Genome-wide in silico identification and expression analysis of beta-galactosidase family members in sweetpotato [Ipomoea batatas (L.) Lam.]

Genome-wide in silico identification and expression analysis of beta-galactosidase family members in sweetpotato [Ipomoea batatas (L.) Lam.] | 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 in silico identification and expression analysis of beta-galactosidase family members in sweetpotato [ Ipomoea batatas (L.) Lam.] Fuyun Hou, Zhen Qin, Taifeng Du, Tao Xu, Aixian Li, Shunxu Dong, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-32133/v3 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 27 Feb, 2021 Read the published version in BMC Genomics → Version 3 posted 5 You are reading this latest preprint version Show more versions Abstract Background: Sweetpotato ( Ipomoea batatas (L.) Lam.) serves as an important food source for human beings. β-galactosidase (bgal) is a glycosyl hydrolase involved in cell wall modification, which plays essential roles in plant development and environmental stress adaptation. However, the function of bgal genes in sweetpotato remains unclear. Results: In this study, 17 β-galactosidase genes ( Ibbgal ) were identified in sweetpotato, which were classified into seven subfamilies using interspecific phylogenetic and comparative analyses. The promoter regions of Ibbgal s harbored several stress, hormone and light responsive cis-acting elements. Quantitative real-time PCR results displayed that Ibbgal genes had the distinct expression patterns across different tissues and varieties. Moreover, the expression profiles under various hormonal treatments, abiotic and biotic stresses were highly divergent in leaves and root. Conclusions: Taken together, these findings suggested that Ibbgals might play an important role in plant development and stress responses, which provided evidences for further study of bgal function and sweetpotato breeding. Epigenetics & Genomics sweetpotato β-Galactosidase gene expression stress Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Background β-galactosidases (EC 3.2.1.23; bgal) widely exist in higher plants. Plant β-galactosidase belongs to the glycoside hydrolase 35 (GH35) families [1], which catalyzes the removal of terminal galactosyl residues from carbohydrates, glycoproteins and galactolipids [2, 3]. In plants, β-galactosidase has been reported to degrade structural polysaccharides in plant cell walls to release free galactose during a variety of biological processes, including cell wall expansion and degradation, metabolic recycling of galactolipids and glycoproteins, and turnover of signaling molecules during ripening [4, 5]. In higher plants, bgals have been grouped into two classes based on their substrate preference [6]. Enzymes in the first class prefer pectic β-(1→4)-galactan as the substrate, and enzymes in the other prefer the β-(1→3) and (1→6)-galactan backbones of arabinogalactan proteins [7, 8]. A typical bgal protein contains the GH35 conserved site in the N-terminal region [9]. Like other glycosidase families, bgal genes are ubiquitously expressed in many plants, such as tomato [2], papaya [10], Arabidopsis [11], Brassica campestris [12] and rice [13]. Plant bgal genes are widely involved in the modification of the architecture of cell walls and intercellular attachments [14, 15]. bgal genes also respond to plant growth and development including fruit development and ripening [16, 17], seed germination [18, 19], and root development [20, 21]. In most fruits, bgal genes exhibit differential expression patterns during flowering and fruit development [16, 12]. In Cicer arietinum , Canbgal-5 expression is relevant to young and meristematic stages with a high cell division rate, while CanBGal-1 and CanBGal-4 are strongly related to later stages of epicotyl growth [3]. In addition, bgal genes can be regulated by abiotic and biotic stresses [22]. For example, Atbgal1 was reported to be induced by salt stress or pathogen attack [23]. Likewise, the transcription level of β-galactosidase in cowpea is reduced under salt treatments [24], and the bgal mRNA level in peach is highly suppressed by water stress [25]. In addition, bgal genes have been found to play a role in a variety of biological processes through ethylene signal transduction [26, 11]. However, the function of bgal has not been studied in sweetpotato ( Ipomoea batatas (L.) Lam). Sweetpotato is an important food crop which is widely grown in tropical and subtropical areas, especially in Asia and sub-Saharan Africa. Due to its outcrossing hexaploidy (2n=6×=90), the genomic research in sweetpotato is very complicated [27, 28]. So far, no high-quality genome sequence of sweetpotato has been available. Although bgal genes are widely isolated from many plant species, its function in sweetpotato remains unknown. In the present study, we firstly identified 17 bgal genes ( Ibbgal ) in sweetpotato, and then investigated their phylogeny, motif compositions and predicted cis-elements using various bioinformatics tools. In addition, the expression patterns of these 17 Ibbgals in different tissues of two cultivars were investigated under three exogenous hormones, two abiotic and one biotic stress conditions. Our study will lay the foundation for further research on the function of bgal gene in plants, and provide new insight into different regulatory mechanisms in plant growth through bgal -mediated responses to environmental stresses in sweetpotato. Results Identification and characterization of Ibbgal genes in sweetpotato A total of 17 Ibbgal genes were isolated from sweetpotato after local BLAST using the conserved bgal domain. The deduced amino acid sequences of the Ibbgal proteins were used to predict their protein lengths, signal peptides, pI values, molecular weights, sub-cellular localization and the possible N-glycosylation sites (Table 1). Characteristic analysis showed that these 17 Ibbgals were 673 to 1110 aa in length, the predicted MWs and pIs ranged from 74.8 kDa to 125.1 kDa and 5.31 to 6.16, respectively. The predicted localization of most Ibbgals varied and included the chloroplast, vacuole, and nucleus. Only one Ibbgal, Ibbgal7, was found to be located in the extracellular. Signal peptides analysis revealed that all Ibbgals, except for Ibbgal4, Ibbgal5, Ibbgal10, Ibbgal13 and Ibbgal17, contained a signal peptide. The number of N-glycosylation sites varied from 1 to 6, wherein Ibbgal13 and Ibbgal16 contained 6 N-glycosylation sites. Conserved motifs and phylogenetic analysis of the Ibbgal proteins In this study, the β-galactosidase active site was found in all Ibbgal proteins. However, all but Ibbgal13 have the active site consensus sequence GGP[LIVM]xQxENE[FY] of the GH35 β-galactosidase family. In addition, all Ibbgal members carried a Gal-lectin domain at the C-terminus of the protein sequence, except for Ibbgal2, Ibbgal5, Ibbgal12, Ibbgal13, and Ibbgal17. Motif analysis showed that motif 1 was found in all Ibbgals except Ibbgal13, and motifs 2-6 were found in all Ibbgals except Ibbgal11 and Ibbgal17 (Fig. 1). A total of 34 bgal genes from sweetpotato and Arabidopsis were classified into seven subgroups, designated as A, B, C, D, E, F and G, using phylogenetic analysis (Fig. 2). Among these groups , groups A and D were the largest groups with four Ibbgal genes in each. Groups B and E had three Ibbgal genes. However, Ibbgal9 , Ibbgal17 and Ibbgal13 were classified into group C, F and E, respectively. Cis-element prediction of Ibbgal genes To understand the potential transcriptional regulatory mechanisms of the Ibbgal genes, the cis-elements of each Ibbgal promoter sequences were predicted and analyzed (Table 2). The promoters of Ibbgals were classified into at least four types of cis-elements, including plant hormone responsive elements, light responsive elements, stress responsive elements, and other elements. Most Ibbgal promoters had the GARE (gibberellin-responsive element), ERE (ethylene-responsive element) cis-elements, AuxRE and CATATGGMSAUR motifs which were involved in plant hormone response. Most Ibbgal promoters, except Ibbgal6 , Ibbgal16 and Ibbgal17 , contained circadian and EE elements participated in circadian regulation. In addition, at least five light response elements were found in each Ibbgal gene, which might be essential for plant growth and development. Interestingly, the Ibbgal s contained the MYC-like and ABRE (Abscisic acid response element) cis-elements mediated the responses to abotic stresses. Expression profiles of Ibbgal genes in tissues and different root development stages To identify the potential functions of Ibbgal genes, we analyzed the transcript levels of Ibbgal s in various tissues of cv . Jishu25 and Jishu29, including leaf, stem lip, stem, fibrous root, and storage root. 47% of Ibbgal s had similar expression patterns in five tissues of two cultivars (Fig. 3A). For example, Ibbgal4 , Ibbgal10 , Ibbgal13 and Ibbgal17 were highly expressed in five tissues, whereas Ibbgal14 , Ibbgal15 and Ibbgal16 were poorly expressed in these tissues. Intriguingly, the expression of Ibbgal4 in fibrous root was significantly higher than that of storage root, while Ibbgal3 and Ibbgal10 were expressed at higher levels in lip than other tissues. However, the transcript of Ibbgal17 mRNA in cv . Jishu25 was prominently higher in storage root than fibrous root, whereas that in cv . Jishu29 had no significant difference in the roots. Similarly, the expression of Ibbgal11 had the opposite pattern in the storage and fibrous roots between cv . Jishu25 and Jishu29. In root development stages, 6 (35.3%) Ibbgal transcripts were down-regulated including Ibbgal2 , Ibbgal3 , Ibbgal4 , Ibbgal6 , Ibbgal10 , and Ibbgal16 , whereas 6 Ibbgal transcripts were up-regulated, two Ibbgal genes ( Ibbgal14 and Ibbgal15 ) were not detected in root development. It is interesting that the Ibbgal11 and Ibbgal12 transcripts had the opposite expression pattern between cv. Jishu25 and Jishu29 (Fig. 3B). Expression profiles of Ibbgal genes in response to abiotic and biotic stresses Besides their functions in plant growth and development, Ibbgal genes may also be involved in biotic and abiotic stress responses. For sweetpotato, salinity and drought are the most dominant factors which limit the growth and yield among various abiotic stresses.Under salt stress, all Ibbgal genes were up-regulated in these two cultivars (Fig.4). Some genes had the highest expression levels at 12 h in the leaves, whereas other Ibbgal genes in roots were expressed at a high level at 6 h and 48 h after salt stress. In addition, Ibbgal2 , Ibbgal4 , Ibbgal5 and Ibbgal13 in the leaves were unregulated remarkably by at least 10-fold induction after salt stress. These results indicated that Ibbgal genes were involved in salt stress response in sweetpotato. Under drought stress (Fig. 4), all Ibbgal genes were up-regulated in the leaves and roots of cv . Jishu29, while Ibbgal3 , Ibbgal6 , Ibbgal10 , and Ibbgal17 were down-regulated in the leaves of Jishu25, and Ibbgal1 , Ibbgal3 and Ibbgal16 expression were also reduced in the root of Jishu25. Amongst the up-regulated genes, the expression of Ibbgal2, Ibbgal4, Ibbgal8, Ibbgal9 and Ibbgal13 reached the peak at 12 h after stress, and Ibbgal4 was the most up-regulated gene with at least 81-fold induction in the two cultivars leaves, suggesting that Ibbgals in the different cultivars responded to drought treatment differently. Black spot, caused by Ceratocystis fimbriata ( C. fimbriata ), is one of the main diseases in sweetpotato production, which seriously affects the quality and yield of sweetpotato. After the pathogen infection, Ibbgal genes had different expression patterns in the leaves and roots of these two cultivars (Fig. 4). Ibbgal5, Ibbgal10 , Ibbgal11 and Ibbgal16 transcripts were induced by the pathogen infection in these two cultivars. It is worth noting that Ibbgal15 expression in the leaves and roots of cv. Jishu25 was up-regulated, whereas down-regulated in cv. Jishu29. Collectively, these results implied that Ibbgals in the different cultivars might have different functions under abiotic and biotic stresses. Expression profiles of Ibbgal genes in response to various hormone treatments To survey the role of Ibbgal genes in plant hormone response, the expression patterns of Ibbgals were analysed under three different hormone treatments. After the uniconazole treatment, the expressions of eight Ibbgal genes (including Ibbgal3 , Ibbgal6 , Ibbgal9-12 , Ibbgal16 and Ibbgal17 ) were induced to varying degrees in the leaves and roots of these two cultivars (Fig. 5). Interestingly, Ibbgal4 and Ibbgal8 expression were up-regulated in cv. Jishu25, whereas down-regulated in cv. Jishu29 after the uniconazole treatment, indicating that sweetpotato same bgal genes could respond to uniconazole treatment differently in the different genotypes. After the GA 3 treatment, the accumulation of four Ibbgals (including Ibbgal4 , Ibbgal6, Ibbgal11 , and Ibbgal12 ) were unregulated, while Ibbgal5 was down-regulated in two cultivars (Fig. 5). Among these Ibbgals , Ibbgal4 was the most up-regulated gene, whereas Ibbgal12 was the least up-regulated