	5.97	42.58	89.59	0.029
VvUGT129	VIT_216s0050g01580.1	16	18494181	18503321	+	471	51742.88	6.03	52.84	97.3	0.106
VvUGT130	VIT_216s0050g01600.1	16	18520803	18522209	+	468	51371.11	5.58	45.05	94	0.095
VvUGT131	VIT_216s0050g01610.1	16	18525704	18527116	+	470	51851.81	5.58	50.2	96.28	0.09
VvUGT132	VIT_216s0050g01660.1	16	18554821	18556254	-	477	52269.35	6.24	50.3	92.58	0.039
VvUGT133	VIT_216s0050g01680.1	16	18585339	18586769	-	476	52021	6.84	45.28	92.35	0.015
VvUGT134	VIT_217s0000g04750.1	17	5148100	5149542	+	480	52554.92	5.62	36.71	94.6	-0.046
VvUGT135	VIT_217s0000g04760.1	17	5149856	5151301	-	481	52744.58	5.71	35.15	97.32	0.042
VvUGT136	VIT_217s0000g07015.3	17	7726000	7727572	-	462	50727.25	5.22	38.77	96.69	0.007
VvUGT137	VIT_217s0000g07070.1	17	7777018	7784341	-	745	82475.66	5.14	38.15	94.36	0.011
VvUGT138	VIT_217s0000g07080.1	17	7802591	7804070	-	458	50609.3	5.52	40.91	94.76	-0.031
VvUGT139	VIT_217s0000g07100.1	17	7856461	7857894	-	456	50271.84	5.82	36.67	95.39	-0.024
VvUGT140	VIT_217s0000g07110.1	17	7861011	7862576	-	445	49540.88	5.66	38.85	94.63	-0.052
VvUGT141	VIT_217s0000g07790.1	17	8768709	8770406	+	457	50395.88	5.3	45.92	86.35	-0.077
VvUGT142	VIT_218s0001g05910.1	18	4550434	4552215	+	495	54979.1	5.03	39.04	82.95	-0.123
VvUGT143	VIT_218s0001g05950.1	18	4564181	4565715	-	474	52938.12	5.78	35.02	87.85	-0.101
VvUGT144	VIT_218s0001g05990.1	18	4584527	4586041	-	472	52868.82	4.84	39.06	90.53	-0.085
VvUGT145	VIT_218s0001g06060.1	18	4605469	4606985	-	473	52901.99	5.04	34.83	89.49	-0.1
VvUGT146	VIT_218s0001g06080.1	18	4610506	4612054	-	468	53162.76	5.44	33.56	81.69	-0.242
VvUGT147	VIT_218s0001g06090.1	18	4613744	4615282	-	476	53382.23	5.16	35.55	87.29	-0.184
VvUGT148	VIT_218s0001g06120.1	18	4629602	4631139	-	476	53226.94	5.1	35.3	85.65	-0.194
VvUGT149	VIT_218s0001g10110.1	18	8470824	8472314	-	496	55873.34	6.1	41.56	88.69	-0.189
VvUGT150	VIT_218s0001g12040.1	18	10250572	10252002	+	476	52698.72	5.06	47.66	99.71	0.022
VvUGT151	VIT_218s0041g00710.1	18	25251260	25252690	-	476	52866.64	5.46	43.96	89.12	-0.087
VvUGT152	VIT_218s0041g00740.1	18	25261347	25262789	-	480	53292.95	5.78	39.35	86.4	-0.134
VvUGT153	VIT_218s0041g00790.1	18	25300595	25302052	-	485	53881.57	5.38	50.54	89.48	-0.106
VvUGT154	VIT_218s0041g00800.1	18	25305917	25307515	+	532	59100.63	5.61	52.76	91.47	-0.094
VvUGT155	VIT_218s0041g00810.1	18	25313263	25314864	-	443	48901	5.58	46.7	87.86	-0.034
VvUGT156	VIT_218s0041g00830.1	18	25326336	25327823	-	495	54873.12	6.25	47.15	90.79	-0.035
VvUGT157	VIT_218s0041g00840.1	18	25334444	25335895	-	483	53289.99	5.74	42.18	88.24	-0.022
VvUGT158	VIT_218s0041g00900.1	18	25392826	25394373	+	515	57105.33	5.8	53.07	91.07	-0.1
VvUGT159	VIT_218s0041g00910.1	18	25406239	25407690	-	483	53122.95	6.03	48.46	86.83	-0.047
VvUGT160	VIT_218s0041g00920.1	18	25418483	25419934	-	483	53664.65	6.37	45.97	86.42	-0.071
