{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T11:48:19Z","timestamp":1788349699784,"version":"build-2803163510"},"reference-count":40,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2022,8,22]],"date-time":"2022-08-22T00:00:00Z","timestamp":1661126400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Natural Science Foundation of China","award":["No. 31971770"],"award-info":[{"award-number":["No. 31971770"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Hydroxycinnamoyl-CoA shikimate\/quinate hydroxycinnamoyl transferase (HCT) is considered to be an essential enzyme for regulating the biosynthesis and composition of lignin. To investigate the properties and function of ZjHCT4, the ZjHCT4 gene was cloned from Zoysia japonica with a completed coding sequence of 1284-bp in length, encoding 428 amino acids. The ZjHCT4 gene promoter has several methyl jasmonate (MeJA) response elements. According to analysis of expression patterns, it was up-regulated by MeJA, GA3 (Gibberellin), and SA (Salicylic acid), and down-regulated by ABA (Abscisic acid). Ectopic ZjHCT4 expression in creeping bentgrass causes excessive plant elongation. In addition, the content of G-lingnin and H-lingnin fell in transgenic plants, whereas the level of S-lingnin increased, resulting in a considerable rise in the S\/G unit ratio. Analysis of the expression levels of lignin-related genes revealed that the ectopic expression of ZjHCT4 altered the expression levels of a number of genes involved in the lignin synthesis pathway. Simultaneously, MeJA, SA, GA3, IAA, BR (Brassinosteroid), and other hormones were dramatically enhanced in transgenic plants relative to control plants, whereas ABA concentration was significantly decreased. Expression of ZjHCT4 impacted lignin composition and plant growth via altering the phenylpropionic acid metabolic pathway and hormone response, as revealed by transcriptome analysis. HCTs may influence plant lignin composition and plant development by altering hormone content. These findings contributed to a deeper comprehension of the lignin synthesis pathway and set the stage for further investigation and application of the HCTs gene.<\/jats:p>","DOI":"10.3390\/ijms23169500","type":"journal-article","created":{"date-parts":[[2022,8,22]],"date-time":"2022-08-22T21:30:34Z","timestamp":1661203834000},"page":"9500","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":27,"title":["Expression of a Hydroxycinnamoyl-CoA Shikimate\/Quinate Hydroxycinnamoyl Transferase 4 Gene from Zoysia japonica (ZjHCT4) Causes Excessive Elongation and Lignin Composition Changes in Agrostis stolonifera"],"prefix":"10.3390","volume":"23","author":[{"given":"Di","family":"Dong","sequence":"first","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhuoxiong","family":"Yang","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuan","family":"Ma","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuwen","family":"Li","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mengdi","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yinruizhi","family":"Li","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5376-4878","authenticated-orcid":false,"given":"Zhuocheng","family":"Liu","sequence":"additional","affiliation":[{"name":"School of Biological and Food Engineering, Suzhou University, Suzhou 234000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenyan","family":"Jia","sequence":"additional","affiliation":[{"name":"Inner Mongolia M-Grass Ecology and Environment (Group) Co., Ltd., Hohhot 010000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liebao","family":"Han","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0591-1092","authenticated-orcid":false,"given":"Yuehui","family":"Chao","sequence":"additional","affiliation":[{"name":"School of Grassland Science, Beijing Forestry University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,8,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1104\/pp.113.217836","article-title":"Elucidation of the Structure and Reaction Mechanism of Sorghum Hydroxycinnamoyltransferase and Its Structural Relationship to Other Coenzyme A-Dependent Transferases and Synthases","volume":"162","author":"Walker","year":"2013","journal-title":"Plant Physiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1104\/pp.110.155119","article-title":"Lignin Biosynthesis and Structure","volume":"153","author":"Vanholme","year":"2010","journal-title":"Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"190215","DOI":"10.1098\/rsob.190215","article-title":"Lignin Biosynthesis: Old Roads Revisited and New Roads Explored","volume":"9","author":"Dixon","year":"2019","journal-title":"Open Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1146\/annurev.arplant.54.031902.134938","article-title":"Lignin Biosynthesis","volume":"54","author":"Boerjan","year":"2003","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1111\/j.1469-8137.2010.03327.x","article-title":"The Origin and Evolution of Lignin Biosynthesis","volume":"187","author":"Weng","year":"2010","journal-title":"New Phytol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1093\/plcell\/koab044","article-title":"Function of the HYDROXYCINNAMOYL-CoA:SHIKIMATE HYDROXYCINNAMOYL TRANSFERASE Is Evolutionarily Conserved in Embryophytes","volume":"33","author":"Kriegshauser","year":"2021","journal-title":"Plant Cell"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1074\/jbc.M209362200","article-title":"Purification, Cloning, and Properties of an Acyltransferase Controlling Shikimate and Quinate Ester Intermediates in Phenylpropanoid Metabolism","volume":"278","author":"Hoffmann","year":"2003","journal-title":"J. Biol. Chem."