gene. In addition, GA 3 treatment increased the expression of Ibbgal5 and Ibbgal10 in cv. Jishu29, but decreased the expression in cv. Jishu25. For the ABA treatment, most Ibbgal transcripts were induced in the leaves of these two cultivars (Fig. 5). In the roots, most Ibbgal transcripts were up-regulated under the stress, excepte for Ibbgal1 and Ibbgal15 . Among the up-regulated genes, Ibbgal4 was significantly induced in cv . Jishu25, while it was only slightly up-regulated in cv . Jishu29. These data indicated that sweetpotato bgal genes might play pivotal roles in hormone-response pathways. Discussion β-galactosidase participates in cell wall biogenesis and modification during plant growth [15, 17]. In this study, 17 β-galactosidase cDNAs were isolated from sweetpotato, which have the same number of β-galactosidases as in Arabidopsis , tomato and peach [29, 17]. All Ibbgals except Ibbgal13 had the active site consensus sequences GGP[LIVM]xQxENE[FY]. Most Ibbgal members contained a Gal-lectin domain at the C-terminus, which might be responsible for substrate specificity of bgals [11, 29]. In addition, most Ibbgals were predicted to have signal peptides in the N-terminus, which might be involved in cell wall-related biological processes [29]. The phylogenetic tree was constructed using the bgal proteins from sweetpotato and Arabidopsis , which was similar to those of tomato and rice [29, 13]. This result implied that the bgals in the same branch might have similar and distinct functions, and bgal diversification might occur in the early stage of plant evolution. Ibbgal4 and Atbgal1 of groups A shared the same clade, suggesting that they might have similar functions. In a previous study, Esteban et al . (2005) found that bgal genes participate in the development of vegetative organs in Cicer arietinum [3]. Atbgal genes were reported to have differential tissue-specific expression patterns [11]. Similarly, the expression patterns of Ibbgal s were distinct in different tissues of sweetpotato in this study. Most Ibbgal genes were expressed in all tissues, whereas Ibbgal14 , Ibbgal15 and Ibbgal16 had low expression levels in five tissues. The results are consistent with the observations in Arabidopsis reported by Gantulga et al . (2009) [30]. A number of cis-elements related to development, such as GCN4_motif, TATA box and RY-element, were found in the promoter of Ibbgal genes [31, 32], suggesting that these genes might be related to the development of sweetpotato. Ibbgal2-4 , Ibbgal6 , Ibbgal10 , Ibbgal12 and Ibbgal17 were highly expressed in the early stages of root development. Previous reports have shown that Atbgal5 is involved in root elongation through modifying the cell wall [21, 33]. Lovas et al . (2003) found that Stubgal83 might participate in root and tuber development by altering the metabolic sugar status of the leaves [34]. Thus, we deduced that Ibbgal s might be associated with root development by modifying the cell wall and carbohydrate metabolism. Further study is needed to investigate the function of Ibbgal genes during root development in sweetpotato. To date, increasing evidence manifests that bgal genes are involved in response to various hormonal, biotic and abiotic stresses. PaGAL3 and PaGAL4 trancripts in avocado fruit were found to be inhibited by ethylene and ripening signals [26]. In plant coleoptile tissues, auxin-induced increase of elongation rate is closely associated with the β-galactosidase activity [35, 3]. Li et al. (2003) reported that the β-galactosidase genes in calamander were down-regulated through IAA, JA and ethylene after infection by fungus C . acutatum of citrus flower [36]. Our study showed that the upstream region of all Ibbgal s contained three to seven cis-elements related to phytohormone responses, such as GARE, ERE, AuxRE, CATATGGMSAUR. GARE and PYRIMIDINEBOXHVEPB1, which are involved in plant hormone responses [37, 38]. In this study, the expression of eight Ibbgal genes was significantly up-regulated by the uniconazole treatment. Meanwhile, the majority of the Ibbgal genes were regulated by the GA 3 treatment in leaves and stems of these two cultivars. ABA is a requisite factor in response to stress, senescence, and fruit development [39, 40]. We found that most Ibbgal genes were induced under ABA treatment. These results revealed that Ibbgal genes mignt play important roles in phytohormone responses. Spadoni et al . (2014) found that the expression levels of bgal genes decrease in peach fruit after hot water treatment [25]. Several bgal genes are regulated by abiotic and biotic stresses in A. thaliana and Brassica campestris [23, 12, 41]. In addition, the cis-elements related to stress responses, such as MYC-like, LRT, W-BOX, MBS and ACGT-motif, have been found in the promoter region of ibbgal genes, which might regulate gene expression during biotic and abiotic stresses [42, 43]. Similarly, our result showed that most Ibbgal transcripts were related to salt stress, drought stress, ABA treatment and pathogen infection. For example, the expression of all Ibbgal4 was greatly up-regulated by salt and ABA treatments in the leaves of sweetpotato. Taken together, these Ibbgal genes play essential functions in biotic and abiotic stress responses and their related signal transduction pathways. In particular, Ibbgal s exhibited different stress and hormone response patterns between leaves and roots, and have distinct expression profiles in the two cultivars. There are different in root pectin content from sweetpotato cultivars. β-galactosidase functions in the degradation of galactan side chains of pectin leading to cell wall loosening and softening [44, 45], suggesting that β-galactosidase may be involved in the regulation of the pectin content, and different bgal-mediated pathways might be activated in the storage root development. In respond to stresses, the accumulated sugar has been reported to involve in osmotic adjustments to sustain cell structure and photosynthesis in plant [46, 47]. Pandy et al . (2017) found that loss of sugar was the key regulator for activation of the cell wall hydrolase during senescence [48]. β-galactosidase under abiotic and biotic stresses might be induce the initial structural modification of cell wall and activated to degrade cell wall polysaccharides for producing sugar. Therefore, Ibbgal genes were mainly up-regulated expressed under abiotic and biotic stresses. Further studies need to be performed to investigate the functions of bgals on the stress-response system in sweetpotato. Conclusion We characterized 17 Ibbgal genes and then analyzed their motif compositions and N-glycosylation site. Based on the phylogenetic analysis, the bgals were divided into six subgroups. We also investigated their promoter regions and sub-cellular location. In addition, we systematically investigated the expression profiles in different tissues, and different development stages of storage roots, as well as the expression of the bgals under six different environmental treatments. The diversification of the bgal genes provides a solid foundation for further elaborating the bgal-mediated stress-response system in sweetpotato. Methods Identification and isolation of Ibbgals in sweetpotato To identify Ibbgal genes, we performed local BLAST and domain search for genes containing the conserved domain of bgals in two transcriptase databases (SRP068179 and CRA000288). The obtained transcript sequences were translated and analyzed by the PFAM program (http://pfam.xfam.org) to examine the presence of the bgal conserved domains. The transcripts encoding proteins which were less than 120 amino acids were removed. The bgal domain was confirmed by analyzing transcripts deduced proteins screened in the NCBI BLAST. If two or more transcripts had the identity of amino acids equal to or higher than 97%, only one of these transcripts was kept in the final list of the genes. Pooled samples including 9 tissues of shoot, leaf, stem, fibrous root, storage toot, flower, salt-treated, drought-treated and ABA-treated plants were collected from two sweetpotato cultivars (Jishu25 and Jishu29). The total RNA was isolated from the pooled sample using TRIzol, and cDNA was synthesized using a reverse transcription Kit (Transgene, China). To isolate the Ibbgal genes, the gene-specific primers were designed used for PCR amplification (Additional file 1: Table S1). The obtained sequences were compared to the corresponding transcripts, and the related protein data are summarized in Table 1. Protein properties, N-glycosylation site and subcellular location of the Ibbgal proteins The molecular weights (MW) and isoelectric points (pI) of Ibbgal genes were analyzed using the ExPasy server (http://web.expasy.org/protparam/)[49]. N-glycosylation site analysis of Ibbgal genes was conducted using the NetNGlyc 1.0 server ( http://www.cbs.dtu.dk/services/NetNGlyc/ )[12]. The WoLF PSORT tools（https://wolfpsort.hgc.jp/）were used to predict the subcellular location of the Ibbgal proteins [50]. Conserved motifs, phylogenetic analysis and promoter region prediction of the Ibbgal proteins The conserved domains were identified by the online program SMART ( http://smart.embl-heidelberg.de/ ). These 17 Ibbgal protein sequences were aligned with the MEME server ( http://meme-suite.org/tools/meme ). The protein sequences of Ibbgal s were aligned, and the phylogenetic tree was constructed using the Neighbor-Joining (NJ) method of MEGA software 7.0 [51]. The bgal protein sequences from different species, including Arabidopsis [29], were obtained based on the description in the literature or downloaded from the Plantgdb database ( http://www.plantgdb.org/ ). The promoter sequences (1.5 kb) of Ibbgal genes was obtained from sweetpotato genomic DNA ( https://ipomoea-genome.org/# ), and then the cis-acting elements were predicted using the PLACE tool (http://www.dna.affrc.go.jp/PLACE/) [52]. Quantitative real-time PCR analysis To investigate the function of 17 Ibbgals in sweetpotato, the expression patterns were analysed in various organs, hormonal treatments, abiotic and biotic stresses using qRT-PCR. The primer sequences of the examined genes were listed in Table S2 (Additional file 2). Total RNA was extracted from the frozen samples by using an RNAprep pure plant kit (TIANGEN, Beijing, China) according to the manufacturer's instructions. qRT-PCR was performed using a Roche LightCycler ® 480II system under the following conditions: 95°C for 15 s, followed by 40 cycles of 95°C for 15 s, 55°C for 15 s and 72°C for 15 s. The Ib-Actin gene was used as an internal reference to evaluate the relative gene expression level. The experiments were conducted for three replicates, and the data were calculated according to the 2 − △△ CT method [53]. Plant materials and stress treatments The seedlings of two sweetpotato cultivars ( cv . Jishu25 and Jishu29) were collected from the Crop Ressearch Institue, Shandong Academy of Agricultural Sciences, China. The uniform seedlings of the two cultivars were grown in the Hoagland solution at 26°C under a photoperiod of 16 h light/8 h dark. When the seedlings had five to six functional leaves and adventitious roots of 8 to 10 cm, these seedlings were subjected to six different stresses, respectively. To study the expression patterns under these stresses, the adventitious roots of seedlings were submerged in the solution containing 150 mM NaCl, 20% PEG 6000, 100 mM ABA, 50 mg/L uniconazole, and 50 mg/L gibberellic acid (GA 3 ) respectively [54]. For black spot pathogen treatment, C. fimbriata conidia was collected after growing in potato dextrose agar (PDA) at 28 °C for 7 days, then were diluted to 1×10 4 spores/mL with sterile water, and then the roots of sweetpotato seedlings were cultivated in the 1×10 4 spores/mL conidia suspension. The treated roots and leaves were collected after 0, 3, 6, 12, 24, and 48 h. To investigate the Ibbgals transcript levels in different tissues, the fifth expanded leaves, lips, stems, fibrous roots and storage roots of the two cultivars were sampled at 125 days after transplanting, and the storage roots were sampled at 40, 55, 70, 95, 110, 125 and 150 days after transplanting in the sweetpotato field. Statistical analysis Statistical analysis was performed using the SPSS software package (v13.0), and the datas were presented as means of three replicates. Differences between means were subjected to ANOVA, and the statistical significance of the difference between means was calculated with Duncan's new multiple ranges test and marked with asterisks at p < 0.05. Abbreviations ABA: abscisic acid; BLAST: Basic Local Alignment Search Tool ; bgal: β-galactosidase; GH35: glycoside hydrolase 35; GA3: gibberellins; IAA: indolyl-3-acetic acid; JA: Jasmonic acid ; MW: Molecular weights; NJ: Neighbor-Joining; pI: Isoelectric points; qRT-PCR: Quantitative reverse transcription polymerase chain reaction Declarations Ethics approval and consent to participate Not applicable Consent for publication Not applicable Availability of data and materials All data generated or analyzed during this study are included in this published article and its supplementary information files. Competing interests The authors declare they have no competing interests. Funding This research was supported by National Key R&D Program of China (2018YFD1000706,2018YFD1000700), Postgraduate Research & Practice Innovation Program of Jiangsu Province (KYCX19_2200), the China Agriculture Research System of sweetpotato (CARS-10-B7), Taishan industry leading talents project（2020-2023）and Shandong agricultural application technology project(2018-2020). Authors' contributions FH designed and performed the experiments and wrote the paper. ZQ and TD performed some experiments and analyzed the data, AL and SD analyzed the data. TX, DM and QW revised the paper. ZL and LZ conceived the experiment. Acknowledgements We thank the Ipomoea Genome Hub project team for sharing the Ipomoea batatas genome annotation data (https://ipomoea-genome.org/). References Letunic I, Bork P. Interactive Tree Of Life v2: online annotation and display of phylogenetic trees made easy. Nucleic Acids Res. 2011; 39:475–8. Smith DL, Gross KC. 