Genome-Wide Identification and Investigation of MST Gene Family Based on Their Evolution and Expression Analysis under Abiotic Stress and Hormone Treatments in Maize (Zea mays L.)

Genome-Wide Identification and Investigation of MST Gene Family Based on Their Evolution and Expression Analysis under Abiotic Stress and Hormone Treatments in Maize (Zea mays L.) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Genome-Wide Identification and Investigation of MST Gene Family Based on Their Evolution and Expression Analysis under Abiotic Stress and Hormone Treatments in Maize (Zea mays L.) Jialun Zhu, Tianfeng Li, Jing Ma, Wenyu Li, Hanyu Zhang, Tsyganova Nadezhda, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3219446/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 12 You are reading this latest preprint version Abstract Background Monosaccharide transporter (MST) family, as a carrier for monosaccharide transport, plays an important role in carbon partitioning and widely involves in plant growth and development, stress response, and signaling transduction. However, little information on the MST family genes is reported in maize ( Zea mays ), especially in response to abiotic stresses. In this study, the genome-wide identification of MST family genes was performed in maize. Result A total of sixty-six putative members of MST gene family were identified and divided into seven subfamilies (including SPT, PMT, VGT, INT, pGlcT, TMT, and ERD) using bioinformatics approaches, and gene information, phylogenetic tree, chromosomal location, gene structure, motif composition, and cis -acting elements were investigated. Eight tandem and twelve segmental duplication events were identified, which played an important role in the expansion of the ZmMST family. Synteny analysis revealed the evolutionary features of MST genes in three gramineous crop species. The expression analysis indicated that most of the PMT, VGT, and ERD subfamilies members responded to osmotic and cadmium stresses, and some of them were regulated by ABA signaling, while only a few members of other subfamilies responded to stresses. In addition, only five genes were induced by NaCl stress in MST family. Conclusion These results serve to understand the evolutionary relationships of the ZmMST family genes and supply some insight into the processes of monosaccharide transport and carbon partitioning on the balance between plant growth and development and stress response in maize. Genome-wide identification Zea mays L. Monosaccharide transporter (MST) Gene family Abiotic stress Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Background The industrial activities of human caused the trends of climate warming and constant changes in the natural environment [ 1 , 2 ]. Many abiotic stresses such as drought, salinity, low temperature, and heavy metals pollution affect the growth and development of plants and the yield of crops, and further threaten food security and human health[ 3 ]. Sugars are the main form of long-distance transportation and distribution of photosynthate and play an important role in plant growth and development [ 4 ]. Sugar not only constitutes metabolites, nutrients, and signal molecules, but also can be used as osmotic substances. One way that plants protect cell structure and alleviate the damage of abiotic stress is by accumulating osmotic substances or compatible solutes [ 4 , 5 ]. In Arabidopsis thaliana , sugar plays an important role in protecting plant structure by increasing the concentration of soluble sugars (sucrose, glucose and fructose) in sink leaves and sucrose in phloem sap, maintaining water potential and improving plant stress tolerance [ 6 – 8 ]. Therefore, sugar transport and distribution are key for crops in yielding and coping with abiotic stress [ 9 , 10 ]. In plants, various sugar transporters are involved in the transport of sugar from source to sink, of which the major facilitator superfamily (MFS) is an important member [ 11 ]. MFS can be generally divided into the sucrose transporter (SUT) family and MST family according to the different transport substrates. MSTs are complete membrane proteins that can participate in the transmembrane transport of monosaccharides [ 12 ]. The MST family can also be further divided into seven subfamilies based on their substrate specificities and sequence features, including Sugar Transport Protein (STP), Polyol/Monosaccharide Transporter (PMT), Vacuolar Glucose Transporter (VGT), Inositol Transporter (INT), Plastidic Glucose Transporter (pGlcT), Tonoplast Membrane Transporter (TMT), and Early-Responsive to Dehydration six-like (ERD) [ 13 ]. STP is a sugar transporter that mainly transports hexose. In Arabidopsis, most STPs exhibited a broad spectrum of absorption characteristics of substrates. AtSTP1 was the function of sugar transport and expressed in germinating seeds and the roots of seedlings, which played a key role in germination and root development [ 14 ]. In atstp1 mutant, the ability to transport sugar alcohols (D-glucose, D-galactose, and D-mannose) was greatly weakened compared with the wild type [ 14 ]. By investigating AtSTP6 -promoter:: GUS plants and conducting in situ hybridization experiments at the late stage of pollen development, it was determined that AtSTP6 was expressed. A transposon-tagged Arabidopsis mutant also demonstrated that the atstp6 mutation may have an impact on pollen vitality, pollen germination, fertilization, and seed production [ 15 ]. PMT proteins (PMTs) not only transport mannitol, sorbitol, xylitol, and other polyols, but also transport monosaccharides. The first PMT gene was found in Apium graveolens L., named AgMAT1 , and it played an important role in the loading process of phloem mannitol [ 16 ]. Arabidopsis genome contained six PMT subfamily genes, named AtPMT1 to AtPMT6 . AtPMT5 was located in the plasma membrane and could transport pentoses such as sugar alcohols (sorbitol, xylitol, erythritol, and glycerol), hexose, and ribose. AtPMT5 was highly expressed in Arabidopsis roots and played an important role in plant morphological construction [ 17 ]. In plants, the VGT subfamily generally had only 2–3 members, and played a key role in the process of seed germination, flowering, and other growth and development [ 18 , 19 ]. In Arabidopsis, AtVGT1 was H + /glucose reverse transporter located on the vacuolar plastid, which was involved in the transport and storage of monosaccharides in vacuoles. AtVGT1 also played a key role in the process of seed germination, flowering, and other growth and development, and the atvgt1 mutant had a lower germination rate and delayed flowering [ 20 ]. INT proteins were highly specific H + -inositol symporters, and the functions of them were in the transportation and distribution of inositol. Four INT genes ( AtINT1 - 4 ) were identified in Arabidopsis, of which AtINT4 was the first identified INT subfamily member. AtINT4 was highly expressed in Arabidopsis pollen and phloem companion tissue, which was related to plant pollen development and participated in inositol loading in phloem [ 21 ]. pGlcT proteins (pGlcTs) could transport glucose. The pGlcT gene was cloned in spinach ( Spinacia oleracea L.) and proved the role in transporting starch and hydrolyzing glucose [ 21 ]. In Arabidopsis, the atpglct1 , atpglct2 , and atpglct1 / atpglct2 mutants showed growth and development inhibition in varying degrees [ 22 ]. TMT proteins (TMTs, also known as ATZ) were also localized on vacuolar plastids. Previous studies reported that TMTs are involved in the transport of sucrose on vacuoles. In the Attmt1 / Attmt2 double mutant, the sucrose transportation to vacuoles was impaired [ 23 ]. The majority of the ERD6-like protein was found on the vacuolar membrane and was in charge of the transmembrane transport of sugar in the vacuoles [ 24 ]. In Arabidopsis, ERD (also known as SFPs) is the largest subfamily in the MST family and responds to abiotic stresses. For instance, AtERD1 and AtERD2 also responded to different stress treatments, and the expression patterns of them showed significantly different [ 25 ]. Genome-wide identification of the MST family had been performed in many plants, such as Arabidopsis, rice, grape, and tobacco [ 12 , 26 – 28 ]. The distribution of monosaccharide transporters in maize plants directly affects plant growth and development and abiotic stress response. However, genome-wide identification, evolutionary analysis, and response to abiotic stresses of the maize monosaccharide transporter family have not yet been reported. In this study, 66 MST genes were identified from the maize genome. Phylogenetic analysis, gene structure analysis, and synteny analysis were performed to understand the evolution and amplification of the ZmMST family, and the expression analysis in different tissues and under different treatments was performed to explore the response to abiotic stresses. These results provide insights into the evolution of the maize MST family and their role in maize growth and development and abiotic stress response. The identification and characterization of ZmMST genes may provide opportunities for the optimization of maize variety selection and breeding. Results Identification and phylogenetic analysis of the MST gene family members in maize The Arabidopsis MST proteins were queried against the maize genome using BLASTP to search for maize MST genes. The gene domain was manually confirmed through the NCBI CDD, SMART, and Pfam websites, and the protein sequence length (number of amino acids), molecular weight, and isoelectric point were determined by the ExPASy proteomics system. The sequences of the monosaccharide transporter family revealed by screening have conserved structural domains. Finally, 66 complete monosaccharide transporter sequences were identified and divided into seven subfamilies in maize, including STP (23 members), PMT (20 members), VGT (2 members), INT (4 members), pGlcT (4 members), TMT (4 members), and ERD (9 members) subfamilies. Gene names, AA accession number, length of the gene, amino acid numbers, molecular weights, chromosomal locations, and pIs were listed in Table 1 . Meanwhile, 64, 69, and 77 genes of the MST family were identified in rice, sorghum, and millet, respectively, as shown in Additional file 1. Table 1 The identification of MST members in maize. Gene name AA accession number Length of gene(bp) Number of amino acid(aa) Molecular weight (Da) Theoretical(pI) Chr strand Transcript ZmSTP1 Zm00001d027268 1578 525 57494.9 9.26 Chr1:103477Chr7:1037365 - 1 ZmSTP2 Zm00001d028230 1545 514 56685 9.12 Chr1:27307350:27313346 - 1 ZmSTP3 Zm00001d032409 1611 536 56846.1 9.44 Chr1:22583003Chr1:225834001 - 1 ZmSTP4 Zm00001d032906 1527 508 54319 9.19 Chr1:24270208Chr1:242703904 + 1 ZmSTP5 Zm00001d003468 1131 376 39731.1 9.92 Chr2:4523420Chr6:45238299 + 1 ZmSTP6 Zm00001d003469 1542 513 54767.7 9.04 Chr2:4523873Chr8:45242608 - 2 ZmSTP7 Zm00001d003471 1674 557 59538.2 10.18 Chr2:4528734Chr1:45292164 - 1 ZmSTP8 Zm00001d005594 1557 518 56633 9.23 Chr2:18012113Chr9:180123254 + 1 ZmSTP9 Zm00001d007078 1473 490 53551.9 9.61 Chr2:221769030:221780225 + 1 ZmSTP10 Zm00001d044245 1530 509 55644.6 9.88 Chr3:22292368Chr9:222925949 - 1 ZmSTP11 Zm00001d049467 1545 514 55430.5 7.72 Chr4:3137898Chr1:31380525 - 1 ZmSTP12 Zm00001d050860 1566 521 56348.6 9.29 Chr4:12723237Chr2:127244717 + 1 ZmSTP13 Zm00001d053846 1656 551 59924.1 8.75 Chr4:24217007Chr3:242172587 - 2 ZmSTP14 Zm00001d016919 1341 447 48442.5 9.85 Chr5:18005825Chr7:180059691 + 1 ZmSTP15 Zm00001d018627 1572 523 57035.4 9.67 Chr7:13357Chr10:1339124 - 1 ZmSTP16 Zm00001d019138 1542 513 56137.9 8.85 Chr7:1871972Chr8:18721630 - 1 ZmSTP17 Zm00001d020071 1569 522 57267.6 9.6 Chr7:9039738Chr4:90401326 + 1 ZmSTP18 Zm00001d020463 1572 523 7884 9.19 Chr7:11645175Chr9:116453072 + 2 ZmSTP19 Zm00001d021775 1575 524 56912.7 9.25 Chr7:16284589Chr1:162848484 - 1 ZmSTP20 Zm00001d045395 1557 518 54517.2 10.13 Chr9:2053261Chr2:20535038 + 1 ZmSTP21 Zm00001d025572 1524 507 53521.9 9.46 Chr10:12264185Chr7:122645133 - 1 ZmSTP22 Zm00001d025573 1371 456 49861.4 10.02 Chr10:12270886Chr5:122710678 - 1 ZmSTP23 Zm00001d000183 1563 520 56829.3 9.58 ctg18Chr1:30749Chr1:312293 + 1 ZmPMT1 Zm00001d028144 1578 525 56032.4 8.05 Chr1:2401225Chr4:24014074 - 1 ZmPMT2 Zm00001d028151 1575 524 56153.1 9.23 Chr1:2429641Chr3:24298623 - 4 ZmPMT3 Zm00001d029645 1560 519 55295.4 9.91 Chr1:8094488Chr6:80947089 + 1 ZmPMT4 Zm00001d030464 1527 508 54026.8 9.3 Chr1:13452100Chr8:134522680 + 1 ZmPMT5 Zm00001d001817 1617 538 55880.6 7.13 Chr2:142570Chr1:1428025 - 1 ZmPMT6 Zm00001d001818 1650 549 57360.9 7.13 Chr2:143207Chr7:1434514 - 1 ZmPMT7 Zm00001d002864 1602 533 57519.7 6.53 Chr2:2496079Chr7:24963400 + 1 ZmPMT8 Zm00001d006688 1551 516 54367.5 9.39 Chr2:21447246Chr4:214474594 + 1 ZmPMT9 Zm00001d006697 1563 520 55079.1 9.37 Chr2:21453545Chr7:214537722 - 1 ZmPMT10 Zm00001d048771 1446 481 50201.4 8.55 Chr4:52585Chr10:5260332 + 1 ZmPMT11 Zm00001d048774 1506 501 52136.4 9.3 Chr4:528403Chr7:5285944 + 1 ZmPMT12 Zm00001d048775 1350 449 46171.2 9.63 Chr4:532866Chr1:5330696 + 1 ZmPMT13 Zm00001d048776 1464 487 50935.3 8.71 Chr4:538077Chr9:5382734 + 1 ZmPMT14 Zm00001d021935 1542 513 54830.1 9.75 Chr7:16637624Chr6:166378812 + 2 ZmPMT15 Zm00001d021936 1533 510 53894 8.37 Chr7:16638310Chr8:166385301 + 2 ZmPMT16 Zm00001d021938 1542 513 54255.2 9.09 Chr7:16655059Chr1:166552978 - 1 ZmPMT17 Zm00001d021942 1485 495 52501.1 8.86 Chr7:16660746Chr4:166609263 - 1 ZmPMT18 Zm00001d048178 1596 531 56736.5 8.86 Chr9:15171082Chr9:151713323 - 5 ZmPMT19 Zm00001d023939 1437 478 50163.6 8.9 Chr10:3031494Chr4:30317071 + 2 ZmPMT20 Zm00001d023941 1470 489 50575.4 8.55 Chr10:30402000:30405639 + 2 ZmVGT1 Zm00001d012938 1398 465 49263.3 8.75 Chr5:223036Chr5:2234904 - 4 ZmVGT2 Zm00001d014435 1554 517 55327 5.77 Chr5:4670830Chr3:46712407 + 8 ZmINT1 Zm00001d018803 1758 585 62441.4 8.58 Chr7:567092Chr8:5673511 + 1 ZmINT2 Zm00001d019537 1458 485 51911.3 8.89 Chr7:41043850:41045307 - 1 ZmINT3 Zm00001d025749 1602 533 56679.5 5.92 Chr10:12828174Chr3:128287246 - 2 ZmINT4 Zm00001d025834 1746 581 62805.8 8.44 Chr10:13139539Chr4:131399111 - 2 ZmpGlcT1 Zm00001d039973 1620 539 56585.5 9.11 Chr3:2131176Chr2:21317041 + 11 ZmpGlcT2 Zm00001d053334 1488 495 58078 6.71 Chr4:22658326Chr6:226590664 - 1 ZmpGlcT3 Zm00001d020374 1653 550 56806.8 9.36 Chr7:11002322Chr5:110030221 + 10 ZmpGlcT4 Zm00001d008567 1620 539 53360.3 8.31 Chr8:1314614Chr1:13153169 + 11 ZmTMT1 Zm00001d029762 1305 434 60523.5 4.96 Chr1:8613001Chr4:86134527 + 13 ZmTMT2 Zm00001d048823 1125 374 59874 6.46 Chr4:601307Chr3:6018723 - 7 ZmTMT3 Zm00001d014872 2292 763 80956.6 4.46 Chr5:6660522Chr8:66608036 - 3 ZmTMT4 Zm00001d016274 1806 601 68874 5.73 Chr5:15513335Chr4:155138132 - 11 ZmERD1 Zm00001d029254 1533 510 51871.2 5.74 Chr1:6385984Chr5:63864152 - 2 ZmERD2 Zm00001d040243 1452 483 39799.5 8.96 Chr3:3341091Chr8:33414667 - 4 ZmERD3 Zm00001d039051 1503 500 52615.2 9.25 Chr6:16929398Chr5:169298831 + 9 ZmERD4 Zm00001d039052 1233 410 53240.9 9.6 Chr6:16932079Chr3:169327039 + 1 ZmERD5 Zm00001d008374 1470 489 31835.5 10.37 Chr8:669913Chr9:6703444 + 1 ZmERD6 Zm00001d009600 1125 374 54327.8 7.31 Chr8:7249771Chr8:72504565 + 36 ZmERD7 Zm00001d009603 1491 496 53739.2 8.44 Chr8:72763170Chr8:72767938 + 4 ZmERD8 Zm00001d009605 1497 498 44025.2 9.34 Chr8:7286116Chr4:72868595 + 2 ZmERD9 Zm00001d009669 1521 506 51148.7 8.31 Chr8:7514623Chr3:75149733 - 10 As shown in Fig. 1 , the MST genes were randomly distributed on 10 maize chromosomes. Among them, 12 MST genes were distributed on chromosome (chr) 7, and only two genes existed on chr 6 and 9. The STP subfamily genes were dispersed across 8 of the 10 chromosomes, and ZmPMT genes were dispersed across 6 of the 10 chromosomes. ZmVGT genes had only two genes and were located on chr 5. ZmTMT genes were located on chr 1, 4, and 5, respectively. ZmINT genes were located on chr 7 and 10, respectively. ZmpGlcT genes were located on chr 3, 4, 7, and 8, respectively. ZmERD genes were located on chr 1, 3, and 8. To explore the systematic evolution, a phylogenetic tree of MST family genes in maize was constructed. As shown in Fig. 2 and Fig. 3 A, ZmMST genes were divided into five main branches and seven groups, which was consistent with the results of existing studies. The VGT subfamily and INT subfamily possessed a closed relationship, while the relationship of ERD and pGlcT was close. Meanwhile, phylogenetic trees were constructed in seven subfamilies, including members from three species of maize, rice, and Arabidopsis. (Additional file 2–8) Gene structure and motif composition of the maize MST gene family The exon-intron organizations of the MST family genes were detected to comprehend the evolution of the MST family in maize. As shown in Fig. 3 B, ZmSTP genes possessed one to four exons (20 with two to three exons, 3 with four exons, and ZmSTP11 with only one the most exon). Although the ZmPMT subfamily has many genes, the gene structure tends to be conserved, and all ZmPMT genes possess two to three exons (14 members with two exons and 6 members with three exons). The ZmVGT subfamily contains only two genes, ZmVGT1 possesses twelve exons and ZmVGT2 possesses fourteen exons. ZmINT and ZmTMT genes possesses one to six exons. ZmpGlcT genes possess 12 to 14 exons. The ZmERD subfamily is different from other subfamilies, which contain an extensive number of exons, from 7 to 18. The conserved motifs of MST proteins were identified by the MEME motif program. The result was shown in Fig. 3 C. Almost all MST family members contained six conserved domains, motifs 1 to 6, and each subfamily contained different conserved motifs. Except for the ERD and TMT subfamilies, the other five subfamilies all contain motif 7. Motif 8 exists in almost all subfamilies except the TMT and motif 11 exists in all subfamilies except the ERD and STP. While motifs 10, 13, 14, and 15 only exist in the STP subfamily. Overall, the gene structures and conserved motif compositions of MSTs were similar in the same subfamily. Synteny analysis of MST genes To analyze the expansion of the MST family, gene duplication events, both tandem duplication (TD) and segmental duplication (SD) were detected. Eight pairs of tandem replication gene pairs (Fig. 1 ), including ZmSTP5 / 6 , ZmSTP21 / 22 , ZmPMT1 / 2 , ZmPMT2 / 3 , ZmPMT5 / 6 , ZmPMT14 / 15 , ZmPMT15 / 16 and ZmERD3 / 4 , and 12 segmental duplication events, including ZmSTP5 / 13 , ZmSTP6 / 15 , ZmSTP9 / 18 , ZmSTP13 / 15 , ZmSTP15 / 22 , ZmSTP13 / 22 , ZmPMT1 / 8 , ZmPMT3 / 4 , ZmPMT8 / 14 , ZmpGlcT1 / 4 , ZmERD2 / 5 , and ZmERD3 / 6 were identified in maize (Fig. 4 ). To further explore the evolutionary mechanisms of the MST family, the syntenic maps of maize were constructed and associated with four representative plant species, including the monocotyledons Sorghum bicolor , Setaria italica , Oryza sativa , and the dicotyledon Medicago truncatula (Fig. 5 ). A total of 41 MST family genes in maize showed syntenic relationships with those in millet, followed by sorghum (47) and rice (33), and there was only one pair of homologous genes in alfalfa (on chr 8), and specific gene pairs are shown in Additional file 9. To better understand the evolutionary constraints acting on the MST gene family, the Ka/Ks ratios of the tandem and segmental duplications in MST gene pairs were calculated (Additional file 10). Five pairs of genes had Ka/Ks > 1 ( ZmSTP5 / 6 , ZmSTP5 / 12 , ZmERD2 / 5 , ZmERD3 / 4 , and ZmERD3 / 6 ). The Ka/Ks of others are all less than 1. Analysis of the cis -acting element in MST gene promoter regions To explore the potential regulatory mechanisms of MST family genes, the 1.5 kb upstream promoter region of MST genes was submitted into Plant CARE to detect the conserved cis -elements. Then, the identified cis -acting elements were divided into three categories: abiotic and biotic stress, phytohormone responsive, and plant growth and development (Fig. 6 ). In the categories of abiotic and biotic stresses, MYB (CCAAT-box) and MYC (CACAT-box) were the two types of cis -elements with the largest proportion in the MST family. Meanwhile, the anaerobic response element ARE, drought response element DRE, and low-temperature element LTR were also widely distributed in the promoter regions of MST genes. For phytohormone responsive elements, the ABRE, TCA, as-1, and CGTCA-motif which are involved in Abscisic Acid (ABA), salicylic acid (SA), and methyl jasmonate (MeJA) responses respectively were detected in the most of MST gene promoter regions. In addition, 18 cis -elements related to plant growth and development were identified, most of which were involved in the light response, accounting for more than half of the group. Expression analysis of MST genes in different maize tissues and in response to different treatments To further analyze the function of MST family members in response to abiotic stresses, the expression patterns of MST family genes were detected in maize under abiotic stresses and ABA treatment (Fig. 7 ). Under PEG treatment, the expression levels of most MST members were increased, especially in the PMT, ERD, and VGT subfamilies. Under NaCl treatment, the expression of only five genes was induced, of which ZmPMT9 was the most significantly increased. Under Cd stress, the transcript levels of ZmPMT8 /12/13/ 15 /16/17/19/ 20 and ZmERD2 / 3 / 4 / 6 were significantly increased. Abscisic acid is a phytohormone involved in regulating plant responses to abiotic stresses. The expression of most MST members was induced under exogenous ABA treatment. Overall, the expression of 61 ZmMST genes showed different alterations under the abiotic stresses and exogenous ABA treatment, and some genes were affected by multiple treatments (Additional file 11). The expression of MST genes in various tissues, including roots, stems, leaves, and seeds, was also examined (Fig. 8 , Additional files 12 and 13). ZmSTP1 , ZmSTP2 , and ZmSTP5 were relatively high and stable in various tissues and periods, while the expression levels of ZmSTP8 / 9 / 10 / 11 / 12 / 13 / 14 / 16 and 22 were relatively low in various tissues. However, with the development process, the expression of ZmSTP2 in roots, stems, leaves, and seeds were rapidly up-regulated, while ZmSTP5 showed a trend of high expression in all tissues and stages. The expression patterns of members of the same subfamily were complementary. For example, ZmSTP19 and ZmSTP23 were highly expressed in leaves but almost not in seeds. ZmSTP23 was highly expressed in leaves, but expressed at low levels in roots, stems, and seeds. ZmPMT1 and ZmPMT13 also had similar expression patterns. According to these results, ZmMST genes played a variety of roles in maize growth and development. Discussion Since monosaccharides such as glucose and fructose are essential for metabolism, storage, and transport, MST is crucial to the processes of carbon partitioning and abiotic stress response in plants [ 12 , 26 , 29 ]. Genome-wide analysis of the MST gene family has been widely carried out in many species. Fifty-three, sixty-four, and sixty-nine genes have been identified in Arabidopsis, rice, and sorghum respectively [ 26 , 29 ]. However, the MST gene family has not been identified in maize. In this study, 66 MST genes were identified in the maize genome, and 77 MST family genes were identified in millet (Table 1 and Additional file 1). By the phylogenetic tree results, ZmMST genes were further divided into 7 subfamilies [ 30 ]. Each subfamily of MST was shown to be specific in the differentiation and evolution of the different Gramineae based on our findings that the number of genes in each subfamily of MST varied in maize, rice, sorghum, and millet. It was reported that the STP subfamily is the largest subfamily in rice. The ERD subfamily is the largest subfamily in Arabidopsis [ 26 ]. In maize, the largest subfamily is also the STP family, which indicates that the MST family has species-specific subfamily expansion in different plants. These expansions may be caused by gene duplication events, which may play a key role in the evolution of the MST gene family. The previous study proved that most genes in the Arabidopsis genome were produced by species-specific expansion of the gene family [ 31 ]. Maize underwent three genome-wide replication events occurred, including approximately 110 million years ago before the differentiation of monocotyledons and dicotyledons, before the emergence of Gramineae 50 million years ago, and the genome-wide replication event after the differentiation of maize and sorghum 12 million years ago [ 32 , 33 ]. Three ways of gene family expansion and doubling were found: whole-genome duplication (WGD), tandem duplication (TD), and segmental duplication (SD) [ 34 ]. WGD is a massive chromosome doubling event that increases the dose of all genes of a species at once, resulting in a large number of chromosomally doubled segments retained in the genome. Tandem duplication occurs frequently in chromosomal recombination domains, where members of tandemly duplicated gene families are typically tightly aligned on the same chromosome, forming a cluster of genes with related sequences and functions [ 35 ]. Segmental duplication occurs when duplicated genes are distant or even located on different chromosomes. In this study, multiple gene replication events were identified, including eight pairs of tandem replication gene pairs ( ZmSTP5 / 6 , ZmSTP21 / 22 , ZmPMT5 / 6 , ZmPMT11 / 12 , ZmPMT12 / 13 , ZmPMT15 / 14 , ZmPMT15 / 16 , and ZmERD3 / 4 ) with highly similar sequences in adjacent positions of chromosomes and twelve pairs of segmental duplication genes ( ZmSTP12 / 5 , ZmSTP5 / 14 , ZmSTP8 / 17 , ZmSTP14 / 21 , ZmSTP12 / 14 , ZmSTP12 / 21 , ZmPMT8 / 14 , ZmPMT14 / 3 , ZmPMT1 / 18 , ZmpGlcT1 / 4 , ZmERD2 / 5 , ZmERD6 / 3 ). The results showed that gene tandem duplication and chromosome segmental duplication are the main forms of monosaccharide transporter replication in maize. Gene family expansion and doubling can provide new adaptability for plant growth and development to resist biotic and abiotic stress, leading to gene functional diversity, and affecting the evolution process of species [ 36 ]. Ka/Ks analysis was used to determine the relative divergence time and whether the functional differentiation of replication genes was subject to selection pressure. According to previous studies, Ka > > Ks or Ka/Ks > > 1, Ka = Ks or Ka/Ks = 1, Ka/Ks or Ka/Ks < < 1, and Ka/Ks or Ka/Ks < < 1 denoted that the gene was susceptible to positive selection, neutral evolution, and purifying selection, respectively [ 37 , 38 ]. In this study, the Ka/Ks of all the duplicated gene pairs were calculated, and most of them showed less than one, indicating that they were subjected to purifying selection in the process of evolution. Five gene pairs subjected to strong positive selection were ZmSTP5 / ZmSTP12 , ZmERD2 / ZmERD5 , and ZmERD3 / ZmERD6 for segmental duplication and ZmSTP5 / ZmSTP6 and ZmSTP3 / ZmSTP4 for tandem duplication. This indicated that they were positively selected and rapidly evolved genes in a short period, and gene functions may have diverged. Of these three gene pairs belong to the ERD subfamily, suggesting that the ERD subfamily may be more important for maize to respond to environmental change. Synteny analysis was performed to analyze the expansion of the MST family between species. There were 47, 41, and 33 collinear gene pairs identified between maize and sorghum, foxtail millet and rice, respectively. Only one collinear pair between maize and alfalfa ( ZmpGlcT4 / AES80568 ) and ZmpGlcT4 also formed collinear pairs with SORBI_3003G084000 and Os01g0133400 between maize and sorghum, rice, respectively. These results showed a closer evolutionary relationship between the two species containing more collinear gene pairs, and most of the no-collinear genes may be produced in earlier replication events. Gene structure and conserved motif analyses were performed to further explore the evolutionary relationship in the MST family of maize. MST genes could be divided into seven subfamilies, and the PMT, STP, and TMT subfamilies contained fewer exons and simpler gene structures, while the pGlcT, ERD, INT, and VGT subfamilies had more exons and more complex gene structures, and gene structures were conserved in the same subfamily (Fig. 3 ). Motifs 1–6 were present in almost all MST family members and were vital for transport function and membrane localization, suggesting that these motifs were highly conserved domains during the evolution of the MST family. Other motifs existed in different subfamilies, indicating that these subfamily members transport different substrates. In addition, STP subfamily members contained all motifs except for motif 11, which implied a broader transport capability for different substrates. The effect of genes on plants depends not only on the function of genes themselves, but also on the regulation of gene expression. The cis -elements in the promoter region were involved in the regulation of the gene expression, and therefore the cis -element analysis was important for the preliminary