VvUGT161	VIT_218s0041g00950.1	18	25452355	25453806	-	483	53290.95	5.74	43.83	87.2	-0.053
VvUGT162	VIT_218s0041g01000.1	18	25522586	25524037	-	483	53658.65	6.03	46.05	87.43	-0.072
VvUGT163	VIT_218s0041g01040.1	18	25565624	25567075	-	483	53323.92	5.44	43	87.99	-0.055
VvUGT164	VIT_218s0041g01080.1	18	25604405	25605856	-	483	53450.29	5.74	41.95	88.8	-0.018
VvUGT165	VIT_219s0014g01950.1	19	2128632	2130017	+	461	50560.24	6.15	40.78	87.42	0.004
VvUGT166	VIT_219s0014g01970.1	19	2134035	2135411	-	458	50658.31	6.24	40.42	85.81	-0.077
VvUGT167	VIT_219s0014g01980.1	19	2155950	2157323	-	457	50542.45	6.63	45.6	88.14	-0.06
VvUGT168	VIT_219s0014g02000.1	19	2189219	2190597	-	452	50051.89	7.27	40.66	88.69	-0.056
VvUGT169	VIT_219s0014g02030.1	19	2206227	2207558	+	443	48943.33	6.38	35.58	91.58	-0.058
VvUGT170	VIT_219s0014g02070.1	19	2231065	2232444	+	459	50598.01	5.68	39.61	88.82	-0.058
VvUGT171	VIT_219s0085g00760.1	19	23206320	23207738	+	472	52174.27	5.06	42.41	102.22	0.067
VvUGT172	VIT_207s0151g00540.1	7_random	964388	966182	-	456	50600.23	5.52	42.2	88.88	-0.066
VvUGT173	VIT_200s0218g00130.1	Un	14055159	14056598	+	479	53732.8	5.78	47.63	91.82	-0.14
VvUGT174	VIT_200s0218g00140.1	Un	14060242	14061681	+	479	53863.87	5.83	50.97	91.42	-0.161
VvUGT175	VIT_200s0218g00150.1	Un	14064350	14071318	+	876	98904.47	5.61	49.49	90.92	-0.202
VvUGT176	VIT_200s0218g00160.1	Un	14074251	14075645	+	464	52762.05	5.92	46.74	93.92	-0.177
VvUGT177	VIT_200s0324g00050.1	Un	23430045	23431602	-	482	54596.8	6.14	40.76	74.9	-0.197
VvUGT178	VIT_200s0324g00060.1	Un	23438469	23440020	-	480	53977.71	5.42	36.62	80.19	-0.194
VvUGT179	VIT_200s0324g00070.1	Un	23446978	23448704	-	483	54572.24	5.88	45.66	80.14	-0.311
VvUGT180	VIT_200s0324g00100.1	Un	23464790	23466333	-	487	54473.85	6.28	39.16	87.1	-0.073
VvUGT181	VIT_200s0391g00010.1	Un	27126122	27137865	-	1397	156468.12	5.8	40.01	85.03	-0.08

No competing interests reported.

TableS1.docx
Appendix A. Supplementary data Table1. S1 Tandem duplication of UGTs in grape

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Editorial decision: Revision requested
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Identification, evolution and expression analysis of the UDP-glycosyltransferase gene family in grape (Vitis vinifera L.)

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Version 1

Abstract

Figures

Introduction

Materials and methods

Phylogenetic, gene structure and conserved motif analysis

Detection of cis-regulatory elements and enrichment analysis

Results

Analysis of phylogenetic, gene structure and conserved motif

Interspecific and intraspecific collinearity analysis

Enrichment analysis of grape

Discussion

Conclusion

Declarations

References

Tables 1

Additional Declarations

Supplementary Files

Status:

Version 1