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Genome-wide identification, structural and gene expression analysis of the bZIP transcription factor family in sweet potato wild relative Ipomoea trifida . BMC Genet. 2019; 20(1):41. Lescot M, Déhais P, Thijs G, Marchal K, Moreau Y, Rouzé P, et al. PlantCARE, a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences. Nucleic Acids Res. 2002; 30(1):325–7. Livak KJ, Schmittgen TD . Analysis of relative gene expression data using real-time quantitative PCR and the 2 −ΔΔCT method. Methods . 2001; 25(4): 402-8. Yang Z, Zhu P, Kang H, Liu L, Cao Q, Sun J, et al. High-throughput deep sequencing reveals the important role that microRNAs play in the salt response in sweet potato (Ipomoea batatas L.). BMC Genomics. 2020; 21(1):164. Tables Table 1 Gene and protein analysis of bgals in Ipomoea batatas. Gene name CDS a Length(aa) b MW(kDa) c pI d Subcellular localization Signal peptides e N-glycosylation site f Ibbgal1 2529 842 94.005 5.98 chloroplast + 3 Ibbgal2 2196 731 81.393 8.39 chloroplast + 2 Ibbgal3 2526 841 93.635 7.27 vacuole + 1 Ibbgal 4 2529 842 93.578 8.71 vacuole - 1 Ibbgal 5 2022 673 74.792 6.32 nucleus - 1 Ibbgal 6 2529 842 93.665 7.94 chloroplast + 1 Ibbgal7 2481 826 7.22 9.32 extracellular + 4 Ibbgal8 2541 846 91.829 6.37 vacuole + 2 Ibbgal9 2463 820 92.0858 5.31 vacuole + 2 Ibbgal10 2391 796 89.004 6.83 nucleus - 4 Ibbgal11 2505 834 94.335 8.57 chloroplast + 5 Ibbgal12 2187 728 80.867 9.13 vacuole + 2 Ibbgal13 3333 1110 125.149 5.5 chloroplast - 6 Ibbgal14 2487 828 93.578 8.71 vacuole + 5 Ibbgal15 2475 824 93.72 8.58 chloroplast + 5 Ibbgal16 2412 803 89.731 6.34 chloroplast + 6 Ibbgal17 2145 714 79.382 7.99 chloroplast - 2 a The length of Ibbgals coding sequence b The length of Ibbgals protein. c Molecular weight d Theoretical isoelectric point e “+ ” means contain signal peptide, “_” means lack signal peptide. f Predicted using NetNGlyc. Table 2. The putative cis-elements in the promoters of 17 Ibbgal s genes. gene Plant hormone response elements Stress response elements Light response elements Other elements Ibbgal 1 ABRE 4 , AuxRE 2 , GARE 2 , TATC-BOX, PYRIMIDINEBOXHVEPB1 box-W 2 , MYC-like 18 , ACGT 10 INR 8 , GT1-motif 5 , Box 4 8 , IBOX 5 , GBOX 3 , GATAbox 10 , GAG-motif, TCT-motif 3 , Box II EEs, TATA-box 21 , GT 15 , CCAAT-box 3 , AAGAA-motif Ibbgal 2 GARE 4 , TGACG-motif2, DPBFCOREDCDC3 2 , CATATGGMSAUR 4 MBS 2 , MYC-like 18 , ACGT 2 INR 3 , IBOX 2 , GATAbox 14 ,GAG-motif,TBOX 2 , TCT-motif 2 ,AT1-motif Circadian 2 , TATA-box 18 , CCAAT-box 9 , GCN4-motif, RY-element 4 , GT 12 Ibbgal 3 ABRE,ERE, DPBFCOREDCDC3 3 , MYC-like 16 , ACGT 2 INR 2 , GT1-motif, IBOX 6 , DRE 2 , GATAbox 15 , GAG-motif, TBOX 3 , TCT-motif, Box II 2 Circadian, TATA-box 17 , CCAAT-box 6 , RY-element 2 , GT 12 Ibbgal 4 ABRE 5 , GARE, AuxRE 2 , PYRIMIDINEBOXHVEPB1 box-W, MYC-like1 8 , ACGT 10 INR8, GT1-motif5, Box 4 8 , IBOX5, GATAbox10, GAG-motif, TCT-motif 3 , Box II EEs,TATA-box21,CCAAT-box3,GT 15 , AAGAA-motif Ibbgal 5 ABRE 3 , ERE, GARE, CGTCA-motif 2 , TGACG-motif4, DPBFCOREDCDC3 4 , PYRIMIDINEBOXHVEPB1 LRT, box-W, MYC-like1 2 , ACGT 8 , MBS 3 ,GT1 8 INR6, GT1-motif2, Box 4 3 , IBOX3, GATAbox15, Box A, TBOX,TCT-motif2, Box II2 Circadian3, TATA-box15, CCAAT-box6, Box A , Ibbgal 6 ABRE 2 , ERE, GARE 2 , CGTCA-motif 2 , TGACG-motif4, DRE2COREZMRAB17, PYRIMIDINEBOXHVEPB1 LRT 3 , MYC-like 10 , ACGT 12 INR 4 , GT1-motif, Box 4, IBOX 8 , GATAbox 22 , TBOX, TCT-motif 5 , Box II 4 TATA-box 21 , CCAAT-box 4 , RY-element, GT 13 Ibbgal 7 ERE, GARE 2 , AuxRE, CGTCA-motif, TGACG-motif 3 , DPBFCOREDCDC3 2 , CATATGGMSAUR 2 MYC-like 14 , ACGT 4 , GT-1 5 INR 4 , Box 4 2 , IBOX14, GATAbox 17 Circadian 4 , TATA-box 17 , CCAAT-box 9 , RY-element2 Ibbgal 8 ABRE 3 , ERE, GARE, DPBFCOREDCDC3 4 , CATATGGMSAUR 4 LRT 2 , MYC-like 20 , DRE 2 , ACGT 12 , MBS2 ,GT-1 9 INR 3 , GT1-motif, Box 4 4 , IBOX 8 , GATAbox 18 , TCT-motif 3 , Box II 3 Circadian 2 , TATA-box 20 , CCAAT-box 3 , RY-element Ibbgal 9 ABRE, ERE, GARE 2 LRT 3 , MYC-like 8 , ACGT 6 , GT-1 5 INR 3 , GT1-motif, Box 4 2 , IBOX 13 ,GATAbox 22 , Tbox 2 , Box II 3 Circadian 5 , EEs, TATA-box 28 , CCAAT-box 3 ,GCN4-motif, RY-element 4 Ibbgal10 ABRE 2 ,GARE,DPBFCOREDCDC3, CATATGGMSAUR 2 ,PYRIMIDINEBOXHVEPB1 box-W, MYC-like 18 , ACGT 12 , MBS 3 , GT-1 2 INR 2 , Box 4 3 , IBOX 7 TATA-box 16 , CCAAT-box 3 , RY-element 3 , Box A 2 Ibbgal11 GARE 3 ,CATATGGMSAUR 2 , PYRIMIDINEBOXHVEPB1 MYC-like 8 , ACGT 4 , MBS 2 , GT-1 2 INR 5 , GT1-motif, Box 4 3 , IBOX 7 , GATAbox 18 , GAG-motif, TBOX 2 , TCT-motif, Box II Circadian,TATA-box 23 , CCAAT-box 4 ,AAGAA-motif, RY-element 2 Ibbgal12 ABRE 3 , ERE, GARE 4 , TGACG-motif, PYRIMIDINEBOXHVEPB1 LRT 3 , box-W, MYC-like 18 , DRE 4 , ACGT 8 ,GT-1 8 INR 8 ，GT1-motif, Box 43, IBOX3, GATAbox21, TCT-motif, Box II 2 Circadian 2 , TATA-box 27 , CCAAT-box 3 ,RY-element Ibbgal13 ABRE 3 , ERE, TGACG-motif, DPBFCOREDCDC3 LRT 2 , MYC-like 18 , ACGT 6 , MBS 2 , GT-1 4 INR 4 , GT1-motif 3 , IBOX 15 , GATAbox 15 , GAG-motif, TBOX, Box II 3 Circadian, TATA-box 12 , CCAAT-box 4 , RY-element Ibbgal14 ABRE 3 , ERE, GARE, TGACG-motif, DPBFCOREDCDC3 2 , CATATGGMSAUR 4 LRT 4 , box-W, MYC-like 14 , ACGT 6 , MBS,GT-1 3 INR 3 , GT1-motif 2 , Box 4, IBOX 10 , GATAbox 18 ,CATT, TBOX 3 , Box II 3 Circadian, TATA-box 13 , CCAAT-box 6 , RY-element 3 Ibbgal15 GARE 2 , DPBFCOREDCDC3 2 LRT 3 , box-W 2 , MYC-like 28 , GT-1 2 INR 4 , GT1-motif 2 , IBOX 3 , GATAbox 10 , TBOX 2 , TCT-motif, Box II Circadian, TATA-box 2 , CCAAT-box 5 , RY-element Ibbgal16 ERE, GARE 2 , DPBFCOREDCDC3 3 , CATATGGMSAUR 2 LRT 2 , box-W, MYC-like 8 , DRE 3 , GT-1 6 INR 4 , Box 4 5 , IBOX 2 , GATAbox 13 , GAG-motif, TBOX, TCT-motif TATA-box 36 , CCAAT-box 3 , RY-element Ibbgal17 ABRE 7 , ERE, GARE 3 , TGACG-motif 4 , DPBFCOREDCDC3 6 , CATATGGMSAUR 2 , GCCCORE LRT 2 , box-W 3 , MYC-like 10 , ACGT 6 , MBS 2 , GT-1 INR 2 , GT1-motif, Box 4, IBOX 9 , GATAbox 24 , TBOX, Box II TATA-box 18 , CCAAT-box 4 , GCN4-motif, RY-element 4 Superscript numbers represent the repeats (2 or more than 2) of each cis-element in the Ibbgal promoter, while the others only contain one copy of corresponding cis-element. ABRE and ACGT, cis-acting elements involved in the abscisic acid responsiveness;AuxRE, cis-acting regulatory element involved in auxin responsiveness; AAGAA-motif, cis-element involved in secondary xylem development; Box A, cis-acting elements of phenylalanine ammonia-lyase; Box II, part of a light responsive element; Box-W, fungal elicitor responsive element; Box 4, part of a conserved DNA module involved in light responsiveness; CATATGGMSAUR , cis-acting element involved in auxin responsiveness; CCAAT-box, MYBHv1 binding site; Circadian, cis-acting regulatory element involved in circadian control; DPBFCOREDCDC3, induced by ABA; DRE, cis-acting element involved in drought response;EEs, part of evening and circadian response; ERE, ethylene-responsive element; GARE, gibberellin-responsive element; GATA-motif,part of a light responsive element; Gbox, cis-acting regulatory element involved in light responsiveness; GATAbox, part of a light responsive element; GAG-motif, part of a light responsive element; GCCCORE , cis-acting element involved in jasmonate responsiveness; GCN4-motif, cis-regulatory element involved in endosperm; GT1-motif, light responsive element; GT-1, cis-acting element involved in the salt stress; INR, part of a light responsive element; IBOX, part of a light responsive element; LTR, cis-acting element involved in low-temperature responsiveness; MBS, MYB binding site involved in drought-inducibility; MYC-like , cis-acting elements of drought-responsive; PYRIMIDINEBOXHVEPB1, cis- and trans-acting elements involved in gibberellins and abscisic acid responsiveness; RY-element, cis-acting regulatory element involved in seedspecific regulation; TATA-box, core promoter element around −30 of transcription start; TATC-box, cis-acting element involved in gibberellin-responsiveness;TBOX, part of a light responsive element; TCT-motif, part of a light responsive element; TGACG-motif, cis-acting regulatory element involved in the MeJA-responsiveness. 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The unrooted tree was generated using MEGA7.0 by the NJ method, and the bootstrap test was set to 1000 replicates. B: Motif distribution in Ibbgal genes. The upper part represents the composition and position of motifs of Ibbgals with six motifs shown in distinct colors. The lower part shows the motifs of Ibbgals with the symbol of each residue.\n","description":"","filename":"Fig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/feee9e4c00a4d05f9fe43591.png"},{"id":4163466,"identity":"fb954006-b1c5-4fa2-a738-d9fc92bd9a7d","added_by":"auto","created_at":"2020-12-10 16:36:01","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":552623,"visible":true,"origin":"","legend":"Phylogenetic tree of bgal proteins in sweetpotato, and Arabidopsis.\nThe bgal protein sequences of Arabidopsis were downloaded from the database of Arabidopsis from the NCBI database (https://www.ncbi.nlm.nih.gov/).The tree was classified into 7 different subfamilies indicated by outer rings with blue color.\n","description":"","filename":"Fig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/c6f2a18bb451ca57379406f1.png"},{"id":4163467,"identity":"9d939ba2-6624-4ab4-936f-ec6242efd50a","added_by":"auto","created_at":"2020-12-10 16:36:02","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":644617,"visible":true,"origin":"","legend":"Relative mRNA expression levels of Ibbgal genes in tissues and storage root development of two sweetpotato varieties.\nA. Tissues including leaf, tip, young-stem, old-stem, fibrous root and storage root. B. Expression profiles in the storage root development .Gene expression were detected by quantitative real-time polymerase chain reaction. Log-transformed fold-change data were used for creating the heatmaps by TBtools. The coloured scale varying from blue to red indicates relatively low or high expression.\n","description":"","filename":"Fig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/19255d3f2ee8ab2faeedf3fa.png"},{"id":4163468,"identity":"2bccc9b5-0a5c-4cef-9247-e25d3423a629","added_by":"auto","created_at":"2020-12-10 16:36:02","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":776084,"visible":true,"origin":"","legend":"Expression patterns of Ibbgal genes under salt, drought stress and balck spot pathogen infection in the leave and root of the two cultivars. \nThe y-axis represents relative expression. Bars represent the mean of three biological replicates ± SE.The asteridk indicated that the expression level between the treatment times is significantly different (P\u003c0.05).","description":"","filename":"FIg4.png","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/e549a88b57cf6502608a63f6.png"},{"id":4163469,"identity":"04b163db-3842-4238-91e7-ed79c8ff451d","added_by":"auto","created_at":"2020-12-10 16:36:02","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":457702,"visible":true,"origin":"","legend":"Expression profiles of Ibbgal genes after uniconazole, GA3 and ABA treatment in the leave and root of the two cultivars. \nGene expression was detected by quantitative real-time polymerase chain reaction. Log-transformed fold-change data were used for creating the heatmaps by TBtools. The coloured scale varies from blue to red, which indicates the low or high expression of each gene.\n","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/60ed89fa25b682f61cd7c5bf.png"},{"id":15670230,"identity":"89e88023-5418-497e-8d69-d769b4a4d9be","added_by":"auto","created_at":"2021-11-18 13:57:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1931781,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/6edb8499-10e4-4b5a-ae0f-516f83f64162.pdf"},{"id":4163463,"identity":"42c1bb50-1d05-4ad7-8025-6c4b89007295","added_by":"auto","created_at":"2020-12-10 16:36:01","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":13948,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile1.docx","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/7bf5cc4a28898cde24ecee68.docx"},{"id":4163465,"identity":"ce221c27-9699-41cf-aad7-e222cdf9cb8f","added_by":"auto","created_at":"2020-12-10 16:36:01","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":14366,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile2.docx","url":"https://assets-eu.researchsquare.com/files/rs-32133/v3/779f685f202fbffcfac1a1e0.docx"}],"financialInterests":"","formattedTitle":"\u003cp\u003eGenome-wide in silico identification and expression analysis of beta-galactosidase family members in sweetpotato [\u003cem\u003eIpomoea batatas\u003c/em\u003e (L.) Lam.]