prediction of gene expression. The type and number of cis -elements in promoters of MST family genes were identified in this study. The ABA response element ABRE belongs to the phytohormone response element and exists in most MST member promoter regions, meaning that the expression of MST members might be involved in the ABA signaling pathway. Additionally, many stress response elements were also found in the promoter region of MST members, such as DRE, MYB, and MYC, suggesting that MST family genes have an important role in maize response to environmental stresses. To further investigate the functional response to environmental stresses, the expression mode of MST family genes in the maize seedling stage was detected under different stress treatments. The expression of half of MST genes was significantly induced under PEG treatment, while fewer genes were induced by salt and Cd stresses (Fig. 7 A, B, and C). Meanwhile, the response of each subfamily in MST to different stresses was also different. Most of the PMT and ERD members responded to osmotic stress, and all ERD members were induced to expression by Cd stresses. The expression of most members was not induced in the STP subfamily under osmotic or ionic stresses. The expression patterns of MST members in response to stress treatments suggested that each subfamily might have different roles in balancing maize growth and development and responding to abiotic stress, implying that the members in each subfamily had also shown functional divergence during evolution and family expansion. ABA is a very important phytohormone in the plant response to abiotic stresses, and many stress response genes are regulated by ABA signaling [ 39 ]. In this study, the expression levels of some MST family members were induced by both ABA treatment and drought stress, indicating that these members might respond to stress in the ABA signaling pathway, while others were only induced by drought stress, meaning that they might respond to stress in the non-ABA signaling pathway. Together, we have identified maize MST family genes. Our findings could contribute to future research on maize MST family genes and provide the foundation for additional investigation of the fundamental functions of this significant monosaccharide transporter family. These findings provide insight into the possible roles of genetic improvement in the capacity of maize to respond to abiotic challenges and may be used to identify relevant candidate MST family genes for functional research. Conclusions In summary, a total of 66 ZmMST genes were identified from maize and divided into seven subfamilies. Phylogenetic tree, synteny, and collinearity analyses provide preliminary insights into dissecting the evolution and expansion of the MST family. Meanwhile, the expression analysis provides valuable clues for exploring the potential function of MST in balancing maize growth and development and abiotic stress response. These findings will be helpful for us to deeply understand the functions of maize MST genes and provide some important information for functional analysis in the future. Methods Identification and Evolutionary Analysis The complete amino acid and nucleotide sequences of Zea mays B73 RefGen_v4 were downloaded from MaizeGDB ( https://maizegdb.org/ ). In addition, MaizeGDB ( https://maizegdb.org/ ) transcriptome data were obtained. Seventy-nine different samples comprised the maize inbred line B73 RNA-seq gene map [ 40 ]. The transcriptome information was chosen from eight distinct maize tissues and developmental stages. From Ensembl ( https://asia.ensembl.org/index.html ), protein sequences for the rice MST and the Arabidopsis MST were acquired. The hidden Markov model repository was built using known MST protein sequences, and HMMER ( http://hmmer.org/ ) was used to query the maize dataset [ 41 ]. By utilizing MST genes from rice and Arabidopsis as queries in a BLAST search, MST genes from maize were investigated. Using the PFAM ( http://pfam.xfam.org/ ) and CDD ( https://www.ncbi.nlm.nih.gov/cdd/ ) databases, the conserved domains of the discovered ZmMST genes were predicted [ 42 , 43 ]. Using MEGA 7.0 ( https://www.megasoftware.net/ ) and ClustalW software ( https://www.genome.jp/tools-bin/clustalw ), evolutionary trees were created for the MST proteins from Arabidopsis, rice, and maize (with 1000 bootstrap replicates) [ 44 , 45 ]. Using a MapChart ( http://mg2c.iask.in/mg2c_v2.0/ ) and the chromosomal start and termination data received from MaizeGDB ( https://maizegdb.org/ ), the chromosomal locations of ZmMST genes were determined [ 46 ]. Tandem duplicated genes were found using the tools for multiple covariance scanning (MCScanX, http://chibba.pgml.uga.edu/mcscan2/MCScanX.zip ) [ 47 ]. Sequence Analysis The molecular weight (MW) and isoelectric point (pI) of the ZmMST proteins were predicted using the ExPASy proteomics system ( http://web.expasy.org/protparam/ ) (Table 1 ) [ 48 ]. The conserved protein motifs of ZmMST genes were discovered using MEME Suite ( http://meme-suite.org/ ), and they were further annotated with TBtools ( https://github.com/CJ-Chen/TBtools ) [ 49 , 50 ]. Fifteen motifs, with lengths ranging from 6 to 50 bp, made up the domain structure (Additional file 14). Using GSDS ( http://gsds.gao-lab.org ), the gene structure was evaluated [ 51 ]. The PlantCARE database ( http://bioinformatics.psb.ugent.be/webtools/plantcare/html/ ) was used to predict the 1500 bp sequence upstream of the cis -acting components of the coding sequences (Additional file 15). Further examination was performed on the components (ABRE, DRE, LTRE, ERE, and MBS) connected to the abiotic stress response [ 52 , 53 ]. Replication Events and Ka/Ks Analysis of MST Genes MCScanX ( http://chibba.pgml.uga.edu/mcscan2/MCScanX.zip ) was used to analyze correlations between ZmMST genes and single or multiple intergenomic variables as well as associations within genomes [ 54 ]. Gene family expansion and doubling can occur through several different processes, including whole-genome duplication or polyploidization, tandem duplication, segmental duplication, transposon-mediated transposon duplication, and retro-position. Finally, a graph of the intragenomic duplication events and gene density findings was created using TBtools software ( https://github.com/CJ-Chen/TBtools ) (Fig. 4 ). ClustalW ( https://www.genome.jp/tools-bin/clustalw ) was used to determine the ratio of Ka (non-synonymous substitution rate) and Ks (synonymous substitution rate) to investigate the selection pressure on the ZmMST family. The time of occurrence of segmental duplication events for homologous genes was calculated as T = Ks/2λ × 10 − 6 , where λ is the rate of molecular substitution in grasses (6.5×10 − 9 ), and expressed as a million years ago (Mya) [ 55 ]. Plant Treatments and Quantitative Real-Time PCR Analysis The autogamous maize cultivar “inbred line” was used in the study. Seeds were preserved in our laboratory and incubated in the seedling culture room of the Laboratory of Plant Physiology and Germplasm, Shenyang Agricultural University. The seeds were disinfected with 75% ethanol and washed with distilled water after 1 min to remove the residual ethanol. The cleaned seeds were evenly sown in seedling pots and irrigated with distilled water to allow the vermiculite to absorb sufficient water. Hoagland’s nutrient solution (pH 6.0) was added to the basal tray in which the seedling pots were placed to ensure that the roots could access the nutrient solution. The nutrient solution was replaced every 3 days until the seedlings attained the three-leaf stage [ 56 ]. The three-leaf seedlings were then treated with drought, salt, Cd stresses, and exogenous ABA by application of half-strength Hoagland’s nutrient solution supplemented with 20% PEG for drought stress treatment, 200 mol/L NaCl for salt stress treatment, 40 mg/L CdCl 2 for Cd stress treatment, and 100 µmol/L ABA for ABA stress treatment. The uppermost mature leaves were collected at 0, 6, 12, and 24 h after initiation of the stress treatment, with three biological replicates at each time point. The tested leaves were immediately frozen in liquid nitrogen and kept at -80°C. Total RNA isolation and quantitative real-time PCR (qRT-PCR) analysis were performed to analyze the expression of maize genes under salt, drought, Cd stress and exogenous ABA treatment [ 57 ]. A total of 61 maize MST genes were used for the analysis. Total RNA from plant leaves was extracted using TRIzol Reagent (CW Biotech) and subjected to DNase I treatment to remove genomic DNA contamination. The RNA concentration was determined utilizing a BioDrop ultramicro ultraviolet nucleic acid assay. First-strand cDNA was synthesized from 1 µg of total RNA using the UEIris II RT-PCR system. qRT-PCR assays were performed using a real-time PCR analyzer (Bio-Rad, Applied Biosystems PCR, SCILOGEX Gradient Thermal Cycler PCR Instrument TC1000-G). Each reaction mixture contained 10 µL of 2×SYBR® Green Pro Taq HS Premix, 1.0 µL cDNA sample, 0.4 µL forward primer (final concentration 10 µM), and 0.4 µL reverse primer (final concentration 10 µM) in a final volume of 20 µL. The thermal-cycling protocol was as follows: 95°C for 5 min, then 45 cycles of 95°C for 15 s and 60°C for 1 min. Melting curve analysis was used to verify the specificity of the reaction. Three technical replicates of each cDNA sample were analyzed. The Zm00001d013367 genes were selected as an internal control to normalize the transcript levels of ZmMST genes. The relative gene expression levels were calculated using the 2 −∆∆CT method (Additional file 16). The normalized data were processed with TBtools and plotted as a heatmap to visualize the changes in MST gene expression ( https://github.com/CJ-Chen/TBtools ) [ 58 ]. Row-scale and log-scale normalization calculations and row clustering were performed on the heatmaps. The number of genes with relative gene expression greater than 2 and relative gene expression less than 0.5 under the four treatments were shown by using the online site Venny 2.1.0 ( https://bioinfogp.cnb.csic.es/tools/venny/ ) to create Venn diagrams. All primer pairs were designed with Primer (v5.0) software ( http://www.broadinstitute.org/ftp/pub/software/Primer5.0/ ) and were listed in Additional file 17. Abbreviations Full Name Abbreviated Name Monosaccharide transporter MST major facilitator superfamily MFS sucrose transporter SUT Sugar Transport Protein STP Polyol/Monosaccharide Transporter PMT Vacuolar Glucose Transporter VGT Inositol Transporter INT Plastidic Glucose Transporter pGlcT Tonoplast Membrane Transporter TMT Early-Responsive to Dehydration six-like ERD chromosome chr whole-genome duplication WGD molecular weight MW isoelectric point pI non-synonymous substitution rate Ka synonymous substitution rate Ks million years ago Mya quantitative real-time PCR qRT-PCR Abscisic Acid ABA salicylic acid SA methyl jasmonate MeJA Declarations Acknowledgements We are grateful to all of the lab members who contributed helpful technical support and data analysis. "The College of Bioscience and Biotechnology Shenyang Agricultural University and Shenyang City Key Laboratory of Maize Genomic Selection Breeding" have our sincere gratitude. We also value the suggestions made for improvements by the reputable editors and reviewers of the text. Authors' contributions Conceptualization, JingJuan Fan and Cong Li; methodology, YanShu Zhu; validation, JiaLun Zhu, TianFeng Li, Jing Ma, and WenYu Li; formal analysis, JiaLun Zhu, TianFeng Li, and HanYu Zhang; data curation, JiaLun Zhu; writing-original draft preparation, JiaLun Zhu and TianFeng Li; writing-review and editing, Cong Li; visualization, Tsyganova Nadezhda and XiaoMei Dong; supervision, YanShu Zhu; project administration, JingJuan Fan; funding acquisition, JingJuan Fan. All authors have read and agreed to the published version of the manuscript. Funding This work was supported by the international cooperation program for universities in Liaoning Province Department of Education, China (No. 2023GJ0008), the scientific research projects of Liaoning Provincial Department of Education, China (No. LJKMZ20221010). Availability of data and materials Not applicable. 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Supplementary Files Additionalfile1.xlsx Additionalfile2.pdf Additionalfile3.pdf Additionalfile4.pdf Additionalfile5.pdf Additionalfile6.pdf Additionalfile7.pdf Additionalfile8.pdf Additionalfile9.xlsx Additionalfile10.xlsx Additionalfile11.pdf Additionalfile12.pdf Additionalfile13.pdf Additionalfile14.xlsx Additionalfile15.xlsx Additionalfile16.xlsx Additionalfile17.xlsx Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 17 Mar, 2024 Reviews received at journal 16 Mar, 2024 Reviews received at journal 11 Mar, 2024 Reviewers agreed at journal 05 Mar, 2024 Reviews received at journal 14 Sep, 2023 Reviewers agreed at journal 01 Sep, 2023 Reviewers agreed at journal 29 Aug, 2023 Reviewers invited by journal 26 Aug, 2023 Editor invited by journal 18 Aug, 2023 Editor assigned by journal 13 Aug, 2023 Submission checks completed at journal 04 Aug, 2023 First submitted to journal 31 Jul, 2023 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3219446","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":224015687,"identity":"7738501f-95c8-43a6-b180-bf4f1a937336","order_by":0,"name":"Jialun Zhu","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Jialun","middleName":"","lastName":"Zhu","suffix":""},{"id":224015688,"identity":"406e8759-38e7-4c2d-81e5-5808b54c5e7b","order_by":1,"name":"Tianfeng Li","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Tianfeng","middleName":"","lastName":"Li","suffix":""},{"id":224015689,"identity":"904d3c8c-cc20-47b6-8353-a6b4ac8e9f76","order_by":2,"name":"Jing Ma","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Jing","middleName":"","lastName":"Ma","suffix":""},{"id":224015690,"identity":"5291fae1-3f1a-4444-bd23-7ad41a2e57d1","order_by":3,"name":"Wenyu Li","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Wenyu","middleName":"","lastName":"Li","suffix":""},{"id":224015691,"identity":"3a0a6fa7-7495-49d8-8771-b706fa6ea706","order_by":4,"name":"Hanyu Zhang","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Hanyu","middleName":"","lastName":"Zhang","suffix":""},{"id":224015692,"identity":"61e75ffd-4649-4fec-b23f-53d7acc1b470","order_by":5,"name":"Tsyganova Nadezhda","email":"","orcid":"","institution":"Saint-Petersburg State Agrarian University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Tsyganova","middleName":"","lastName":"Nadezhda","suffix":""},{"id":224015693,"identity":"5bc13fec-f0f0-4bf4-8ca8-1597b15d5771","order_by":6,"name":"Yanshu Zhu","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yanshu","middleName":"","lastName":"Zhu","suffix":""},{"id":224015694,"identity":"086e7627-a357-4009-bad2-4d7d828327a5","order_by":7,"name":"Xiaomei Dong","email":"","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xiaomei","middleName":"","lastName":"Dong","suffix":""},{"id":224015695,"identity":"82b06490-de92-47c4-a5d3-6c863530f00a","order_by":8,"name":"Cong Li","email":"","orcid":"","institution":"Shenyang City Key Laboratory of Maize Genomic Selection Breeding","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Cong","middleName":"","lastName":"Li","suffix":""},{"id":224015696,"identity":"a0b7277f-c6ca-48a6-901e-8efba6953ff8","order_by":9,"name":"Jinjuan Fan","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2ElEQVRIiWNgGAWjYBACxmYGBgMgzc8vAeZLyBCtRXLmDAbGBqAWHqJtk9xwA6yFgbAW5nbeA8U8NXckjG83H390o8aCh4H98NEN+B3Gl2DMc+yZhNmdY4nNOceADuNJS7uBXwuPgTEP2+E6sxs5hs05bEAtEjxmRGj5d1jCeAZIyz9itfC2HZYwkABqyW0jUovh3L7DEhI30hJn5/ZJ8LAR8oth/xkzgzffDkvwz0g+8DnnW50cP/vhY/i1NDCwGaCIsOFTDgLywKh5QEjRKBgFo2AUjHAAADT8Qr2VMDGqAAAAAElFTkSuQmCC","orcid":"","institution":"College of Bioscience and Biotechnology, Shenyang Agricultural University","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Jinjuan","middleName":"","lastName":"Fan","suffix":""}],"badges":[],"createdAt":"2023-07-31 06:44:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3219446/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3219446/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":41308910,"identity":"a921b170-1cf6-44b8-a682-59d7efa20ffb","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":303042,"visible":true,"origin":"","legend":"\u003cp\u003eChromosomal distribution of ZmMST genes. ZmPMT genes, ZmpGlcT genes, ZmERD genes, ZmSTP genes, ZmINT genes, ZmVGT genes, and ZmTMT genes are highlighted in purple, yellow-green, red, dark brown, dark blue, dark green, and light blue, respectively. Tandem repeat genes are highlighted in yellow.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/f50c4218390b94d7aee19058.png"},{"id":41309894,"identity":"9d3fa002-846b-4e40-8dfd-c6ac7592d9d3","added_by":"auto","created_at":"2023-08-09 14:22:18","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":916376,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree for MST proteins of \u003cem\u003eZea\u003c/em\u003e \u003cem\u003emays\u003c/em\u003e. Multiple sequence alignment of the MST domains was performed using MUSCLE, and the phylogenetic tree was constructed using MEGA 7.0 with the maximum likelihood method with 1000 bootstrap replicates. Proteins of ZmPMT, ZmpGlcT, ZmERD, ZmSTP, ZmINT, ZmVGT, and ZmTMT are highlighted in purple, yellow-green, red, dark brown, dark blue, dark green, and light blue, respectively.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/20b63e3428d6f5faa3c4009a.png"},{"id":41308904,"identity":"8facba91-5745-4756-b8ec-e11cbe101b95","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":714433,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic relationships, gene structure, and conserved motif analyses of ZmMST genes based on phylogenetic relationships. All motifs were identified with the MEME Suite using the complete amino acid sequences. Exon-intron structure analyses were performed with TBtools. \u003cstrong\u003eA\u003c/strong\u003eNeighbor-joining tree indicating evolutionary relationships. \u003cstrong\u003eB\u003c/strong\u003eExon-intron structure. Green boxes, yellow boxes, and black lines indicate the untranslated region, coding sequence, and gene length, respectively. \u003cstrong\u003eC\u003c/strong\u003eConserved motifs.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/5baab395a534d88a0387a997.png"},{"id":41308907,"identity":"b9964740-6f8f-4f42-9e47-9eaff9d2425d","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2967172,"visible":true,"origin":"","legend":"\u003cp\u003eSynteny analysis of the MST family in maize. The two outermost circles are the density of the maize genome, the third circle is the numbering of maize chromosomes, the fourth circle is the ID of genes with segmental repeat relationships, and the innermost circle is the result of the analysis of covariance within the maize genome. Red curves linking ZmMST genes indicate duplicated gene pairs in the ZmMST family.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/ab4997952cc245e605cc9db1.png"},{"id":41309896,"identity":"37bbf4d9-d3cf-4136-a5fb-5797cdc4f243","added_by":"auto","created_at":"2023-08-09 14:22:19","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":6923907,"visible":true,"origin":"","legend":"\u003cp\u003eSynteny analysis of MST genes between maize and three representative gramineous species and \u003cem\u003eMedicago truncatula\u003c/em\u003e. \u003cem\u003eZ\u003c/em\u003e.\u003cem\u003emays\u003c/em\u003e, \u003cem\u003eS\u003c/em\u003e.\u003cem\u003ebicolor\u003c/em\u003e, \u003cem\u003eS\u003c/em\u003e.\u003cem\u003eitalica\u003c/em\u003e, \u003cem\u003eO\u003c/em\u003e.\u003cem\u003esativa\u003c/em\u003e, and \u003cem\u003eM\u003c/em\u003e.\u003cem\u003etruncatula\u003c/em\u003eindicate \u003cem\u003eZea\u003c/em\u003e \u003cem\u003emays\u003c/em\u003e, \u003cem\u003eSorghum bicolor\u003c/em\u003e, \u003cem\u003eSetaria italica\u003c/em\u003e, \u003cem\u003eOryza sativa\u003c/em\u003e, and \u003cem\u003eMedicago truncatula\u003c/em\u003e, respectively\u003cem\u003e. \u003c/em\u003eGray lines in the background show collinear blocks in the genomes of maize and sorghum, foxtail millet, rice, and \u003cem\u003eMedicago truncatula, \u003c/em\u003eand red lines highlight the collinear MST gene pairs.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/facdc5cd2a8d523d63a73cab.png"},{"id":41308911,"identity":"df372758-2bb0-48f5-81c7-d558dcc313ec","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":3725156,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eCis\u003c/em\u003e-acting elements in the ZmMST gene family. \u003cstrong\u003eA\u003c/strong\u003e Number of different elements in the promoter region of the ZmMST genes, as indicated by different color intensities and numbers in the grid. \u003cstrong\u003eB\u003c/strong\u003e Total number of \u003cem\u003ecis\u003c/em\u003e-acting elements in each response category. Sandy-brown indicates abiotic and biotic stresses, goldenrod indicates phytohormones, and chocolate represents plant growth and development.\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/832c81b90bde3c61877bf39f.png"},{"id":41308922,"identity":"adac674f-ad08-4e89-b714-f0d7edb4ab7b","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":5924735,"visible":true,"origin":"","legend":"\u003cp\u003eExpression analysis of ZmMST genes in response to abiotic stresses and exogenous ABA. \u003cstrong\u003eA\u003c/strong\u003e The expression of ZmMST genes at 0, 6, 12, and 24 h under PEG treatment. \u003cstrong\u003eB\u003c/strong\u003e The expression of ZmMST genes at 0, 6, 12, and 24 h under NaCl treatment. \u003cstrong\u003eC\u003c/strong\u003e The expression of ZmMST genes at 0, 6, 12, and 24 h under Cd treatment. \u003cstrong\u003eD\u003c/strong\u003e The expression of ZmMST genes at 0, 6, 12, and 24 h under ABA treatment. Red represents induced expression; blue represents repressed expression. The 0 h relative expression level of each treatment was taken as 1.\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/a69f6361d29c065e1e68e7f1.png"},{"id":41308925,"identity":"e9aadfb9-ce45-4d22-b27a-28707a5af4a7","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":310542,"visible":true,"origin":"","legend":"\u003cp\u003eExpression profiles of ZmSTP genes in different tissues. Log\u003csub\u003e2\u003c/sub\u003e-based fold change data were used to create the heatmap. Fold changes in gene expression are indicated by the color scale. DAS: Days of growth after sprouting. DAP: Days after pollination.\u003c/p\u003e","description":"","filename":"8.png","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/93e98649eec283e585e8311e.png"},{"id":41310765,"identity":"794ca8cb-7213-4ad5-b465-4f84e511190a","added_by":"auto","created_at":"2023-08-09 14:38:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":6371201,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/fa5a0237-47f3-4eeb-9dba-a976dc52c076.pdf"},{"id":41308902,"identity":"1a303d3a-6436-44bc-897c-8ce4594c34ca","added_by":"auto","created_at":"2023-08-09 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14:14:19","extension":"pdf","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":430713,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile13.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/d2e67430c0a09e3bef3df96b.pdf"},{"id":41308927,"identity":"92883904-71aa-4879-b52a-41ba53f07e5b","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"xlsx","order_by":14,"title":"","display":"","copyAsset":false,"role":"supplement","size":10335,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile14.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/2633c1282336f5b64eafcf2b.xlsx"},{"id":41308917,"identity":"aaf0491e-7a37-40d8-850d-c0d73d04d774","added_by":"auto","created_at":"2023-08-09 14:14:19","extension":"xlsx","order_by":15,"title":"","display":"","copyAsset":false,"role":"supplement","size":21359,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile15.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/48df487309d2e8b562a601f0.xlsx"},{"id":41310270,"identity":"c0c29af3-0f74-4b41-818e-56f0885a906b","added_by":"auto","created_at":"2023-08-09 14:30:19","extension":"xlsx","order_by":16,"title":"","display":"","copyAsset":false,"role":"supplement","size":39784,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile16.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/975298fefc7da16ff45211a7.xlsx"},{"id":41309931,"identity":"5fc9a1cd-bede-4572-bca7-64d169de38fd","added_by":"auto","created_at":"2023-08-09 14:22:19","extension":"xlsx","order_by":17,"title":"","display":"","copyAsset":false,"role":"supplement","size":12965,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile17.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-3219446/v1/4b239cd624dd6b542ffb2d0a.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Genome-Wide Identification and Investigation of MST Gene Family Based on Their Evolution and Expression Analysis under Abiotic Stress and Hormone Treatments in Maize (Zea mays L.)","fulltext":[{"header":"Background","content":"\u003cp\u003eThe industrial activities of human caused the trends of climate warming and constant changes in the natural environment [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. Many abiotic stresses such as drought, salinity, low temperature, and heavy metals pollution affect the growth and development of plants and the yield of crops, and further threaten food security and human health[\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. Sugars are the main form of long-distance transportation and distribution of photosynthate and play an important role in plant growth and development [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. Sugar not only constitutes metabolites, nutrients, and signal molecules, but also can be used as osmotic substances. One way that plants protect cell structure and alleviate the damage of abiotic stress is by accumulating osmotic substances or compatible solutes [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. In \u003cem\u003eArabidopsis thaliana\u003c/em\u003e, sugar plays an important role in protecting plant structure by increasing the concentration of soluble sugars (sucrose, glucose and fructose) in sink leaves and sucrose in phloem sap, maintaining water potential and improving plant stress tolerance [\u003cspan additionalcitationids=\"CR7\" citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. Therefore, sugar transport and distribution are key for crops in yielding and coping with abiotic stress [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e, \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eIn plants, various sugar transporters are involved in the transport of sugar from source to sink, of which the major facilitator superfamily (MFS) is an important member [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. MFS can be generally divided into the sucrose transporter (SUT) family and MST family according to the different transport substrates. MSTs are complete membrane proteins that can participate in the transmembrane transport of monosaccharides [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. The MST family can also be further divided into seven subfamilies based on their substrate specificities and sequence features, including Sugar Transport Protein (STP), Polyol/Monosaccharide Transporter (PMT), Vacuolar Glucose Transporter (VGT), Inositol Transporter (INT), Plastidic Glucose Transporter (pGlcT), Tonoplast Membrane Transporter (TMT), and Early-Responsive to Dehydration six-like (ERD) [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eSTP is a sugar transporter that mainly transports hexose. In Arabidopsis, most STPs exhibited a broad spectrum of absorption characteristics of substrates. AtSTP1 was the function of sugar transport and expressed in germinating seeds and the roots of seedlings, which played a key role in germination and root development [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. In \u003cem\u003eatstp1\u003c/em\u003e mutant, the ability to transport sugar alcohols (D-glucose, D-galactose, and D-mannose) was greatly weakened compared with the wild type [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. By investigating \u003cem\u003eAtSTP6\u003c/em\u003e-promoter::\u003cem\u003eGUS\u003c/em\u003e plants and conducting in situ hybridization experiments at the late stage of pollen development, it was determined that \u003cem\u003eAtSTP6\u003c/em\u003e was expressed. A transposon-tagged Arabidopsis mutant also demonstrated that the \u003cem\u003eatstp6\u003c/em\u003e mutation may have an impact on pollen vitality, pollen germination, fertilization, and seed production [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. PMT proteins (PMTs) not only transport mannitol, sorbitol, xylitol, and other polyols, but also transport monosaccharides. The first PMT gene was found in \u003cem\u003eApium graveolens\u003c/em\u003e L., named \u003cem\u003eAgMAT1\u003c/em\u003e, and it played an important role in the loading