\u003c/p\u003e","fulltext":[{"header":"Background","content":"\u003cp\u003e\u0026beta;-galactosidases (EC 3.2.1.23; bgal) widely exist in higher plants. \u0026nbsp; Plant \u0026beta;-galactosidase belongs to the glycoside hydrolase 35 (GH35) families [1], which catalyzes the removal of terminal galactosyl residues from carbohydrates, glycoproteins and galactolipids [2, 3]. In plants, \u0026beta;-galactosidase has been reported to degrade structural polysaccharides in plant cell walls to release free galactose during a variety of biological processes, including cell wall expansion and degradation, metabolic recycling of galactolipids and glycoproteins, and turnover of signaling molecules during ripening [4, 5].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn higher plants, bgals have been grouped into two classes based on their substrate preference [6]. Enzymes in the first class prefer pectic \u0026beta;-(1\u0026rarr;4)-galactan as the substrate, and enzymes in the other prefer the \u0026beta;-(1\u0026rarr;3) and (1\u0026rarr;6)-galactan backbones of arabinogalactan proteins [7, 8]. A typical bgal protein contains the GH35 conserved site in the N-terminal region [9]. Like other glycosidase families, bgal genes are ubiquitously expressed in many plants, such as tomato [2], papaya [10], \u003cem\u003eArabidopsis\u003c/em\u003e [11], \u003cem\u003eBrassica campestris\u0026nbsp;\u003c/em\u003e[12] and rice [13].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003ePlant \u003cem\u003ebgal\u0026nbsp;\u003c/em\u003egenes are widely involved in the modification of the architecture of cell walls and intercellular attachments [14, 15]. \u003cem\u003ebgal\u0026nbsp;\u003c/em\u003egenes also respond to plant growth and development including fruit development and ripening [16, 17], seed germination [18, 19], and root development [20, 21]. In most fruits, \u003cem\u003ebgal\u0026nbsp;\u003c/em\u003egenes exhibit differential expression patterns during flowering and fruit development [16, 12]. In \u003cem\u003eCicer arietinum\u003c/em\u003e, \u0026nbsp;\u003cem\u003eCanbgal-5\u003c/em\u003e expression is\u0026nbsp;\u0026nbsp;\u003ca href=\"javascript%3A;\"\u003erelevant\u003c/a\u003e\u0026nbsp; to young and meristematic stages with a high cell division rate, while\u0026nbsp;\u003cem\u003eCanBGal-1\u0026nbsp;\u003c/em\u003eand\u003cem\u003e\u0026nbsp;CanBGal-4\u0026nbsp;\u003c/em\u003eare strongly related to later stages of epicotyl growth [3]. In addition, \u003cem\u003ebgal\u0026nbsp;\u003c/em\u003egenes can be regulated by abiotic and biotic stresses [22]. For example, \u003cem\u003eAtbgal1\u003c/em\u003e was reported to be induced by salt stress or pathogen attack [23]. Likewise, the transcription level of \u0026beta;-galactosidase in cowpea is reduced under salt treatments [24], and the \u003cem\u003ebgal\u003c/em\u003e mRNA level in peach\u003cem\u003e\u0026nbsp;\u003c/em\u003eis highly suppressed by water stress [25]. In addition, \u003cem\u003ebgal\u0026nbsp;\u003c/em\u003egenes have been found to play a role \u0026nbsp;in a variety of biological processes through ethylene signal transduction [26, 11]. However, the function of \u003cem\u003ebgal\u003c/em\u003e has not been studied in sweetpotato (\u003cem\u003eIpomoea batatas\u003c/em\u003e (L.) Lam).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSweetpotato is an important food crop which is widely grown in tropical and subtropical areas, especially in Asia and sub-Saharan Africa. Due to its outcrossing hexaploidy (2n=6\u0026times;=90), the genomic research in sweetpotato is very complicated [27, 28]. So far, no high-quality genome sequence of sweetpotato has been available. Although \u003cem\u003ebgal\u003c/em\u003e genes are widely isolated from many plant species, its function in sweetpotato remains unknown. In the present study, we firstly identified 17 \u003cem\u003ebgal\u003c/em\u003e genes (\u003cem\u003eIbbgal\u003c/em\u003e) in sweetpotato, and then investigated their phylogeny, motif compositions and predicted cis-elements using various bioinformatics tools. In addition, the expression patterns of these 17 \u003cem\u003eIbbgals\u003c/em\u003e in different tissues of two cultivars were investigated under three exogenous hormones, two abiotic and one biotic stress conditions. Our study will lay the foundation for further research on the function of \u003cem\u003ebgal\u003c/em\u003e gene in plants, and provide new insight into different regulatory mechanisms in plant growth through \u003cem\u003ebgal\u003c/em\u003e-mediated responses to environmental stresses in sweetpotato.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eIdentification and characterization of \u003cem\u003eIbbgal\u003c/em\u003e genes in sweetpotato\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eA total of 17 \u003cem\u003eIbbgal\u003c/em\u003e genes were isolated from sweetpotato after local BLAST using the conserved bgal domain. The deduced amino acid sequences of the \u003cem\u003eIbbgal\u003c/em\u003e proteins were used to predict their protein lengths, signal peptides, pI values, molecular weights, sub-cellular localization and the possible N-glycosylation sites (Table 1). Characteristic analysis showed that these 17 Ibbgals were 673 to 1110 aa in length, the predicted MWs and pIs ranged from 74.8 kDa to 125.1 kDa and 5.31 to 6.16, respectively. The predicted localization of most Ibbgals varied and included the chloroplast, vacuole, and nucleus. Only one Ibbgal, Ibbgal7, was found to be located in the extracellular. Signal peptides analysis revealed that all Ibbgals, except for Ibbgal4, Ibbgal5, Ibbgal10, Ibbgal13 and Ibbgal17, contained a signal peptide. The number of N-glycosylation sites varied from 1 to 6, wherein Ibbgal13 and Ibbgal16 contained 6 N-glycosylation sites.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConserved motifs and phylogenetic analysis of the Ibbgal proteins\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn this study, the \u0026beta;-galactosidase active site was found in all Ibbgal proteins. However, all but Ibbgal13 have the active site consensus sequence GGP[LIVM]xQxENE[FY] of the GH35 \u0026beta;-galactosidase family. In addition, all Ibbgal members carried a Gal-lectin domain at the C-terminus of the protein sequence, except for Ibbgal2, Ibbgal5, Ibbgal12, Ibbgal13, and Ibbgal17. Motif analysis showed that motif 1 was found in all Ibbgals except Ibbgal13, and motifs 2-6 were found in all Ibbgals except Ibbgal11 and Ibbgal17 (Fig. 1). A total of 34 \u003cem\u003ebgal\u003c/em\u003e genes from sweetpotato and\u003cem\u003e\u0026nbsp;Arabidopsis\u0026nbsp;\u003c/em\u003ewere classified into seven subgroups, designated as A, B, C, D, E, F and G, using phylogenetic analysis (Fig. 2). Among these\u003cem\u003e\u0026nbsp;groups\u003c/em\u003e, groups A and D were the largest groups with four \u003cem\u003eIbbgal\u0026nbsp;\u003c/em\u003egenes in each. Groups B and E had three \u003cem\u003eIbbgal\u003c/em\u003e genes. However, \u003cem\u003eIbbgal9\u003c/em\u003e, \u003cem\u003eIbbgal17\u003c/em\u003e and \u003cem\u003eIbbgal13\u003c/em\u003e were classified into group C, F and E, respectively.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCis-element prediction of \u003cem\u003eIbbgal\u003c/em\u003e genes\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo understand the potential transcriptional regulatory mechanisms of the \u003cem\u003eIbbgal\u003c/em\u003e genes, the cis-elements of each \u003cem\u003eIbbgal\u003c/em\u003e promoter sequences were predicted and analyzed (Table 2). The promoters of \u003cem\u003eIbbgals\u003c/em\u003e were classified into at least four types of cis-elements, including plant hormone responsive elements, light responsive elements, stress responsive elements, and other elements. Most \u003cem\u003eIbbgal\u003c/em\u003e promoters had the GARE (gibberellin-responsive element), ERE (ethylene-responsive element) cis-elements, AuxRE and CATATGGMSAUR motifs which were involved in plant hormone response. Most \u003cem\u003eIbbgal\u003c/em\u003e promoters, except \u003cem\u003eIbbgal6\u003c/em\u003e, \u003cem\u003eIbbgal16\u003c/em\u003e and \u003cem\u003eIbbgal17\u003c/em\u003e, contained circadian and EE elements participated in circadian regulation. In addition, at least five light response elements were found in each \u003cem\u003eIbbgal\u003c/em\u003e gene, which might be essential for plant growth and development. Interestingly, the \u003cem\u003eIbbgal\u003c/em\u003es contained the MYC-like and ABRE (Abscisic acid response element) cis-elements mediated the responses to abotic stresses.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eExpression profiles of \u003cem\u003eIbbgal\u003c/em\u003e genes in tissues and different root development stages\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo identify the potential functions of \u003cem\u003eIbbgal\u003c/em\u003e genes, we analyzed the transcript levels of \u003cem\u003eIbbgal\u003c/em\u003es in various tissues of \u003cem\u003ecv\u003c/em\u003e. Jishu25 and Jishu29, including leaf, stem lip, stem, fibrous root, and storage root. 47% of \u003cem\u003eIbbgal\u003c/em\u003es had similar expression patterns in five tissues of two cultivars (Fig. 3A). For example, \u003cem\u003eIbbgal4\u003c/em\u003e, \u003cem\u003eIbbgal10\u003c/em\u003e, \u003cem\u003eIbbgal13\u003c/em\u003e and \u003cem\u003eIbbgal17\u003c/em\u003e were highly expressed in five tissues, whereas \u003cem\u003eIbbgal14\u003c/em\u003e,\u003cem\u003e\u0026nbsp;Ibbgal15\u003c/em\u003e and \u003cem\u003eIbbgal16\u0026nbsp;\u003c/em\u003ewere poorly expressed in these tissues. Intriguingly, the expression of \u003cem\u003eIbbgal4\u003c/em\u003e in fibrous root was significantly higher than that of storage root, while \u003cem\u003eIbbgal3\u003c/em\u003e and \u003cem\u003eIbbgal10\u003c/em\u003e were expressed at higher levels in lip than other tissues. However, the transcript of \u003cem\u003eIbbgal17\u003c/em\u003e mRNA in \u003cem\u003ecv\u003c/em\u003e. Jishu25 was prominently higher in storage root than fibrous root, whereas that in \u003cem\u003ecv\u003c/em\u003e. Jishu29 had no significant difference in the roots. Similarly, the expression of \u003cem\u003eIbbgal11\u003c/em\u003e had the opposite pattern in the storage and fibrous roots between \u003cem\u003ecv\u003c/em\u003e. Jishu25 and Jishu29.\u003c/p\u003e\n\u003cp\u003eIn root development stages, 6 (35.3%) \u003cem\u003eIbbgal\u003c/em\u003e transcripts were down-regulated including \u003cem\u003eIbbgal2\u003c/em\u003e, \u003cem\u003eIbbgal3\u003c/em\u003e, \u003cem\u003eIbbgal4\u003c/em\u003e, \u003cem\u003eIbbgal6\u003c/em\u003e, \u003cem\u003eIbbgal10\u003c/em\u003e, and \u003cem\u003eIbbgal16\u003c/em\u003e, whereas 6 \u003cem\u003eIbbgal\u003c/em\u003e transcripts were up-regulated, two \u003cem\u003eIbbgal\u003c/em\u003e genes (\u003cem\u003eIbbgal14\u003c/em\u003e and \u003cem\u003eIbbgal15\u003c/em\u003e) were not detected in root development. It is interesting that the \u003cem\u003eIbbgal11\u003c/em\u003e and \u003cem\u003eIbbgal12\u003c/em\u003e transcripts had the opposite expression pattern between\u003cem\u003e\u0026nbsp;cv.\u003c/em\u003e Jishu25 and Jishu29 (Fig. 3B).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eExpression profiles of\u003cem\u003e\u0026nbsp;Ibbgal\u003c/em\u003e\u0026nbsp; genes in response to abiotic and biotic stresses\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBesides their functions in plant growth and development, \u003cem\u003eIbbgal\u003c/em\u003e genes may also be involved in biotic and abiotic stress responses.\u0026nbsp;For sweetpotato, salinity and drought are the most dominant factors which limit the growth and yield among various abiotic stresses.Under salt stress, all \u003cem\u003eIbbgal\u003c/em\u003e genes were up-regulated in these two cultivars (Fig.4). Some genes had the highest expression levels at 12 h in the leaves, whereas other \u003cem\u003eIbbgal\u003c/em\u003e genes in roots were expressed at a high level at 6 h and 48 h after salt stress. In addition,\u003cem\u003e\u0026nbsp;Ibbgal2\u003c/em\u003e, \u003cem\u003eIbbgal4\u003c/em\u003e, \u003cem\u003eIbbgal5\u003c/em\u003e and \u003cem\u003eIbbgal13\u003c/em\u003e in the leaves were unregulated remarkably by at least 10-fold induction after salt stress. These results indicated that \u003cem\u003eIbbgal\u003c/em\u003e genes were involved in salt stress response in sweetpotato. Under drought stress (Fig. 4), all \u003cem\u003eIbbgal\u003c/em\u003e genes were up-regulated in the leaves and roots of \u003cem\u003ecv\u003c/em\u003e. Jishu29, while \u003cem\u003eIbbgal3\u003c/em\u003e, \u003cem\u003eIbbgal6\u003c/em\u003e, \u003cem\u003eIbbgal10\u003c/em\u003e, and \u003cem\u003eIbbgal17\u003c/em\u003e were down-regulated in the leaves of Jishu25, and \u003cem\u003eIbbgal1\u003c/em\u003e, \u003cem\u003eIbbgal3\u003c/em\u003e and \u003cem\u003eIbbgal16\u003c/em\u003e expression were also reduced in the root of Jishu25. Amongst the up-regulated genes, the expression of \u003cem\u003eIbbgal2,\u003c/em\u003e \u003cem\u003eIbbgal4,\u003c/em\u003e \u003cem\u003eIbbgal8, Ibbgal9\u0026nbsp;\u003c/em\u003eand \u003cem\u003eIbbgal13\u0026nbsp;\u003c/em\u003ereached the peak at 12 h after stress, and \u003cem\u003eIbbgal4\u003c/em\u003e was the most up-regulated gene with at least 81-fold induction in the two cultivars leaves, suggesting that \u003cem\u003eIbbgals\u003c/em\u003e in the different cultivars responded to drought treatment differently.\u0026nbsp;Black spot,\u0026nbsp;caused by\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003cem\u003eCeratocystis fimbriata\u003c/em\u003e(\u003cem\u003eC. fimbriata\u003c/em\u003e),\u0026nbsp;is one of the main diseases in sweetpotato production, which seriously affects the quality and yield of sweetpotato. After the pathogen infection, \u003cem\u003eIbbgal\u003c/em\u003e genes had different expression patterns in the leaves and roots of these two cultivars (Fig. 4). \u003cem\u003eIbbgal5,\u003c/em\u003e \u003cem\u003eIbbgal10\u003c/em\u003e, \u003cem\u003eIbbgal11\u003c/em\u003e and \u003cem\u003eIbbgal16\u003c/em\u003e transcripts were induced by the pathogen infection in these two cultivars. It is worth noting that \u003cem\u003eIbbgal15\u0026nbsp;\u003c/em\u003eexpression in the leaves and roots of \u003cem\u003ecv.\u003c/em\u003e Jishu25 was up-regulated, whereas down-regulated in \u003cem\u003ecv.\u003c/em\u003e Jishu29. Collectively, these results implied that \u003cem\u003eIbbgals\u003c/em\u003e in the different cultivars might have different functions under abiotic and biotic stresses.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eExpression profiles of \u003cem\u003eIbbgal\u003c/em\u003e genes in response to various hormone treatments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo survey the role of \u003cem\u003eIbbgal\u003c/em\u003e genes in plant hormone response, the expression patterns of \u003cem\u003eIbbgals\u003c/em\u003e were analysed under three different hormone treatments. After the uniconazole treatment, the expressions of eight \u003cem\u003eIbbgal\u003c/em\u003e genes (including \u003cem\u003eIbbgal3\u003c/em\u003e, \u003cem\u003eIbbgal6\u003c/em\u003e, \u003cem\u003eIbbgal9-12\u003c/em\u003e, \u003cem\u003eIbbgal16\u003c/em\u003e and \u003cem\u003eIbbgal17\u003c/em\u003e) were induced to varying degrees in the leaves and roots of these two cultivars (Fig. 5). Interestingly, \u003cem\u003eIbbgal4\u003c/em\u003e and \u003cem\u003eIbbgal8\u0026nbsp;\u003c/em\u003eexpression were up-regulated in \u003cem\u003ecv.\u003c/em\u003e Jishu25, whereas down-regulated in cv. Jishu29 after the uniconazole treatment, indicating that sweetpotato same \u003cem\u003ebgal\u003c/em\u003e genes could \u0026nbsp;respond to uniconazole treatment differently in the different genotypes. After the GA\u003csub\u003e3\u003c/sub\u003e treatment, the accumulation of four \u003cem\u003eIbbgals\u003c/em\u003e (including \u003cem\u003eIbbgal4\u003c/em\u003e, \u003cem\u003eIbbgal6,\u003c/em\u003e \u003cem\u003eIbbgal11\u003c/em\u003e, and\u003cem\u003e\u0026nbsp;Ibbgal12\u003c/em\u003e) were unregulated, while \u003cem\u003eIbbgal5\u0026nbsp;\u003c/em\u003ewas down-regulated in two cultivars (Fig. 5). Among these \u003cem\u003eIbbgals\u003c/em\u003e, \u003cem\u003eIbbgal4\u003c/em\u003e was the most up-regulated gene, whereas \u003cem\u003eIbbgal12\u003c/em\u003e was the least up-regulated gene. In addition, GA\u003csub\u003e3\u003c/sub\u003e treatment increased the expression of \u003cem\u003eIbbgal5\u003c/em\u003e and \u003cem\u003eIbbgal10\u003c/em\u003e in \u003cem\u003ecv.\u0026nbsp;\u003c/em\u003eJishu29, but decreased the expression in \u003cem\u003ecv.\u0026nbsp;\u003c/em\u003eJishu25. For the ABA treatment, most \u003cem\u003eIbbgal\u003c/em\u003e transcripts were induced in the leaves of these two cultivars (Fig. 5). In the roots, most\u003cem\u003e\u0026nbsp;Ibbgal\u003c/em\u003e transcripts were up-regulated under the stress, excepte for \u003cem\u003eIbbgal1\u003c/em\u003e and \u003cem\u003eIbbgal15\u003c/em\u003e. Among the up-regulated genes, \u003cem\u003eIbbgal4\u0026nbsp;\u003c/em\u003ewas significantly induced in \u003cem\u003ecv\u003c/em\u003e. Jishu25, while it was only slightly up-regulated in \u003cem\u003ecv\u003c/em\u003e. Jishu29.\u0026nbsp;These data indicated that sweetpotato\u003cem\u003e\u0026nbsp;bgal\u003c/em\u003e genes might play pivotal roles in hormone-response pathways.\u003c/p\u003e"},{"header":"Discussion ","content":"\u003cp\u003e\u0026beta;-galactosidase participates in cell wall biogenesis and modification during plant growth [15, 17]. In this study, 17 \u0026beta;-galactosidase cDNAs were isolated from sweetpotato, which have the same number of \u0026beta;-galactosidases as in \u003cem\u003eArabidopsis\u003c/em\u003e, tomato and peach [29, 17]. All Ibbgals except Ibbgal13 had the active site consensus sequences GGP[LIVM]xQxENE[FY]. Most Ibbgal members contained a Gal-lectin domain at the C-terminus, which might be responsible for substrate specificity of bgals [11, 29]. In addition, most Ibbgals were predicted to have signal peptides in the N-terminus, which might be involved in cell wall-related biological processes [29]. The phylogenetic tree was constructed using the bgal proteins from sweetpotato and \u003cem\u003eArabidopsis\u003c/em\u003e, which was similar to those of tomato and rice [29, 13]. This result implied that the bgals\u0026nbsp;in the same branch might have similar and distinct functions, and bgal diversification might occur in the early stage of plant evolution. \u003cem\u003eIbbgal4\u003c/em\u003e and \u003cem\u003eAtbgal1\u003c/em\u003e of groups A shared the same clade, suggesting that they might have similar functions.\u003c/p\u003e\n\u003cp\u003eIn a previous study, Esteban \u003cem\u003eet al\u003c/em\u003e. (2005) found that \u003cem\u003ebgal\u003c/em\u003e genes participate in the development of vegetative organs in \u003cem\u003eCicer arietinum\u003c/em\u003e [3].\u003cem\u003e\u0026nbsp;Atbgal\u0026nbsp;\u003c/em\u003egenes were reported to have differential tissue-specific expression patterns [11]. Similarly, the expression patterns of \u003cem\u003eIbbgal\u003c/em\u003es were distinct in different tissues of sweetpotato in this study. Most \u003cem\u003eIbbgal\u003c/em\u003e genes were expressed in all tissues, whereas \u003cem\u003eIbbgal14\u003c/em\u003e, \u003cem\u003eIbbgal15\u0026nbsp;\u003c/em\u003eand \u003cem\u003eIbbgal16\u0026nbsp;\u003c/em\u003ehad low expression levels in five tissues. The results are consistent with the observations in Arabidopsis reported by Gantulga\u003cem\u003e\u0026nbsp;et al\u003c/em\u003e. (2009) [30]. A number of cis-elements related to development, such as GCN4_motif, TATA box and RY-element, were found in the promoter of \u003cem\u003eIbbgal\u003c/em\u003e genes [31, 32], suggesting that these genes might be related to the development of sweetpotato. \u003cem\u003eIbbgal2-4\u003c/em\u003e, \u003cem\u003eIbbgal6\u003c/em\u003e, \u003cem\u003eIbbgal10\u003c/em\u003e, \u003cem\u003eIbbgal12\u003c/em\u003e and \u003cem\u003eIbbgal17\u003c/em\u003e were highly expressed in the early stages of root development. Previous reports have shown that \u003cem\u003eAtbgal5\u003c/em\u003e is involved in root elongation through modifying the cell wall [21, 33]. Lovas \u003cem\u003eet al\u003c/em\u003e. (2003) found that \u003cem\u003eStubgal83\u003c/em\u003e might participate in root and tuber development by altering the metabolic sugar status of the leaves [34]. Thus, we deduced that \u003cem\u003eIbbgal\u003c/em\u003es might be associated with root development by modifying the cell wall and carbohydrate metabolism. Further study is needed to investigate the function of \u003cem\u003eIbbgal\u003c/em\u003e genes during root development in sweetpotato.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp;To date, increasing evidence manifests that \u003cem\u003ebgal\u003c/em\u003e genes are involved in response to various hormonal, biotic and abiotic stresses. \u003cem\u003ePaGAL3\u0026nbsp;\u003c/em\u003eand \u003cem\u003ePaGAL4\u003c/em\u003e trancripts in avocado fruit were found to be inhibited by ethylene and ripening signals [26]. In plant coleoptile tissues, auxin-induced increase of elongation rate is closely associated with the \u0026beta;-galactosidase activity [35, 3]. Li \u003cem\u003eet al.\u003c/em\u003e (2003) reported that the \u0026beta;-galactosidase genes in calamander were down-regulated through IAA, JA and ethylene after infection by fungus C\u003cem\u003e. acutatum\u003c/em\u003e of citrus flower [36]. Our study showed that the upstream region of all \u003cem\u003eIbbgal\u003c/em\u003es contained three to seven cis-elements related to phytohormone responses, such as GARE, ERE, AuxRE, CATATGGMSAUR. GARE and PYRIMIDINEBOXHVEPB1, which are involved in plant hormone responses [37, 38]. In this study, the expression of eight \u003cem\u003eIbbgal\u0026nbsp;\u003c/em\u003egenes was significantly up-regulated by the uniconazole treatment. Meanwhile, the majority of the \u003cem\u003eIbbgal\u003c/em\u003e genes were regulated by the GA\u003csub\u003e3\u003c/sub\u003e treatment in leaves and stems of these two cultivars. ABA is a requisite factor in response to stress, senescence, and fruit development [39, 40]. We found that most \u003cem\u003eIbbgal\u003c/em\u003e genes were induced under ABA treatment. These results revealed that \u003cem\u003eIbbgal\u003c/em\u003e genes mignt play important roles in phytohormone responses. Spadoni \u003cem\u003eet al\u003c/em\u003e. (2014) found that the expression levels of \u003cem\u003ebgal\u003c/em\u003e genes decrease in peach fruit after hot water treatment [25]. Several \u003cem\u003ebgal\u003c/em\u003e genes are regulated by abiotic and biotic stresses in \u003cem\u003eA. thaliana\u003c/em\u003e and \u003cem\u003eBrassica campestris\u0026nbsp;\u003c/em\u003e[23, 12, 41]. In addition, the cis-elements related to stress responses, such as MYC-like, LRT, W-BOX, MBS and ACGT-motif, have been found in the promoter region of\u003cem\u003e\u0026nbsp;ibbgal\u003c/em\u003e genes, which might regulate gene expression during biotic and abiotic stresses [42, 43]. \u0026nbsp;Similarly, our result showed that most \u003cem\u003eIbbgal\u003c/em\u003e transcripts were related to salt stress, drought stress, ABA treatment and pathogen infection. For example, the expression of all \u003cem\u003eIbbgal4\u003c/em\u003e was greatly up-regulated by salt and ABA treatments in the leaves of sweetpotato. Taken together, these \u003cem\u003eIbbgal\u003c/em\u003e genes play essential functions in biotic and abiotic stress responses and their related signal transduction pathways.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp; \u0026nbsp;In particular, \u003cem\u003eIbbgal\u003c/em\u003es exhibited different stress and hormone response patterns between leaves and roots, and have distinct expression profiles in the two cultivars. There are different in root pectin content from sweetpotato cultivars. \u0026beta;-galactosidase functions in the degradation of galactan side chains of pectin leading to cell wall loosening and softening [44, 45], suggesting that \u0026beta;-galactosidase may be involved in the regulation of the pectin content, and different bgal-mediated pathways might be activated in the storage root development. In respond to stresses,\u0026nbsp;the accumulated sugar has been reported to involve in osmotic adjustments to\u0026nbsp;sustain cell structure and photosynthesis\u0026nbsp;in plant [46, 47]. Pandy \u003cem\u003eet al\u003c/em\u003e. (2017) found that loss of sugar was the key regulator for activation of the cell wall hydrolase during senescence [48].\u0026nbsp;\u0026beta;-galactosidase\u0026nbsp;under abiotic and biotic stresses\u0026nbsp;might be induce the initial structural modification of cell wall and\u0026nbsp;activated to\u0026nbsp;degrade\u0026nbsp;cell wall polysaccharides\u0026nbsp;for producing sugar.\u0026nbsp;Therefore, \u003cem\u003eIbbgal\u003c/em\u003e genes were mainly up-regulated expressed under abiotic and biotic stresses. Further studies need to be performed to investigate the functions of \u0026nbsp;bgals on the stress-response system in sweetpotato.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eWe characterized 17 \u003cem\u003eIbbgal\u003c/em\u003e genes and then analyzed their motif compositions and N-glycosylation site. Based on the phylogenetic analysis, the bgals were divided into six subgroups. We also investigated their promoter regions and sub-cellular location. In addition, we systematically investigated the expression profiles in different tissues, and different development stages of storage roots, as well as the expression of the bgals under six different environmental treatments. The diversification of the bgal genes provides a solid foundation for further elaborating the bgal-mediated stress-response system in sweetpotato.