process of phloem mannitol [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. Arabidopsis genome contained six PMT subfamily genes, named \u003cem\u003eAtPMT1\u003c/em\u003e to \u003cem\u003eAtPMT6\u003c/em\u003e. \u003cem\u003eAtPMT5\u003c/em\u003e was located in the plasma membrane and could transport pentoses such as sugar alcohols (sorbitol, xylitol, erythritol, and glycerol), hexose, and ribose. \u003cem\u003eAtPMT5\u003c/em\u003e was highly expressed in Arabidopsis roots and played an important role in plant morphological construction [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. In plants, the VGT subfamily generally had only 2\u0026ndash;3 members, and played a key role in the process of seed germination, flowering, and other growth and development [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. In Arabidopsis, AtVGT1 was H\u003csup\u003e+\u003c/sup\u003e/glucose reverse transporter located on the vacuolar plastid, which was involved in the transport and storage of monosaccharides in vacuoles. AtVGT1 also played a key role in the process of seed germination, flowering, and other growth and development, and the \u003cem\u003eatvgt1\u003c/em\u003e mutant had a lower germination rate and delayed flowering [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. INT proteins were highly specific H\u003csup\u003e+\u003c/sup\u003e-inositol symporters, and the functions of them were in the transportation and distribution of inositol. Four INT genes (\u003cem\u003eAtINT1\u003c/em\u003e-\u003cem\u003e4\u003c/em\u003e) were identified in Arabidopsis, of which \u003cem\u003eAtINT4\u003c/em\u003e was the first identified INT subfamily member. \u003cem\u003eAtINT4\u003c/em\u003e was highly expressed in Arabidopsis pollen and phloem companion tissue, which was related to plant pollen development and participated in inositol loading in phloem [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. pGlcT proteins (pGlcTs) could transport glucose. The pGlcT gene was cloned in spinach (\u003cem\u003eSpinacia oleracea\u003c/em\u003e L.) and proved the role in transporting starch and hydrolyzing glucose [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. In Arabidopsis, the \u003cem\u003eatpglct1\u003c/em\u003e, \u003cem\u003eatpglct2\u003c/em\u003e, and \u003cem\u003eatpglct1\u003c/em\u003e/\u003cem\u003eatpglct2\u003c/em\u003e mutants showed growth and development inhibition in varying degrees [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. TMT proteins (TMTs, also known as ATZ) were also localized on vacuolar plastids. Previous studies reported that TMTs are involved in the transport of sucrose on vacuoles. In the \u003cem\u003eAttmt1\u003c/em\u003e/\u003cem\u003eAttmt2\u003c/em\u003e double mutant, the sucrose transportation to vacuoles was impaired [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. The majority of the ERD6-like protein was found on the vacuolar membrane and was in charge of the transmembrane transport of sugar in the vacuoles [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. In Arabidopsis, ERD (also known as SFPs) is the largest subfamily in the MST family and responds to abiotic stresses. For instance, \u003cem\u003eAtERD1\u003c/em\u003e and \u003cem\u003eAtERD2\u003c/em\u003e also responded to different stress treatments, and the expression patterns of them showed significantly different [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. Genome-wide identification of the MST family had been performed in many plants, such as Arabidopsis, rice, grape, and tobacco [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan additionalcitationids=\"CR27\" citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe distribution of monosaccharide transporters in maize plants directly affects plant growth and development and abiotic stress response. However, genome-wide identification, evolutionary analysis, and response to abiotic stresses of the maize monosaccharide transporter family have not yet been reported. In this study, 66 MST genes were identified from the maize genome. Phylogenetic analysis, gene structure analysis, and synteny analysis were performed to understand the evolution and amplification of the ZmMST family, and the expression analysis in different tissues and under different treatments was performed to explore the response to abiotic stresses. These results provide insights into the evolution of the maize MST family and their role in maize growth and development and abiotic stress response. The identification and characterization of ZmMST genes may provide opportunities for the optimization of maize variety selection and breeding.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eIdentification and phylogenetic analysis of the MST gene family members in maize\u003c/h2\u003e \u003cp\u003eThe Arabidopsis MST proteins were queried against the maize genome using BLASTP to search for maize MST genes. The gene domain was manually confirmed through the NCBI CDD, SMART, and Pfam websites, and the protein sequence length (number of amino acids), molecular weight, and isoelectric point were determined by the ExPASy proteomics system. The sequences of the monosaccharide transporter family revealed by screening have conserved structural domains. Finally, 66 complete monosaccharide transporter sequences were identified and divided into seven subfamilies in maize, including STP (23 members), PMT (20 members), VGT (2 members), INT (4 members), pGlcT (4 members), TMT (4 members), and ERD (9 members) subfamilies. Gene names, AA accession number, length of the gene, amino acid numbers, molecular weights, chromosomal locations, and pIs were listed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Meanwhile, 64, 69, and 77 genes of the MST family were identified in rice, sorghum, and millet, respectively, as shown in Additional file 1.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe identification of MST members in maize.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGene name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAA accession number\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLength of gene(bp)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNumber of amino acid(aa)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMolecular weight (Da)\u003c/p\u003e \u003c/th\u003e 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\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e524\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e56153.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e9.23\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr1:2429641Chr3:24298623\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmPMT3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e 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\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e449\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e46171.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e9.63\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr4:532866Chr1:5330696\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmPMT13\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e 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\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e517\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e55327\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e5.77\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr5:4670830Chr3:46712407\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmINT1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e 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\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e374\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e59874\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e6.46\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr4:601307Chr3:6018723\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmTMT3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d014872\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e2292\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e763\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e80956.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e4.46\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr5:6660522Chr8:66608036\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmTMT4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d016274\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1806\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e601\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e68874\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e5.73\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr5:15513335Chr4:155138132\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e11\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d029254\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1533\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e510\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e51871.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e5.74\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr1:6385984Chr5:63864152\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d040243\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1452\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e483\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e39799.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e8.96\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr3:3341091Chr8:33414667\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD3\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d039051\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1503\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e500\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e52615.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e9.25\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr6:16929398Chr5:169298831\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e9\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d039052\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1233\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e410\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e53240.9\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e9.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr6:16932079Chr3:169327039\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d008374\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1470\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e489\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e31835.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e10.37\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr8:669913Chr9:6703444\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d009600\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1125\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e374\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e54327.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e7.31\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr8:7249771Chr8:72504565\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e36\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d009603\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1491\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e496\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e53739.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e8.44\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr8:72763170Chr8:72767938\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e4\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d009605\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1497\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e498\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e44025.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e9.34\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr8:7286116Chr4:72868595\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e+\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eZmERD9\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003eZm00001d009669\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e1521\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e506\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e51148.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e8.31\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eChr8:7514623Chr3:75149733\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e-\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e10\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eAs shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, the MST genes were randomly distributed on 10 maize chromosomes. Among them, 12 MST genes were distributed on chromosome (chr) 7, and only two genes existed on chr 6 and 9. The STP subfamily genes were dispersed across 8 of the 10 chromosomes, and ZmPMT genes were dispersed across 6 of the 10 chromosomes. ZmVGT genes had only two genes and were located on chr 5. ZmTMT genes were located on chr 1, 4, and 5, respectively. ZmINT genes were located on chr 7 and 10, respectively. ZmpGlcT genes were located on chr 3, 4, 7, and 8, respectively. ZmERD genes were located on chr 1, 3, and 8.\u003c/p\u003e \u003cp\u003eTo explore the systematic evolution, a phylogenetic tree of MST family genes in maize was constructed. As shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA, ZmMST genes were divided into five main branches and seven groups, which was consistent with the results of existing studies. The VGT subfamily and INT subfamily possessed a closed relationship, while the relationship of ERD and pGlcT was close. Meanwhile, phylogenetic trees were constructed in seven subfamilies, including members from three species of maize, rice, and Arabidopsis. (Additional file 2\u0026ndash;8)\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eGene structure and motif composition of the maize MST gene family\u003c/h2\u003e \u003cp\u003eThe exon-intron organizations of the MST family genes were detected to comprehend the evolution of the MST family in maize. As shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB, ZmSTP genes possessed one to four exons (20 with two to three exons, 3 with four exons, and \u003cem\u003eZmSTP11\u003c/em\u003e with only one the most exon). Although the ZmPMT subfamily has many genes, the gene structure tends to be conserved, and all ZmPMT genes possess two to three exons (14 members with two exons and 6 members with three exons). The ZmVGT subfamily contains only two genes, \u003cem\u003eZmVGT1\u003c/em\u003e possesses twelve exons and \u003cem\u003eZmVGT2\u003c/em\u003e possesses fourteen exons. ZmINT and ZmTMT genes possesses one to six exons. ZmpGlcT genes possess 12 to 14 exons. The ZmERD subfamily is different from other subfamilies, which contain an extensive number of exons, from 7 to 18.