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e\u003cstrong\u003eIdentification and isolation of \u003cem\u003eIbbgals\u003c/em\u003e in sweetpotato\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo identify \u003cem\u003eIbbgal\u003c/em\u003e genes, we performed local BLAST and domain search for genes containing the conserved domain of bgals in two transcriptase databases (SRP068179 and CRA000288). The obtained transcript sequences were translated and analyzed by the PFAM program (http://pfam.xfam.org) to examine the presence of the bgal conserved domains. The transcripts encoding proteins which were less than 120 amino acids were removed. The bgal domain was confirmed by analyzing transcripts deduced proteins screened in the NCBI BLAST. If two or more transcripts had the identity of amino acids equal to or higher than 97%, only one of these transcripts was kept in the final list of the genes. Pooled samples including 9 tissues of shoot, leaf, stem, fibrous root, storage toot, flower, salt-treated, drought-treated and ABA-treated plants were collected from two sweetpotato cultivars (Jishu25 and Jishu29). The total RNA was isolated from the pooled sample using TRIzol, and cDNA was synthesized using a reverse transcription Kit (Transgene, China). To isolate the \u003cem\u003eIbbgal\u003c/em\u003e genes, the gene-specific primers were designed used for PCR amplification (Additional file 1: Table S1). The obtained sequences were compared to the corresponding transcripts, and the related protein data are summarized in Table 1.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eProtein properties, N-glycosylation site and subcellular location of the \u003cem\u003eIbbgal\u0026nbsp;\u003c/em\u003eproteins\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe molecular weights (MW) and isoelectric points (pI) of \u003cem\u003eIbbgal\u003c/em\u003e genes were analyzed using the ExPasy server (http://web.expasy.org/protparam/)[49]. N-glycosylation site analysis of \u003cem\u003eIbbgal\u003c/em\u003e genes was conducted using the NetNGlyc 1.0 server (\u003ca href=\"http://www.cbs.dtu.dk/services/NetNGlyc/\"\u003ehttp://www.cbs.dtu.dk/services/NetNGlyc/\u003c/a\u003e)[12]. The WoLF PSORT tools（https://wolfpsort.hgc.jp/）were used to predict the subcellular location of the Ibbgal proteins [50].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConserved motifs, phylogenetic analysis and promoter region prediction of the Ibbgal proteins\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe conserved domains were identified by the online program SMART (\u003ca href=\"http://smart.embl-heidelberg.de/\"\u003ehttp://smart.embl-heidelberg.de/\u003c/a\u003e). These 17 Ibbgal protein sequences were aligned with the MEME server (\u003ca href=\"http://meme-suite.org/tools/meme\"\u003ehttp://meme-suite.org/tools/meme\u003c/a\u003e). The protein sequences of \u003cem\u003eIbbgal\u003c/em\u003es were aligned, and the phylogenetic tree was constructed using the Neighbor-Joining (NJ) method of MEGA software 7.0 [51]. The bgal protein sequences from different species, including \u003cem\u003eArabidopsis\u003c/em\u003e [29], were obtained based on the description in the literature or downloaded from the Plantgdb database (\u003ca href=\"http://www.plantgdb.org/\"\u003ehttp://www.plantgdb.org/\u003c/a\u003e). The promoter sequences (1.5 kb) of \u003cem\u003eIbbgal\u003c/em\u003e genes was obtained from sweetpotato genomic DNA (\u003ca href=\"https://ipomoea-genome.org/\"\u003ehttps://ipomoea-genome.org/#\u003c/a\u003e), and then the cis-acting elements were predicted using the PLACE tool (http://www.dna.affrc.go.jp/PLACE/) [52].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eQuantitative real-time PCR analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo investigate the function \u0026nbsp;of 17 Ibbgals in sweetpotato, the expression patterns were analysed in various organs, hormonal treatments, abiotic and biotic stresses using qRT-PCR. The primer sequences of the examined genes were listed in Table S2 (Additional file 2). Total RNA was extracted from the frozen samples by using an RNAprep pure plant kit (TIANGEN, Beijing, China) according to the manufacturer\u0026apos;s instructions. qRT-PCR was performed using a Roche LightCycler\u003csup\u003e\u0026reg;\u0026nbsp;\u003c/sup\u003e480II system under the following conditions: 95\u0026deg;C for 15 s, followed by 40 cycles of 95\u0026deg;C for 15 s, 55\u0026deg;C for 15 s and 72\u0026deg;C for 15 s. The \u003cem\u003eIb-Actin\u003c/em\u003e gene was used as an internal reference to evaluate the relative gene expression level. The experiments were conducted for three replicates, and the data were calculated according to the 2\u003csup\u003e\u0026minus;\u003c/sup\u003e\u003csup\u003e△△\u003c/sup\u003e\u003csup\u003eCT\u003c/sup\u003e method [53].\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePlant materials and stress treatments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe seedlings of two sweetpotato cultivars (\u003cem\u003ecv\u003c/em\u003e. Jishu25 and Jishu29) were collected from the Crop Ressearch Institue, Shandong Academy of Agricultural Sciences, China. The uniform seedlings of the two cultivars were grown in the Hoagland solution at 26\u0026deg;C under a photoperiod of 16 h light/8 h dark. When the seedlings had five to six functional leaves and adventitious roots of 8 to 10 cm, these seedlings were subjected to six different stresses, respectively. To study the expression patterns under these stresses, the adventitious roots of seedlings were submerged in the solution containing 150 mM NaCl, 20% PEG 6000, 100 mM ABA, 50 mg/L uniconazole, and 50 mg/L gibberellic acid (GA\u003csub\u003e3\u003c/sub\u003e) respectively [54].\u0026nbsp;For\u0026nbsp;black spot\u0026nbsp;pathogen\u0026nbsp;treatment,\u0026nbsp;\u003cem\u003eC. fimbriata\u003c/em\u003e conidia was collected after growing in potato dextrose agar (PDA) at 28 \u0026deg;C for 7 days, then were diluted to 1\u0026times;10\u003csup\u003e4\u003c/sup\u003e spores/mL with sterile water, and then\u0026nbsp;the roots of sweetpotato\u0026nbsp;seedlings were\u0026nbsp;cultivated\u0026nbsp;in the\u0026nbsp;1\u0026times;10\u003csup\u003e4\u0026nbsp;\u003c/sup\u003espores/mL conidia\u0026nbsp;suspension.\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eThe treated roots and leaves were collected after 0, 3, 6, 12, 24, and 48 h. To investigate the \u003cem\u003eIbbgals\u003c/em\u003e transcript levels in different tissues, the fifth expanded leaves, lips, stems, fibrous roots and storage roots of the two cultivars were sampled at 125 days after transplanting, \u0026nbsp;and the storage roots were sampled at 40, 55, 70, 95, 110, 125 and 150 days after transplanting in the sweetpotato field.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStatistical analysis\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eStatistical analysis was performed using the SPSS software package (v13.0), and the datas were presented as means of three replicates. Differences between means were subjected to ANOVA, and the statistical significance of the difference between means was calculated with Duncan\u0026apos;s new multiple ranges test and marked with asterisks at p \u0026lt; 0.05.\u003c/p\u003e"},{"header":"Abbreviations","content":"\u003cp\u003eABA: abscisic acid; BLAST: \u003ca href=\"https://www.baidu.com/link?url=lGiS6Xb1PfCiUwjzEEoDrMDgmkkaC1WaT88rqOvxcBbgi8UuvbpBEw7f3BhAohcB\u0026amp;wd=\u0026amp;eqid=cc251da50000151b000000025ecf230a\"\u003eBasic Local Alignment Search Tool\u003c/a\u003e; bgal: \u0026beta;-galactosidase; GH35: glycoside hydrolase 35; GA3: gibberellins; IAA: indolyl-3-acetic acid; JA: \u003ca href=\"http://www.baidu.com/baidu.php?url=a00000KEJeCxDFezEbu5o8xeY2Mii0X9BChcypuG6X0lq5WP2bm7-LWnqTQJX-jwK9L7tV6CnLUQ2I6xJjtV_rFgUmTqwwsgAhU1ewTr86sIFxCtVssseP2DvQCyupJ6NMgd5wEyTmwT5X7X_FIX17P4Klwm6U8p-Hnrk33qQOmsUaHxdF-FKtJHin2WsF_BQ2eYT4ViF_wofQ2nWA8Su47s9yWB.7b_jizeVu1gETBaLOBXM6thmpjLwItHAOuEzQDk8_____zNtVHQ8gZJyAp7W_vUVS4f.U1Yk0ZDqibZTdnjzzSD0TA-W5H00TZPGuv3quW-WuH6sPW-bnWNWnhPBPvczP1PhnvFhmW64Pjc4ryf0IjLTdnjzzSD0pyYqnWcd0ATqUvNsT1D0Iybqmh7GuZR0TA-b5Hcd0APGujYkrHm0UgfqnH0kPdtknjD4g1csPWFxnW0dnNt1PW0k0AVG5H00TMfqP1c10ANGujY1nHmsP6KBpHYkPHNxnHR3g1csP7tznHT0UynqnH6YPjDzrjRkn7tknj0kg1DYrHmvPHbYnWbkg1c4PjfkPW6vg100TgKGujYs0Z7Wpyfqn0KzuLw9u1Ys0A7B5HKxn0K-ThTqn0KsTjYs0A4vTjYsQW0snj0snj0s0AdYTjYs0AwbUL0qn0KzpWYs0Aw-IWdsmsKhIjYs0ZKC5H00ULnqn0KBI1Ykn0K8IjYs0ZPl5fK9TdqGuAnqTZnVmvY0pywW5Nwj0ZwdT1YknWfLPjndnjnYrj6zPjDLn10z0ZF-TgfqnHR4njm1rjbzPW6zPsK1pyfqrAR3m16snHfsnj0vrynYmsKWTvYqfRFawWfvPWRYfbuKfWm4ffK9m1Yk0ZK85H00TydY5H00Tyd15H00XMfqn0KVmdqhThqV5HKxn7tsg1Kxn0Kbmy4dmhNxTAk9Uh-bT1Ysg1Kxn7tsg100TA7Ygvu_myTqn0Kbmv-b5H00ugwGujYVnfK9TLKWm1Ys0ZNspy4Wm1Ys0Z7VuWYs0AuWIgfqn0KGTvP_5H00XMK_Ignqn0K9uAu_myTqnfK_uhnqn0KbmvPb5fKBuA-b5RcdPbFKPRc1njRYwHRsf1m1n1cswWcYwDDswHR3Pbn10AFY5H00Uv7YI1Ys0AqY5H00ULFsIjYsc10Wc10Wnansc108nj0snj0sc10Wc10WQinsQW0snj0snankQW0snj0sn0KkgLmqna33n-tsQW0sg108njKxna3sPNtsQW0Yg108n1c0ug9Y5H00mMPxTZFEuA-b5H00mLFW5HfvPjR\u0026amp;word=JA%E8%8C%89%E8%8E%89%E9%85%B8\u0026amp;ck=7623.5.137.264.152.134.139.2080\u0026amp;shh=www.baidu.com\u0026amp;sht=50000021_hao_pg\u0026amp;us=1.0.1.0.1.301.0\"\u003eJasmonic acid\u003c/a\u003e; MW: Molecular weights; NJ: Neighbor-Joining; pI: Isoelectric points; qRT-PCR: Quantitative reverse transcription polymerase chain reaction\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data generated or analyzed during this study are included in this published article and its supplementary information files.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare they have no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was supported by National Key R\u0026amp;D Program of China (2018YFD1000706,2018YFD1000700), Postgraduate Research \u0026amp; Practice Innovation Program of Jiangsu Province (KYCX19_2200), the China Agriculture Research System of sweetpotato (CARS-10-B7), Taishan industry leading talents project（2020-2023）and Shandong agricultural application technology project(2018-2020).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026apos; contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFH designed and performed the experiments and wrote the paper. ZQ and TD performed some experiments and analyzed the data, AL and SD analyzed the data. TX, DM and QW revised the paper. ZL and LZ conceived the experiment.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank the Ipomoea Genome Hub project team for sharing the Ipomoea batatas genome annotation data (https://ipomoea-genome.org/).\u003c/p\u003e"},{"header":"References ","content":"\u003col\u003e\n\u003cli\u003eLetunic I, Bork P. Interactive Tree Of Life v2: online annotation and display of phylogenetic trees made easy. Nucleic Acids Res. 2011; 39:475\u0026ndash;8.\u003c/li\u003e\n\u003cli\u003eSmith DL, Gross KC. A family of at least seven \u0026beta;-galactosidase genes is expressed during tomato fruit development. Plant Physiol. 2000; 123: 1173\u0026ndash;83.\u003c/li\u003e\n\u003cli\u003eEsteban R, Labrador E, Dopico B. 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Nucleic Acids Res. 2002; 30(1):325\u0026ndash;7.\u003c/li\u003e\n\u003cli\u003eLivak \u003ca href=\"https://www.sciencedirect.com/science/article/pii/S1046202301912629#!\"\u003eKJ,\u003c/a\u003e Schmittgen TD\u003ca href=\"https://www.sciencedirect.com/science/article/pii/S1046202301912629#!\"\u003e. \u003c/a\u003eAnalysis of relative gene expression data using real-time quantitative PCR and the 2\u003csup\u003e\u0026minus;\u0026Delta;\u0026Delta;CT\u003c/sup\u003e method. \u003ca href=\"https://www.sciencedirect.com/science/journal/10462023\"\u003eMethods\u003c/a\u003e. 2001; 25(4): 402-8.\u003c/li\u003e\n\u003cli\u003eYang Z, Zhu P, Kang H, Liu L, Cao Q, Sun J, et al. High-throughput deep sequencing reveals the important role that microRNAs play in the salt response in sweet potato (Ipomoea batatas L.). BMC Genomics. 2020; 21(1):164.