\u003c/p\u003e \u003cp\u003eThe conserved motifs of MST proteins were identified by the MEME motif program. The result was shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC. Almost all MST family members contained six conserved domains, motifs 1 to 6, and each subfamily contained different conserved motifs. Except for the ERD and TMT subfamilies, the other five subfamilies all contain motif 7. Motif 8 exists in almost all subfamilies except the TMT and motif 11 exists in all subfamilies except the ERD and STP. While motifs 10, 13, 14, and 15 only exist in the STP subfamily. Overall, the gene structures and conserved motif compositions of MSTs were similar in the same subfamily.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eSynteny analysis of MST genes\u003c/h2\u003e \u003cp\u003eTo analyze the expansion of the MST family, gene duplication events, both tandem duplication (TD) and segmental duplication (SD) were detected. Eight pairs of tandem replication gene pairs (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), including \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e, \u003cem\u003eZmSTP21\u003c/em\u003e/\u003cem\u003e22\u003c/em\u003e, \u003cem\u003eZmPMT1\u003c/em\u003e/\u003cem\u003e2\u003c/em\u003e, \u003cem\u003eZmPMT2\u003c/em\u003e/\u003cem\u003e3\u003c/em\u003e, \u003cem\u003eZmPMT5\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e, \u003cem\u003eZmPMT14\u003c/em\u003e/\u003cem\u003e15\u003c/em\u003e, \u003cem\u003eZmPMT15\u003c/em\u003e/\u003cem\u003e16\u003c/em\u003e and \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e, and 12 segmental duplication events, including \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e13\u003c/em\u003e, \u003cem\u003eZmSTP6\u003c/em\u003e/\u003cem\u003e15\u003c/em\u003e, \u003cem\u003eZmSTP9\u003c/em\u003e/\u003cem\u003e18\u003c/em\u003e, \u003cem\u003eZmSTP13\u003c/em\u003e/\u003cem\u003e15\u003c/em\u003e, \u003cem\u003eZmSTP15\u003c/em\u003e/\u003cem\u003e22\u003c/em\u003e, \u003cem\u003eZmSTP13\u003c/em\u003e/\u003cem\u003e22\u003c/em\u003e, \u003cem\u003eZmPMT1\u003c/em\u003e/\u003cem\u003e8\u003c/em\u003e, \u003cem\u003eZmPMT3\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e, \u003cem\u003eZmPMT8\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e, \u003cem\u003eZmpGlcT1\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e, \u003cem\u003eZmERD2\u003c/em\u003e/\u003cem\u003e5\u003c/em\u003e, and \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e were identified in maize (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo further explore the evolutionary mechanisms of the MST family, the syntenic maps of maize were constructed and associated with four representative plant species, including the monocotyledons \u003cem\u003eSorghum bicolor\u003c/em\u003e, \u003cem\u003eSetaria italica\u003c/em\u003e, \u003cem\u003eOryza sativa\u003c/em\u003e, and the dicotyledon \u003cem\u003eMedicago truncatula\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). A total of 41 MST family genes in maize showed syntenic relationships with those in millet, followed by sorghum (47) and rice (33), and there was only one pair of homologous genes in alfalfa (on chr 8), and specific gene pairs are shown in Additional file 9.\u003c/p\u003e \u003cp\u003eTo better understand the evolutionary constraints acting on the MST gene family, the Ka/Ks ratios of the tandem and segmental duplications in MST gene pairs were calculated (Additional file 10). Five pairs of genes had Ka/Ks\u0026thinsp;\u0026gt;\u0026thinsp;1 (\u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e, \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e12\u003c/em\u003e, \u003cem\u003eZmERD2\u003c/em\u003e/\u003cem\u003e5\u003c/em\u003e, \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e, and \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e). The Ka/Ks of others are all less than 1.\u003c/p\u003e \u003cp\u003e \u003cb\u003eAnalysis of the\u003c/b\u003e \u003cb\u003ecis\u003c/b\u003e \u003cb\u003e-acting element in MST gene promoter regions\u003c/b\u003e \u003c/p\u003e \u003cp\u003eTo explore the potential regulatory mechanisms of MST family genes, the 1.5 kb upstream promoter region of MST genes was submitted into Plant CARE to detect the conserved \u003cem\u003ecis\u003c/em\u003e-elements. Then, the identified \u003cem\u003ecis\u003c/em\u003e-acting elements were divided into three categories: abiotic and biotic stress, phytohormone responsive, and plant growth and development (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the categories of abiotic and biotic stresses, MYB (CCAAT-box) and MYC (CACAT-box) were the two types of \u003cem\u003ecis\u003c/em\u003e-elements with the largest proportion in the MST family. Meanwhile, the anaerobic response element ARE, drought response element DRE, and low-temperature element LTR were also widely distributed in the promoter regions of MST genes. For phytohormone responsive elements, the ABRE, TCA, as-1, and CGTCA-motif which are involved in Abscisic Acid (ABA), salicylic acid (SA), and methyl jasmonate (MeJA) responses respectively were detected in the most of MST gene promoter regions. In addition, 18 \u003cem\u003ecis\u003c/em\u003e-elements related to plant growth and development were identified, most of which were involved in the light response, accounting for more than half of the group.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eExpression analysis of MST genes in different maize tissues and in response to different treatments\u003c/h2\u003e \u003cp\u003eTo further analyze the function of MST family members in response to abiotic stresses, the expression patterns of MST family genes were detected in maize under abiotic stresses and ABA treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e). Under PEG treatment, the expression levels of most MST members were increased, especially in the PMT, ERD, and VGT subfamilies. Under NaCl treatment, the expression of only five genes was induced, of which \u003cem\u003eZmPMT9\u003c/em\u003e was the most significantly increased. Under Cd stress, the transcript levels of \u003cem\u003eZmPMT8\u003c/em\u003e/12/13/\u003cem\u003e15\u003c/em\u003e/16/17/19/\u003cem\u003e20\u003c/em\u003e and \u003cem\u003eZmERD2\u003c/em\u003e/\u003cem\u003e3\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e were significantly increased. Abscisic acid is a phytohormone involved in regulating plant responses to abiotic stresses. The expression of most MST members was induced under exogenous ABA treatment. Overall, the expression of 61 ZmMST genes showed different alterations under the abiotic stresses and exogenous ABA treatment, and some genes were affected by multiple treatments (Additional file 11).\u003c/p\u003e \u003cp\u003eThe expression of MST genes in various tissues, including roots, stems, leaves, and seeds, was also examined (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e, Additional files 12 and 13). \u003cem\u003eZmSTP1\u003c/em\u003e, \u003cem\u003eZmSTP2\u003c/em\u003e, and \u003cem\u003eZmSTP5\u003c/em\u003e were relatively high and stable in various tissues and periods, while the expression levels of \u003cem\u003eZmSTP8\u003c/em\u003e/\u003cem\u003e9\u003c/em\u003e/\u003cem\u003e10\u003c/em\u003e/\u003cem\u003e11\u003c/em\u003e/\u003cem\u003e12\u003c/em\u003e/\u003cem\u003e13\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e/\u003cem\u003e16\u003c/em\u003e and \u003cem\u003e22\u003c/em\u003e were relatively low in various tissues. However, with the development process, the expression of \u003cem\u003eZmSTP2\u003c/em\u003e in roots, stems, leaves, and seeds were rapidly up-regulated, while \u003cem\u003eZmSTP5\u003c/em\u003e showed a trend of high expression in all tissues and stages. The expression patterns of members of the same subfamily were complementary. For example, \u003cem\u003eZmSTP19\u003c/em\u003e and \u003cem\u003eZmSTP23\u003c/em\u003e were highly expressed in leaves but almost not in seeds. \u003cem\u003eZmSTP23\u003c/em\u003e was highly expressed in leaves, but expressed at low levels in roots, stems, and seeds. \u003cem\u003eZmPMT1\u003c/em\u003e and \u003cem\u003eZmPMT13\u003c/em\u003e also had similar expression patterns. According to these results, ZmMST genes played a variety of roles in maize growth and development.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eSince monosaccharides such as glucose and fructose are essential for metabolism, storage, and transport, MST is crucial to the processes of carbon partitioning and abiotic stress response in plants [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]. Genome-wide analysis of the MST gene family has been widely carried out in many species. Fifty-three, sixty-four, and sixty-nine genes have been identified in Arabidopsis, rice, and sorghum respectively [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]. However, the MST gene family has not been identified in maize. In this study, 66 MST genes were identified in the maize genome, and 77 MST family genes were identified in millet (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and Additional file 1). By the phylogenetic tree results, ZmMST genes were further divided into 7 subfamilies [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. Each subfamily of MST was shown to be specific in the differentiation and evolution of the different Gramineae based on our findings that the number of genes in each subfamily of MST varied in maize, rice, sorghum, and millet. It was reported that the STP subfamily is the largest subfamily in rice. The ERD subfamily is the largest subfamily in Arabidopsis [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. In maize, the largest subfamily is also the STP family, which indicates that the MST family has species-specific subfamily expansion in different plants. These expansions may be caused by gene duplication events, which may play a key role in the evolution of the MST gene family.\u003c/p\u003e \u003cp\u003eThe previous study proved that most genes in the Arabidopsis genome were produced by species-specific expansion of the gene family [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]. Maize underwent three genome-wide replication events occurred, including approximately 110\u0026nbsp;million years ago before the differentiation of monocotyledons and dicotyledons, before the emergence of Gramineae 50\u0026nbsp;million years ago, and the genome-wide replication event after the differentiation of maize and sorghum 12\u0026nbsp;million years ago [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]. Three ways of gene family expansion and doubling were found: whole-genome duplication (WGD), tandem duplication (TD), and segmental duplication (SD) [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. WGD is a massive chromosome doubling event that increases the dose of all genes of a species at once, resulting in a large number of chromosomally doubled segments retained in the genome. Tandem duplication occurs frequently in chromosomal recombination domains, where members of tandemly duplicated gene families are typically tightly aligned on the same chromosome, forming a cluster of genes with related sequences and functions [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Segmental duplication occurs when duplicated genes are distant or even located on different chromosomes. In this study, multiple gene replication events were identified, including eight pairs of tandem replication gene pairs (\u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e, \u003cem\u003eZmSTP21\u003c/em\u003e/\u003cem\u003e22\u003c/em\u003e, \u003cem\u003eZmPMT5\u003c/em\u003e/\u003cem\u003e6\u003c/em\u003e, \u003cem\u003eZmPMT11\u003c/em\u003e/\u003cem\u003e12\u003c/em\u003e, \u003cem\u003eZmPMT12\u003c/em\u003e/\u003cem\u003e13\u003c/em\u003e, \u003cem\u003eZmPMT15\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e, \u003cem\u003eZmPMT15\u003c/em\u003e/\u003cem\u003e16\u003c/em\u003e, and \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e) with highly similar sequences in adjacent positions of chromosomes and twelve pairs of segmental duplication genes (\u003cem\u003eZmSTP12\u003c/em\u003e/\u003cem\u003e5\u003c/em\u003e, \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e, \u003cem\u003eZmSTP8\u003c/em\u003e/\u003cem\u003e17\u003c/em\u003e, \u003cem\u003eZmSTP14\u003c/em\u003e/\u003cem\u003e21\u003c/em\u003e, \u003cem\u003eZmSTP12\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e, \u003cem\u003eZmSTP12\u003c/em\u003e/\u003cem\u003e21\u003c/em\u003e, \u003cem\u003eZmPMT8\u003c/em\u003e/\u003cem\u003e14\u003c/em\u003e, \u003cem\u003eZmPMT14\u003c/em\u003e/\u003cem\u003e3\u003c/em\u003e, \u003cem\u003eZmPMT1\u003c/em\u003e/\u003cem\u003e18\u003c/em\u003e, \u003cem\u003eZmpGlcT1\u003c/em\u003e/\u003cem\u003e4\u003c/em\u003e, \u003cem\u003eZmERD2\u003c/em\u003e/\u003cem\u003e5\u003c/em\u003e, \u003cem\u003eZmERD6\u003c/em\u003e/\u003cem\u003e3\u003c/em\u003e). The results showed that gene tandem duplication and chromosome segmental duplication are the main forms of monosaccharide transporter replication in maize.\u003c/p\u003e \u003cp\u003eGene family expansion and doubling can provide new adaptability for plant growth and development to resist biotic and abiotic stress, leading to gene functional diversity, and affecting the evolution process of species [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]. Ka/Ks analysis was used to determine the relative divergence time and whether the functional differentiation of replication genes was subject to selection pressure. According to previous studies, Ka\u0026thinsp;\u0026gt;\u0026thinsp;\u0026gt;\u0026thinsp;Ks or Ka/Ks\u0026thinsp;\u0026gt;\u0026thinsp;\u0026gt;\u0026thinsp;1, Ka\u0026thinsp;=\u0026thinsp;Ks or Ka/Ks\u0026thinsp;=\u0026thinsp;1, Ka/Ks or Ka/Ks\u0026thinsp;\u0026lt;\u0026thinsp;\u0026lt;\u0026thinsp;1, and Ka/Ks or Ka/Ks\u0026thinsp;\u0026lt;\u0026thinsp;\u0026lt;\u0026thinsp;1 denoted that the gene was susceptible to positive selection, neutral evolution, and purifying selection, respectively [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. In this study, the Ka/Ks of all the duplicated gene pairs were calculated, and most of them showed less than one, indicating that they were subjected to purifying selection in the process of evolution. Five gene pairs subjected to strong positive selection were \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003eZmSTP12\u003c/em\u003e, \u003cem\u003eZmERD2\u003c/em\u003e/\u003cem\u003eZmERD5\u003c/em\u003e, and \u003cem\u003eZmERD3\u003c/em\u003e/\u003cem\u003eZmERD6\u003c/em\u003e for segmental duplication and \u003cem\u003eZmSTP5\u003c/em\u003e/\u003cem\u003eZmSTP6\u003c/em\u003e and \u003cem\u003eZmSTP3\u003c/em\u003e/\u003cem\u003eZmSTP4\u003c/em\u003e for tandem duplication. This indicated that they were positively selected and rapidly evolved genes in a short period, and gene functions may have diverged. Of these three gene pairs belong to the ERD subfamily, suggesting that the ERD subfamily may be more important for maize to respond to environmental change. Synteny analysis was performed to analyze the expansion of the MST family between species. There were 47, 41, and 33 collinear gene pairs identified between maize and sorghum, foxtail millet and rice, respectively. Only one collinear pair between maize and alfalfa (\u003cem\u003eZmpGlcT4\u003c/em\u003e/\u003cem\u003eAES80568\u003c/em\u003e) and \u003cem\u003eZmpGlcT4\u003c/em\u003e also formed collinear pairs with \u003cem\u003eSORBI_3003G084000\u003c/em\u003e and \u003cem\u003eOs01g0133400\u003c/em\u003e between maize and sorghum, rice, respectively. These results showed a closer evolutionary relationship between the two species containing more collinear gene pairs, and most of the no-collinear genes may be produced in earlier replication events.\u003c/p\u003e \u003cp\u003eGene structure and conserved motif analyses were performed to further explore the evolutionary relationship in the MST family of maize. MST genes could be divided into seven subfamilies, and the PMT, STP, and TMT subfamilies contained fewer exons and simpler gene structures, while the pGlcT, ERD, INT, and VGT subfamilies had more exons and more complex gene structures, and gene structures were conserved in the same subfamily (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Motifs 1\u0026ndash;6 were present in almost all MST family members and were vital for transport function and membrane localization, suggesting that these motifs were highly conserved domains during the evolution of the MST family. Other motifs existed in different subfamilies, indicating that these subfamily members transport different substrates. In addition, STP subfamily members contained all motifs except for motif 11, which implied a broader transport capability for different substrates. The effect of genes on plants depends not only on the function of genes themselves, but also on the regulation of gene expression. The \u003cem\u003ecis\u003c/em\u003e-elements in the promoter region were involved in the regulation of the gene expression, and therefore the \u003cem\u003ecis\u003c/em\u003e-element analysis was important for the preliminary prediction of gene expression. The type and number of \u003cem\u003ecis\u003c/em\u003e-elements in promoters of MST family genes were identified in this study. The ABA response element ABRE belongs to the phytohormone response element and exists in most MST member promoter regions, meaning that the expression of MST members might be involved in the ABA signaling pathway. Additionally, many stress response elements were also found in the promoter region of MST members, such as DRE, MYB, and MYC, suggesting that MST family genes have an important role in maize response to environmental stresses.\u003c/p\u003e \u003cp\u003eTo further investigate the functional response to environmental stresses, the expression mode of MST family genes in the maize seedling stage was detected under different stress treatments. The expression of half of MST genes was significantly induced under PEG treatment, while fewer genes were induced by salt and Cd stresses (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eA, B, and C). Meanwhile, the response of each subfamily in MST to different stresses was also different. Most of the PMT and ERD members responded to osmotic stress, and all ERD members were induced to expression by Cd stresses. The expression of most members was not induced in the STP subfamily under osmotic or ionic stresses. The expression patterns of MST members in response to stress treatments suggested that each subfamily might have different roles in balancing maize growth and development and responding to abiotic stress, implying that the members in each subfamily had also shown functional divergence during evolution and family expansion. ABA is a very important phytohormone in the plant response to abiotic stresses, and many stress response genes are regulated by ABA signaling [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. In this study, the expression levels of some MST family members were induced by both ABA treatment and drought stress, indicating that these members might respond to stress in the ABA signaling pathway, while others were only induced by drought stress, meaning that they might respond to stress in the non-ABA signaling pathway.\u003c/p\u003e \u003cp\u003eTogether, we have identified maize MST family genes. Our findings could contribute to future research on maize MST family genes and provide the foundation for additional investigation of the fundamental functions of this significant monosaccharide transporter family. These findings provide insight into the possible roles of genetic improvement in the capacity of maize to respond to abiotic challenges and may be used to identify relevant candidate MST family genes for functional research.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eIn summary, a total of 66 ZmMST genes were identified from maize and divided into seven subfamilies. Phylogenetic tree, synteny, and collinearity analyses provide preliminary insights into dissecting the evolution and expansion of the MST family. Meanwhile, the expression analysis provides valuable clues for exploring the potential function of MST in balancing maize growth and development and abiotic stress response. These findings will be helpful for us to deeply understand the functions of maize MST genes and provide some important information for functional analysis in the future.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eIdentification and Evolutionary Analysis\u003c/h2\u003e \u003cp\u003eThe complete amino acid and nucleotide sequences of Zea mays B73 RefGen_v4 were downloaded from MaizeGDB (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://maizegdb.org/\u003c/span\u003e\u003cspan address=\"https://maizegdb.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). In addition, MaizeGDB (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://maizegdb.org/\u003c/span\u003e\u003cspan address=\"https://maizegdb.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) transcriptome data were obtained. Seventy-nine different samples comprised the maize inbred line B73 RNA-seq gene map [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. The transcriptome information was chosen from eight distinct maize tissues and developmental stages. From Ensembl (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://asia.ensembl.org/index.html\u003c/span\u003e\u003cspan address=\"https://asia.ensembl.org/index.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), protein sequences for the rice MST and the Arabidopsis MST were acquired. The hidden Markov model repository was built using known MST protein sequences, and HMMER (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://hmmer.org/\u003c/span\u003e\u003cspan address=\"http://hmmer.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to query the maize dataset [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. By utilizing MST genes from rice and Arabidopsis as queries in a BLAST search, MST genes from maize were investigated. Using the PFAM (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://pfam.xfam.org/\u003c/span\u003e\u003cspan address=\"http://pfam.xfam.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and CDD (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ncbi.nlm.nih.gov/cdd/\u003c/span\u003e\u003cspan address=\"https://www.ncbi.nlm.nih.gov/cdd/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) databases, the conserved domains of the discovered ZmMST genes were predicted [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]. Using MEGA 7.0 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.megasoftware.net/\u003c/span\u003e\u003cspan address=\"https://www.megasoftware.net/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and ClustalW software (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.genome.jp/tools-bin/clustalw\u003c/span\u003e\u003cspan address=\"https://www.genome.jp/tools-bin/clustalw\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), evolutionary trees were created for the MST proteins from Arabidopsis, rice, and maize (with 1000 bootstrap replicates) [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]. Using a MapChart (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://mg2c.iask.in/mg2c_v2.0/\u003c/span\u003e\u003cspan address=\"http://mg2c.iask.in/mg2c_v2.0/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and the chromosomal start and termination data received from MaizeGDB (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://maizegdb.org/\u003c/span\u003e\u003cspan address=\"https://maizegdb.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), the chromosomal locations of ZmMST genes were determined [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]. Tandem duplicated genes were found using the tools for multiple covariance scanning (MCScanX, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://chibba.pgml.uga.edu/mcscan2/MCScanX.zip\u003c/span\u003e\u003cspan address=\"http://chibba.pgml.uga.edu/mcscan2/MCScanX.zip\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eSequence Analysis\u003c/h2\u003e \u003cp\u003eThe molecular weight (MW) and isoelectric point (pI) of the ZmMST proteins were predicted using the ExPASy proteomics system (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://web.expasy.org/protparam/\u003c/span\u003e\u003cspan address=\"http://web.expasy.org/protparam/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. The conserved protein motifs of ZmMST genes were discovered using MEME Suite (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://meme-suite.org/\u003c/span\u003e\u003cspan address=\"http://meme-suite.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), and they were further annotated with TBtools (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://github.com/CJ-Chen/TBtools\u003c/span\u003e\u003cspan address=\"https://github.com/CJ-Chen/TBtools\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e]. Fifteen motifs, with lengths ranging from 6 to 50 bp, made up the domain structure (Additional file 14). Using GSDS (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://gsds.gao-lab.org\u003c/span\u003e\u003cspan address=\"http://gsds.gao-lab.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), the gene structure was evaluated [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e]. The PlantCARE database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://bioinformatics.psb.ugent.be/webtools/plantcare/html/\u003c/span\u003e\u003cspan address=\"http://bioinformatics.psb.ugent.be/webtools/plantcare/html/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to predict the 1500 bp sequence upstream of the \u003cem\u003ecis\u003c/em\u003e-acting components of the coding sequences (Additional file 15). Further examination was performed on the components (ABRE, DRE, LTRE, ERE, and MBS) connected to the abiotic stress response [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eReplication Events and Ka/Ks Analysis of MST Genes\u003c/h2\u003e \u003cp\u003eMCScanX (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://chibba.pgml.uga.edu/mcscan2/MCScanX.zip\u003c/span\u003e\u003cspan address=\"http://chibba.pgml.uga.edu/mcscan2/MCScanX.zip\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to analyze correlations between ZmMST genes and single or multiple intergenomic variables as well as associations within genomes [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. Gene family expansion and doubling can occur through several different processes, including whole-genome duplication or polyploidization, tandem duplication, segmental duplication, transposon-mediated transposon duplication, and retro-position. Finally, a graph of the intragenomic duplication events and gene density findings was created using TBtools software (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://github.com/CJ-Chen/TBtools\u003c/span\u003e\u003cspan address=\"https://github.com/CJ-Chen/TBtools\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eClustalW (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.genome.jp/tools-bin/clustalw\u003c/span\u003e\u003cspan address=\"https://www.genome.jp/tools-bin/clustalw\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to determine the ratio of Ka (non-synonymous substitution rate) and Ks (synonymous substitution rate) to investigate the selection pressure on the ZmMST family. The time of occurrence of segmental duplication events for homologous genes was calculated as T\u0026thinsp;=\u0026thinsp;Ks/2λ\u0026thinsp;\u0026times;\u0026thinsp;10\u003csup\u003e\u0026minus;\u0026thinsp;6\u003c/sup\u003e, where λ is the rate of molecular substitution in grasses (6.5\u0026times;10\u003csup\u003e\u0026minus;\u0026thinsp;9\u003c/sup\u003e), and expressed as a million years ago (Mya) [\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003ePlant Treatments and Quantitative Real-Time PCR Analysis\u003c/h2\u003e \u003cp\u003eThe autogamous maize cultivar \u0026ldquo;inbred line\u0026rdquo; was used in the study. Seeds were preserved in our laboratory and incubated in the seedling culture room of the Laboratory of Plant Physiology and Germplasm, Shenyang Agricultural University. The seeds were disinfected with 75% ethanol and washed with distilled water after 1 min to remove the residual ethanol. The cleaned seeds were evenly sown in seedling pots and irrigated with distilled water to allow the vermiculite to absorb sufficient water. Hoagland\u0026rsquo;s nutrient solution (pH 6.0) was added to the basal tray in which the seedling pots were placed to ensure that the roots could access the nutrient solution. The nutrient solution was replaced every 3 days until the seedlings attained the three-leaf stage [\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e]. The three-leaf seedlings were then treated with drought, salt, Cd stresses, and exogenous ABA by application of half-strength Hoagland\u0026rsquo;s nutrient solution supplemented with 20% PEG for drought stress treatment, 200 mol/L NaCl for salt stress treatment, 40 mg/L CdCl\u003csub\u003e2\u003c/sub\u003e for Cd stress treatment, and 100 \u0026micro;mol/L ABA for ABA stress treatment. The uppermost mature leaves were collected at 0, 6, 12, and 24 h after initiation of the stress treatment, with three biological replicates at each time point. The tested leaves were immediately frozen in liquid nitrogen and kept at -80\u0026deg;C.