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1 Gene and protein analysis of bgals in \u003cem\u003eIpomoea batatas. \u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"1\" width=\"609\"\u003e\n\u003ctbody\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003eGene\u003c/p\u003e\n\u003cp\u003ename\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003eCDS\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003eLength(aa)\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003eMW(kDa)\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003epI\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003eSubcellular localization\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003eSignal peptides\u003csup\u003ee\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003eN-glycosylation site\u003csup\u003ef\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal1\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2529\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e842\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e94.005\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e5.98\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e3\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal2\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2196\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e731\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e81.393\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e8.39\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal3\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2526\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e841\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e93.635\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e7.27\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e1\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 4\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2529\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e842\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e93.578\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e8.71\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e1\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 5\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2022\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e673\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e74.792\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e6.32\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003enucleus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e1\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 6\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2529\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e842\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e93.665\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e7.94\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e1\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal7\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2481\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e826\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e7.22\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e9.32\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003eextracellular\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e4\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal8\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2541\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e846\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e91.829\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e6.37\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal9\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2463\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e820\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e92.0858\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e5.31\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal10\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2391\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e796\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e89.004\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e6.83\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003enucleus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e4\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal11\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2505\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e834\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e94.335\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e8.57\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e5\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal12\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2187\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e728\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e80.867\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e9.13\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal13\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e3333\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e1110\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e125.149\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e5.5\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e6\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal14\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2487\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e828\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e93.578\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e8.71\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003evacuole\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e5\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal15\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2475\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e824\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e93.72\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e8.58\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e5\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal16\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2412\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e803\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e89.731\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e6.34\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e+\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e6\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"73\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal17\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"57\"\u003e\n\u003cp\u003e2145\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e714\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"85\"\u003e\n\u003cp\u003e79.382\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"47\"\u003e\n\u003cp\u003e7.99\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"95\"\u003e\n\u003cp\u003echloroplast\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"76\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"91\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003ea\u003c/sup\u003eThe length of Ibbgals coding sequence\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb\u003c/sup\u003eThe length of Ibbgals protein.\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ec\u003c/sup\u003eMolecular weight\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ed\u003c/sup\u003eTheoretical isoelectric point\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ee\u003c/sup\u003e\u0026ldquo;+ \u0026rdquo; means contain signal peptide, \u0026ldquo;_\u0026rdquo; means lack signal peptide.\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ef\u003c/sup\u003e Predicted using NetNGlyc.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 2. The putative cis-elements in the promoters of 17 \u003cem\u003eIbbgal\u003c/em\u003es genes.\u003c/p\u003e\n\u003ctable border=\"1\"\u003e\n\u003ctbody\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003egene\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003ePlant hormone response elements\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eStress response elements\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003e\n\u003cp\u003eLight response elements\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eOther elements\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 1\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e4\u003c/sup\u003e, AuxRE\u003csup\u003e2\u003c/sup\u003e, GARE\u003csup\u003e2\u003c/sup\u003e, TATC-BOX, PYRIMIDINEBOXHVEPB1\u003c/td\u003e\n\u003ctd width=\"170\"\u003ebox-W\u003csup\u003e2\u003c/sup\u003e, MYC-like\u003csup\u003e18\u003c/sup\u003e, ACGT\u003csup\u003e10\u003c/sup\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e8\u003c/sup\u003e, GT1-motif\u003csup\u003e5\u003c/sup\u003e, Box 4 \u003csup\u003e8\u003c/sup\u003e, IBOX\u003csup\u003e5\u003c/sup\u003e, GBOX\u003csup\u003e3\u003c/sup\u003e, GATAbox\u003csup\u003e10\u003c/sup\u003e, GAG-motif, TCT-motif\u003csup\u003e3\u003c/sup\u003e, Box II\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eEEs, TATA-box\u003csup\u003e21\u003c/sup\u003e,\u0026nbsp; GT\u003csup\u003e15\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e, AAGAA-motif\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 2\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eGARE\u003csup\u003e4\u003c/sup\u003e, TGACG-motif2, DPBFCOREDCDC3\u003csup\u003e2\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e4\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eMBS\u003csup\u003e2\u003c/sup\u003e, MYC-like\u003csup\u003e18\u003c/sup\u003e, ACGT\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR \u003csup\u003e3\u003c/sup\u003e, IBOX\u003csup\u003e2\u003c/sup\u003e, GATAbox\u003csup\u003e14\u003c/sup\u003e,GAG-motif,TBOX\u003csup\u003e2\u003c/sup\u003e, TCT-motif\u003csup\u003e2\u003c/sup\u003e,AT1-motif\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian\u003csup\u003e2\u003c/sup\u003e, TATA-box\u003csup\u003e18\u003c/sup\u003e, CCAAT-box\u003csup\u003e9\u003c/sup\u003e, GCN4-motif, RY-element\u003csup\u003e4\u003c/sup\u003e, GT\u003csup\u003e12\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 3\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE,ERE, DPBFCOREDCDC3\u003csup\u003e3\u003c/sup\u003e,\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eMYC-like\u003csup\u003e16\u003c/sup\u003e, ACGT\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e2\u003c/sup\u003e, GT1-motif, IBOX\u003csup\u003e6\u003c/sup\u003e, DRE\u003csup\u003e2\u003c/sup\u003e, GATAbox\u003csup\u003e15\u003c/sup\u003e, GAG-motif, TBOX\u003csup\u003e3\u003c/sup\u003e, TCT-motif, Box II\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian, TATA-box\u003csup\u003e17\u003c/sup\u003e, CCAAT-box\u003csup\u003e6\u003c/sup\u003e, RY-element\u003csup\u003e2\u003c/sup\u003e , GT\u003csup\u003e12\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 4\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e5\u003c/sup\u003e, GARE, AuxRE\u003csup\u003e2\u003c/sup\u003e, PYRIMIDINEBOXHVEPB1\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003ebox-W, MYC-like1\u003csup\u003e8\u003c/sup\u003e, ACGT\u003csup\u003e10\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR8, GT1-motif5, Box 4\u003csup\u003e8\u003c/sup\u003e, IBOX5, GATAbox10, GAG-motif, TCT-motif\u003csup\u003e3\u003c/sup\u003e, Box II\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eEEs,TATA-box21,CCAAT-box3,GT\u003csup\u003e15\u003c/sup\u003e, AAGAA-motif\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 5\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE\u003csup\u003e3\u003c/sup\u003e, ERE, GARE, CGTCA-motif\u003csup\u003e2\u003c/sup\u003e, TGACG-motif4, DPBFCOREDCDC3\u003csup\u003e4\u003c/sup\u003e, PYRIMIDINEBOXHVEPB1\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eLRT, box-W, MYC-like1\u003csup\u003e2\u003c/sup\u003e, ACGT\u003csup\u003e8\u003c/sup\u003e, MBS\u003csup\u003e3\u003c/sup\u003e ,GT1\u003csup\u003e8\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR6, GT1-motif2, Box 4\u003csup\u003e3\u003c/sup\u003e, IBOX3, GATAbox15, Box A, TBOX,TCT-motif2, Box II2\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian3, TATA-box15, CCAAT-box6, Box A ,\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 6\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE\u003csup\u003e2\u003c/sup\u003e, ERE, GARE\u003csup\u003e2\u003c/sup\u003e, CGTCA-motif\u003csup\u003e2\u003c/sup\u003e, TGACG-motif4, DRE2COREZMRAB17, PYRIMIDINEBOXHVEPB1\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eLRT\u003csup\u003e3\u003c/sup\u003e, MYC-like\u003csup\u003e10\u003c/sup\u003e, ACGT\u003csup\u003e12\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e4\u003c/sup\u003e, GT1-motif, Box 4, IBOX\u003csup\u003e8\u003c/sup\u003e, GATAbox\u003csup\u003e22\u003c/sup\u003e, TBOX, TCT-motif\u003csup\u003e5\u003c/sup\u003e, Box II\u003csup\u003e4\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eTATA-box\u003csup\u003e21\u003c/sup\u003e, CCAAT-box\u003csup\u003e4\u003c/sup\u003e, RY-element, GT\u003csup\u003e13\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 7\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eERE, GARE\u003csup\u003e2\u003c/sup\u003e, AuxRE, CGTCA-motif, TGACG-motif\u003csup\u003e3\u003c/sup\u003e, DPBFCOREDCDC3\u003csup\u003e2\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eMYC-like\u003csup\u003e14\u003c/sup\u003e, ACGT\u003csup\u003e4\u003c/sup\u003e , GT-1\u003csup\u003e5\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003e\n\u003cp\u003eINR\u003csup\u003e4\u003c/sup\u003e, Box 4\u003csup\u003e2\u003c/sup\u003e, IBOX14, GATAbox\u003csup\u003e17\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian\u003csup\u003e4\u003c/sup\u003e, TATA-box\u003csup\u003e17\u003c/sup\u003e, CCAAT-box\u003csup\u003e9\u003c/sup\u003e, RY-element2\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 8\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE\u003csup\u003e3\u003c/sup\u003e, ERE, GARE, DPBFCOREDCDC3\u003csup\u003e4\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e4\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eLRT\u003csup\u003e2\u003c/sup\u003e, MYC-like\u003csup\u003e20\u003c/sup\u003e, DRE\u003csup\u003e2\u003c/sup\u003e, ACGT\u003csup\u003e12\u003c/sup\u003e, MBS2 ,GT-1\u003csup\u003e9\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e3\u003c/sup\u003e, GT1-motif, Box 4\u003csup\u003e4\u003c/sup\u003e, IBOX\u003csup\u003e8\u003c/sup\u003e, GATAbox\u003csup\u003e18\u003c/sup\u003e, TCT-motif\u003csup\u003e3\u003c/sup\u003e, Box II\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian\u003csup\u003e2\u003c/sup\u003e, TATA-box\u003csup\u003e20\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e, RY-element\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal 9\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE, ERE, GARE\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eLRT\u003csup\u003e3\u003c/sup\u003e, MYC-like\u003csup\u003e8\u003c/sup\u003e, ACGT\u003csup\u003e6\u003c/sup\u003e , GT-1\u003csup\u003e5\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e3\u003c/sup\u003e, GT1-motif, Box 4\u003csup\u003e2\u003c/sup\u003e, IBOX\u003csup\u003e13\u003c/sup\u003e,GATAbox\u003csup\u003e22\u003c/sup\u003e, Tbox\u003csup\u003e2\u003c/sup\u003e, Box II\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian\u003csup\u003e5\u003c/sup\u003e, EEs, TATA-box\u003csup\u003e28\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e,GCN4-motif, RY-element\u003csup\u003e4\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal10\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eABRE\u003csup\u003e2\u003c/sup\u003e,GARE,DPBFCOREDCDC3, CATATGGMSAUR\u003csup\u003e2\u003c/sup\u003e,PYRIMIDINEBOXHVEPB1\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003ebox-W, MYC-like\u003csup\u003e18\u003c/sup\u003e, ACGT\u003csup\u003e12\u003c/sup\u003e, MBS\u003csup\u003e3\u003c/sup\u003e, GT-1\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003e\n\u003cp\u003eINR\u003csup\u003e2\u003c/sup\u003e, Box 4\u003csup\u003e3\u003c/sup\u003e, IBOX\u003csup\u003e7\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eTATA-box\u003csup\u003e16\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e, RY-element\u003csup\u003e3\u003c/sup\u003e, Box A\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal11\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003e\n\u003cp\u003eGARE\u003csup\u003e3\u003c/sup\u003e,CATATGGMSAUR\u003csup\u003e2\u003c/sup\u003e, PYRIMIDINEBOXHVEPB1\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"170\"\u003e\n\u003cp\u003eMYC-like\u003csup\u003e8\u003c/sup\u003e, ACGT\u003csup\u003e4\u003c/sup\u003e, MBS\u003csup\u003e2\u003c/sup\u003e, GT-1\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e5\u003c/sup\u003e, GT1-motif, Box 4\u003csup\u003e3\u003c/sup\u003e, IBOX\u003csup\u003e7\u003c/sup\u003e, GATAbox\u003csup\u003e18\u003c/sup\u003e, GAG-motif, TBOX\u003csup\u003e2\u003c/sup\u003e, TCT-motif, Box II\u003c/td\u003e\n\u003ctd width=\"188\"\u003e\n\u003cp\u003eCircadian,TATA-box\u003csup\u003e23\u003c/sup\u003e, CCAAT-box\u003csup\u003e4\u003c/sup\u003e,AAGAA-motif, RY-element\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal12\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e3\u003c/sup\u003e, ERE, GARE\u003csup\u003e4\u003c/sup\u003e, TGACG-motif, PYRIMIDINEBOXHVEPB1\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e3\u003c/sup\u003e, box-W, MYC-like\u003csup\u003e18\u003c/sup\u003e, DRE\u003csup\u003e4\u003c/sup\u003e, ACGT\u003csup\u003e8\u003c/sup\u003e,GT-1\u003csup\u003e8\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e8\u003c/sup\u003e，GT1-motif, Box 43, IBOX3, GATAbox21, TCT-motif, Box II\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003eCircadian\u003csup\u003e2\u003c/sup\u003e, TATA-box\u003csup\u003e27\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e,RY-element\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal13\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e3\u003c/sup\u003e, ERE, TGACG-motif, DPBFCOREDCDC3\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e2\u003c/sup\u003e, MYC-like\u003csup\u003e18\u003c/sup\u003e, ACGT\u003csup\u003e6\u003c/sup\u003e, MBS\u003csup\u003e2\u003c/sup\u003e, GT-1\u003csup\u003e4\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e4\u003c/sup\u003e, GT1-motif\u003csup\u003e3\u003c/sup\u003e, IBOX\u003csup\u003e15 \u003c/sup\u003e, GATAbox\u003csup\u003e15\u003c/sup\u003e, GAG-motif, TBOX, Box II\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003eCircadian, TATA-box\u003csup\u003e12\u003c/sup\u003e, CCAAT-box\u003csup\u003e4\u003c/sup\u003e, RY-element\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal14\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e3\u003c/sup\u003e, ERE, GARE, TGACG-motif, DPBFCOREDCDC3\u003csup\u003e2\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e4\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e4\u003c/sup\u003e, box-W, MYC-like\u003csup\u003e14\u003c/sup\u003e, ACGT\u003csup\u003e6\u003c/sup\u003e, MBS,GT-1\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e3\u003c/sup\u003e, GT1-motif\u003csup\u003e2\u003c/sup\u003e, Box 4, IBOX\u003csup\u003e10\u003c/sup\u003e, GATAbox\u003csup\u003e18\u003c/sup\u003e,CATT, TBOX\u003csup\u003e3\u003c/sup\u003e, Box II\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"188\"\u003eCircadian, TATA-box\u003csup\u003e13\u003c/sup\u003e, CCAAT-box\u003csup\u003e6\u003c/sup\u003e, RY-element\u003csup\u003e3\u003c/sup\u003e\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal15\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eGARE\u003csup\u003e2\u003c/sup\u003e, DPBFCOREDCDC3\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e3\u003c/sup\u003e, box-W\u003csup\u003e2\u003c/sup\u003e, MYC-like\u003csup\u003e28\u003c/sup\u003e, GT-1\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e4\u003c/sup\u003e, GT1-motif\u003csup\u003e2\u003c/sup\u003e, IBOX\u003csup\u003e3\u003c/sup\u003e, GATAbox\u003csup\u003e10\u003c/sup\u003e, TBOX\u003csup\u003e2\u003c/sup\u003e, TCT-motif, Box II\u003c/td\u003e\n\u003ctd width=\"188\"\u003eCircadian, TATA-box\u003csup\u003e2\u003c/sup\u003e, CCAAT-box\u003csup\u003e5\u003c/sup\u003e, RY-element\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal16\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eERE, GARE\u003csup\u003e2\u003c/sup\u003e, DPBFCOREDCDC3\u003csup\u003e3\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e2\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e2\u003c/sup\u003e, box-W, MYC-like\u003csup\u003e8\u003c/sup\u003e, DRE\u003csup\u003e3\u003c/sup\u003e, GT-1\u003csup\u003e6\u003c/sup\u003e\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e4\u003c/sup\u003e, Box 4\u003csup\u003e5\u003c/sup\u003e, IBOX\u003csup\u003e2\u003c/sup\u003e, GATAbox\u003csup\u003e13\u003c/sup\u003e, GAG-motif, TBOX, TCT-motif\u003c/td\u003e\n\u003ctd width=\"188\"\u003eTATA-box\u003csup\u003e36\u003c/sup\u003e, CCAAT-box\u003csup\u003e3\u003c/sup\u003e, RY-element\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"71\"\u003e\n\u003cp\u003e\u003cem\u003eIbbgal17\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"257\"\u003eABRE\u003csup\u003e7\u003c/sup\u003e, ERE, GARE\u003csup\u003e3\u003c/sup\u003e, TGACG-motif\u003csup\u003e4\u003c/sup\u003e, DPBFCOREDCDC3\u003csup\u003e6\u003c/sup\u003e, CATATGGMSAUR\u003csup\u003e2\u003c/sup\u003e, GCCCORE\u003c/td\u003e\n\u003ctd width=\"170\"\u003eLRT\u003csup\u003e2\u003c/sup\u003e, box-W\u003csup\u003e3\u003c/sup\u003e, MYC-like\u003csup\u003e10\u003c/sup\u003e, ACGT\u003csup\u003e6\u003c/sup\u003e, MBS\u003csup\u003e2\u003c/sup\u003e, GT-1\u003c/td\u003e\n\u003ctd width=\"258\"\u003eINR\u003csup\u003e2\u003c/sup\u003e, GT1-motif, Box 4, IBOX\u003csup\u003e9\u003c/sup\u003e, GATAbox\u003csup\u003e24\u003c/sup\u003e, TBOX, Box II\u003c/td\u003e\n\u003ctd width=\"188\"\u003eTATA-box\u003csup\u003e18\u003c/sup\u003e, CCAAT-box\u003csup\u003e4\u003c/sup\u003e, GCN4-motif, RY-element\u003csup\u003e4 \u003c/sup\u003e\u003c/td\u003e\n\u003c/tr\u003e\n\u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eSuperscript numbers represent the repeats (2 or more than 2) of each cis-element in the \u003cem\u003eIbbgal\u003c/em\u003e promoter, while the others only contain one copy of corresponding cis-element.\u003c/p\u003e\n\u003cp\u003eABRE and ACGT, cis-acting elements involved in the abscisic acid responsiveness;AuxRE, cis-acting regulatory element involved in auxin responsiveness; AAGAA-motif, cis-element involved in secondary xylem development; Box A, cis-acting elements of phenylalanine ammonia-lyase; Box II, part of a light responsive element; Box-W, fungal elicitor responsive element; Box 4, part of a conserved DNA module involved in light responsiveness; CATATGGMSAUR , cis-acting element involved in auxin responsiveness; CCAAT-box, MYBHv1 binding site; Circadian, cis-acting regulatory element involved in circadian control; DPBFCOREDCDC3, induced by ABA; DRE, cis-acting element involved in drought response;EEs, part of evening and circadian response; ERE, ethylene-responsive element; GARE, gibberellin-responsive element; GATA-motif,part of a light responsive element; Gbox, cis-acting regulatory element involved in light responsiveness; GATAbox, part of a light responsive element; GAG-motif, part of a light responsive element; GCCCORE , cis-acting element involved in jasmonate responsiveness; GCN4-motif, cis-regulatory element involved in endosperm; GT1-motif, light responsive element; GT-1, cis-acting element involved in the salt stress; INR, part of a light responsive element; IBOX, part of a light responsive element; LTR, cis-acting element involved in low-temperature responsiveness; MBS, MYB binding site involved in drought-inducibility; MYC-like , cis-acting elements of drought-responsive; PYRIMIDINEBOXHVEPB1, cis- and trans-acting elements involved in gibberellins and abscisic acid responsiveness; RY-element, cis-acting regulatory element involved in seedspecific regulation; TATA-box, core promoter element around \u0026minus;30 of transcription start; TATC-box, cis-acting element involved in gibberellin-responsiveness;TBOX, part of a light responsive element; TCT-motif, part of a light responsive element; TGACG-motif, cis-acting regulatory element involved in the MeJA-responsiveness.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"bmc-genomics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"gics","sideBox":"Learn more about [BMC Genomics](http://bmcgenomics.biomedcentral.com/)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/gics","title":"BMC Genomics","twitterHandle":"#BMCGenomics","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"em","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"sweetpotato, β-Galactosidase, gene expression, stress","lastPublishedDoi":"10.21203/rs.3.rs-32133/v3","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-32133/v3","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eBackground:\u003c/strong\u003e Sweetpotato (\u003cem\u003eIpomoea batatas\u003c/em\u003e (L.) Lam.) serves as an important food source for human beings. β-galactosidase (bgal) is a glycosyl hydrolase involved in cell wall modification, which plays essential roles in plant development and environmental stress adaptation. However, the function of \u003cem\u003ebgal\u003c/em\u003e genes in sweetpotato remains unclear.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eResults:\u003c/strong\u003e In this study, 17 β-galactosidase genes (\u003cem\u003eIbbgal\u003c/em\u003e) were identified in sweetpotato, which were classified into seven subfamilies using interspecific phylogenetic and comparative analyses. The promoter regions of \u003cem\u003eIbbgal\u003c/em\u003es harbored several stress, hormone and light responsive cis-acting elements. Quantitative real-time PCR results displayed that \u003cem\u003eIbbgal \u003c/em\u003egenes had the distinct expression patterns across different tissues and varieties. Moreover, the expression profiles under various hormonal treatments, abiotic and biotic stresses were highly divergent in leaves and root. \u003c/p\u003e\u003cp\u003e\u003cstrong\u003eConclusions:\u003c/strong\u003e Taken together, these findings suggested that \u003cem\u003eIbbgals\u003c/em\u003e might play an important role in plant development and stress responses, which provided evidences for further study of bgal function and sweetpotato breeding.\u003c/p\u003e","manuscriptTitle":"Genome-wide in silico identification and expression analysis of beta-galactosidase family members in sweetpotato [Ipomoea batatas (L.) Lam.]","msid":"","msnumber":"","nonDraftVersions":[{"code":3,"date":"2020-12-10 16:35:59","doi":"10.21203/rs.3.rs-32133/v3","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Minor revision","date":"2021-01-27T00:00:00+00:00","index":"","fulltext":""},{"type":"reviewersInvited","content":"","date":"2020-11-25T00:00:00+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2020-11-24T00:00:00+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2020-11-23T23:00:00+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2020-11-23T23:00:00+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"bmc-genomics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"gics","sideBox":"Learn more about [BMC 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