\u003c/p\u003e \u003cp\u003eTotal RNA isolation and quantitative real-time PCR (qRT-PCR) analysis were performed to analyze the expression of maize genes under salt, drought, Cd stress and exogenous ABA treatment [\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e]. A total of 61 maize MST genes were used for the analysis. Total RNA from plant leaves was extracted using TRIzol Reagent (CW Biotech) and subjected to DNase I treatment to remove genomic DNA contamination. The RNA concentration was determined utilizing a BioDrop ultramicro ultraviolet nucleic acid assay. First-strand cDNA was synthesized from 1 \u0026micro;g of total RNA using the UEIris II RT-PCR system. qRT-PCR assays were performed using a real-time PCR analyzer (Bio-Rad, Applied Biosystems PCR, SCILOGEX Gradient Thermal Cycler PCR Instrument TC1000-G). Each reaction mixture contained 10 \u0026micro;L of 2\u0026times;SYBR\u0026reg; Green Pro Taq HS Premix, 1.0 \u0026micro;L cDNA sample, 0.4 \u0026micro;L forward primer (final concentration 10 \u0026micro;M), and 0.4 \u0026micro;L reverse primer (final concentration 10 \u0026micro;M) in a final volume of 20 \u0026micro;L. The thermal-cycling protocol was as follows: 95\u0026deg;C for 5 min, then 45 cycles of 95\u0026deg;C for 15 s and 60\u0026deg;C for 1 min. Melting curve analysis was used to verify the specificity of the reaction. Three technical replicates of each cDNA sample were analyzed. The \u003cem\u003eZm00001d013367\u003c/em\u003e genes were selected as an internal control to normalize the transcript levels of ZmMST genes. The relative gene expression levels were calculated using the 2\u003csup\u003e\u0026minus;∆∆CT\u003c/sup\u003e method (Additional file 16). The normalized data were processed with TBtools and plotted as a heatmap to visualize the changes in MST gene expression (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://github.com/CJ-Chen/TBtools\u003c/span\u003e\u003cspan address=\"https://github.com/CJ-Chen/TBtools\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) [\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e]. Row-scale and log-scale normalization calculations and row clustering were performed on the heatmaps. The number of genes with relative gene expression greater than 2 and relative gene expression less than 0.5 under the four treatments were shown by using the online site Venny 2.1.0 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://bioinfogp.cnb.csic.es/tools/venny/\u003c/span\u003e\u003cspan address=\"https://bioinfogp.cnb.csic.es/tools/venny/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) to create Venn diagrams. All primer pairs were designed with Primer (v5.0) software (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.broadinstitute.org/ftp/pub/software/Primer5.0/\u003c/span\u003e\u003cspan address=\"http://www.broadinstitute.org/ftp/pub/software/Primer5.0/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and were listed in Additional file 17.\u003c/p\u003e \u003c/div\u003e"},{"header":"Abbreviations","content":"\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eFull Name\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eAbbreviated Name\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eMonosaccharide transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eMST\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003emajor facilitator superfamily\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eMFS\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003esucrose transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eSUT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eSugar Transport Protein\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eSTP\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003ePolyol/Monosaccharide Transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003ePMT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eVacuolar Glucose Transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eVGT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eInositol Transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eINT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003ePlastidic Glucose Transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003epGlcT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eTonoplast Membrane Transporter\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eTMT\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eEarly-Responsive to Dehydration six-like\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eERD\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003echromosome\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003echr\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003ewhole-genome duplication\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eWGD\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003emolecular weight\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eMW\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eisoelectric point\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003epI\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003enon-synonymous substitution rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eKa\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003esynonymous substitution rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eKs\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003emillion years ago\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eMya\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003equantitative real-time PCR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eqRT-PCR\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003eAbscisic Acid\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eABA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003esalicylic acid\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eSA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"68.68686868686869%\"\u003e\n \u003cp\u003emethyl jasmonate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.31313131313131%\"\u003e\n \u003cp\u003eMeJA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful to all of the lab members who contributed helpful technical support and data analysis. \u0026quot;The College of Bioscience and Biotechnology Shenyang Agricultural University and Shenyang City Key Laboratory of Maize Genomic Selection Breeding\u0026quot; have our sincere gratitude. We also value the suggestions made for improvements by the reputable editors and reviewers of the text.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026apos; contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConceptualization, JingJuan Fan and Cong Li; methodology, YanShu Zhu; validation, JiaLun Zhu, TianFeng Li, Jing Ma, and WenYu Li; formal analysis, JiaLun Zhu, TianFeng Li, and HanYu Zhang; data curation, JiaLun Zhu; writing-original draft preparation, JiaLun Zhu and TianFeng Li; writing-review and editing, Cong Li; visualization, Tsyganova Nadezhda and XiaoMei Dong; supervision, YanShu Zhu; project administration, JingJuan Fan; funding acquisition, JingJuan Fan. All authors have read and agreed to the published version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003ethe international cooperation program for universities in Liaoning Province Department of Education, China (No. 2023GJ0008), the scientific research projects of Liaoning Provincial Department of Education, China (No. LJKMZ20221010).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\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\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no conflicts of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eFedoroff NV, Battisti DS, Beachy RN, Cooper PJ, Fischhoff DA, Hodges CN, et al. 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Rep. 2012;39(3):2401-2415.\u003c/li\u003e\n\u003cli\u003eGaut BS, Morton BR, McCaig BC, Clegg MT. Substitution rate comparisons between grasses and palms: Synonymous rate differences at the nuclear gene Adh parallel rate differences at the plastid gene rbcL. Proc. Natl. Acad. Sci. USA 1996;93(19):10274-10279.\u003c/li\u003e\n\u003cli\u003eHothem SD, Marley KA, Larson RA. Photochemistry in Hoagland\u0026rsquo; s Nutrient Solution. J. Plant Nutr. 2003;26(4):845-854.\u003c/li\u003e\n\u003cli\u003eNaeem M, Shahzad K, Saqib S, Asim S, Nasrullah MY, Muhammad IA. The Solanum melongena COP1LIKE manipulates fruit ripening and flowering time in tomato (Solanum lycopersicum). Plant Growth Regul. 2022;96(3):369-382.\u003c/li\u003e\n\u003cli\u003eTrick AY, Chen FE, Schares JA, Freml BE, Lor P, Yun Y, et al. High resolution estimates of relative gene abundance with quantitative ratiometric regression PCR (qRR-PCR). Analyst 2021;146(21):6463-6469.\u003c/li\u003e\n\u003c/ol\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-plant-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pbio","sideBox":"Learn more about [BMC Plant Biology](http://bmcplantbiol.biomedcentral.com/)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/pbio/default.aspx","title":"BMC Plant Biology","twitterHandle":"BMC_series","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"em","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Genome-wide identification, Zea mays L., Monosaccharide transporter (MST), Gene family, Abiotic stress","lastPublishedDoi":"10.21203/rs.3.rs-3219446/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3219446/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eBackground\u003c/h2\u003e \u003cp\u003eMonosaccharide transporter (MST) family, as a carrier for monosaccharide transport, plays an important role in carbon partitioning and widely involves in plant growth and development, stress response, and signaling transduction. However, little information on the MST family genes is reported in maize (\u003cem\u003eZea mays\u003c/em\u003e), especially in response to abiotic stresses. In this study, the genome-wide identification of MST family genes was performed in maize.\u003c/p\u003e\u003ch2\u003eResult\u003c/h2\u003e \u003cp\u003eA total of sixty-six putative members of MST gene family were identified and divided into seven subfamilies (including SPT, PMT, VGT, INT, pGlcT, TMT, and ERD) using bioinformatics approaches, and gene information, phylogenetic tree, chromosomal location, gene structure, motif composition, and \u003cem\u003ecis\u003c/em\u003e-acting elements were investigated. Eight tandem and twelve segmental duplication events were identified, which played an important role in the expansion of the ZmMST family. Synteny analysis revealed the evolutionary features of MST genes in three gramineous crop species. The expression analysis indicated that most of the PMT, VGT, and ERD subfamilies members responded to osmotic and cadmium stresses, and some of them were regulated by ABA signaling, while only a few members of other subfamilies responded to stresses. In addition, only five genes were induced by NaCl stress in MST family.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e \u003cp\u003eThese results serve to understand the evolutionary relationships of the ZmMST family genes and supply some insight into the processes of monosaccharide transport and carbon partitioning on the balance between plant growth and development and stress response in maize.\u003c/p\u003e","manuscriptTitle":"Genome-Wide Identification and Investigation of MST Gene Family Based on Their Evolution and Expression Analysis under Abiotic Stress and Hormone Treatments in Maize (Zea mays L.)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-08-09 14:14:13","doi":"10.21203/rs.3.rs-3219446/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-03-17T07:16:15+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-03-16T05:06:31+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-03-12T03:15:08+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"d68394a2-bead-41e1-a634-ef49bc3a5f6e","date":"2024-03-06T02:34:50+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2023-09-14T17:52:28+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"8ebda5a7-a22e-4cd5-9f01-790e762d5614","date":"2023-09-01T14:21:25+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"32cb6731-1d7e-4aab-a039-4e79510e4647","date":"2023-08-29T07:31:19+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2023-08-26T06:29:42+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2023-08-18T11:32:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2023-08-13T16:21:08+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2023-08-04T09:11:44+00:00","index":"","fulltext":""},{"type":"submitted","content":"BMC Plant Biology","date":"2023-07-31T06:29:05+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"bmc-plant-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pbio","sideBox":"Learn more about [BMC Plant Biology](http://bmcplantbiol.biomedcentral.com/)","snPcode":"","submissionUrl":"https://www.editorialmanager.com/pbio/default.aspx","title":"BMC Plant Biology","twitterHandle":"BMC_series","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"em","reportingPortfolio":"BMC Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"7c919e3d-5286-4af9-b8c8-639a71081f56","owner":[],"postedDate":"August 9th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2024-05-22T18:17:32+00:00","versionOfRecord":[],"versionCreatedAt":"2023-08-09 14:14:13","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3219446","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3219446","identity":"rs-3219446","version":["v1"]},"buildId":"WrCJVZZCHTDjtuVLN7oU0","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}