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.1016\/j.phytochem.2006.06.028","article-title":"Biosynthesis of Curcuminoids and Gingerols in Turmeric (Curcuma Longa) and Ginger (Zingiber Officinale): Identification of Curcuminoid Synthase and Hydroxycinnamoyl-CoA Thioesterases","volume":"67","author":"Timmermann","year":"2006","journal-title":"Phytochemistry"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1521","DOI":"10.1016\/j.phytochem.2007.03.022","article-title":"Down-Regulation of Hydroxycinnamoyl CoA: Shikimate Hydroxycinnamoyl Transferase in Transgenic Alfalfa Affects Lignification, Development and Forage Quality","volume":"68","author":"Shadle","year":"2007","journal-title":"Phytochemistry"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1111\/j.1469-8137.2010.03621.x","article-title":"Selective Lignin Downregulation Leads to Constitutive Defense Response Expression in Alfalfa (Medicago Sativa L.)","volume":"190","author":"Jikumaru","year":"2011","journal-title":"New Phytol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1186\/s13068-021-01905-1","article-title":"Targeting Hydroxycinnamoyl CoA: Shikimate Hydroxycinnamoyl Transferase for Lignin Modification in Brachypodium Distachyon","volume":"14","author":"Barros","year":"2021","journal-title":"Biotechnol. Biofuels"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"11856","DOI":"10.1073\/pnas.0701428104","article-title":"Exploring Lignification in Conifers by Silencing Hydroxycinnamoyl-CoA:Shikimate Hydroxycinnamoyltransferase in Pinus Radiata","volume":"104","author":"Wagner","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1007\/s11103-013-0152-y","article-title":"Early lignin pathway enzymes and routes to chlorogenic acid in switchgrass (Panicum virgatumL.)","volume":"84","author":"Shen","year":"2014","journal-title":"Plant Mol. Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1446","DOI":"10.1105\/tpc.020297","article-title":"Silencing of hydroxycinnamoyl-coenzyme A shikimate\/quinate hydroxycinnamoyltransferase affects phenylpropanoid biosynthesis","volume":"16","author":"Hoffmann","year":"2004","journal-title":"Plant Cell"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/j.ymben.2009.11.003","article-title":"Production of novel antioxidative phenolic amides through heterologous expression of the plant\u2019s chlorogenic acid biosynthesis genes in yeast","volume":"12","author":"Andrea","year":"2010","journal-title":"Metab. Eng."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.1007\/s00425-011-1367-2","article-title":"Distinct substrate specificities and unusual substrate flexibilities of two hydroxycinnamoyltransferases, rosmarinic acid synthase and hydroxycinnamoyl-CoA: Shikimate hydroxycinnamoyl-transferase, from Coleus blumei Benth","volume":"233","author":"Sander","year":"2011","journal-title":"Planta"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.phytochem.2011.12.015","article-title":"Characterization of hydroxycinnamoyltransferase from rice and its application for biological synthesis of hydroxycinnamoyl glycerols","volume":"76","author":"Kim","year":"2012","journal-title":"Phytochemistry"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1111\/tpj.15354","article-title":"Ectopic expression of the Arabidopsis florigen gene FLOWERING LOCUS T in seeds enhances seed dormancy via the GA and DOG1 pathways","volume":"107","author":"Chen","year":"2021","journal-title":"Plant J."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1007\/s11103-004-2648-y","article-title":"Ectopic expression of OsYAB1 causes extra stamens and carpels in rice","volume":"56","author":"Jang","year":"2004","journal-title":"Plant Mol. Biol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1007\/s13562-019-00533-z","article-title":"Ectopic expression of an antisense BpCCoAOMT gene from Betula platyphylla Suk. affects growth and development of tobacco due to lignin content reductio","volume":"29","author":"Zhang","year":"2019","journal-title":"J. Plant Biochem. Biotechnol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1744-697X.2009.00129.x","article-title":"Effect of Storage Temperature and Duration on the Vigor of Zoysiagrass (Zoysia Japonica Steud.) Seed Harvested at Different Maturity Stages","volume":"55","author":"Mao","year":"2009","journal-title":"Grassl. Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"15181","DOI":"10.1038\/s41598-018-33597-3","article-title":"Comparative Transcriptomic Analysis Reveals Common Molecular Factors Responsive to Heat and Drought Stress in Agrostis Stolonifera","volume":"8","author":"Xu","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"88","DOI":"10.5660\/WTS.2016.5.2.88","article-title":"Performance of Zoysia Spp. and Axonopus Compressus Turf on Turf-Paver Complex under Simulated Traffic","volume":"5","author":"Chin","year":"2016","journal-title":"Weed Turfgrass Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1111\/j.1365-313X.2006.02946.x","article-title":"Characterization of a BAHD Acyltransferase Responsible for Producing the Green Leaf Volatile (Z)-3-hexen-1-yl Acetate in Arabidopsis Thaliana","volume":"49","author":"Pichersky","year":"2007","journal-title":"Plant J."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Sun, C.-H., Yang, C.-Y., and Tzen, J.T.C. (2018). Molecular Identification and Characterization of Hydroxycinnamoyl Transferase in Tea Plants (Camellia sinensis L.). Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19123938"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1093\/pcp\/pcn077","article-title":"Jasmonate-Induced Nicotine Formation in Tobacco Is Mediated by Tobacco COI1 and JAZ Genes","volume":"49","author":"Shoji","year":"2008","journal-title":"Plant Cell Physiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1111\/pbi.12439","article-title":"Combining enhanced biomass density with reduced lignin level for improved forage quality","volume":"14","author":"Shadle","year":"2016","journal-title":"Plant Biotechnol. J."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/S0031-9422(02)00211-X","article-title":"Trends in lignin modification: A comprehensive analysis of the effects of genetic manipulations\/mutations on lignification and vascular integrity","volume":"61","author":"Anterola","year":"2002","journal-title":"Phytochemistry"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1146\/annurev-genet-102209-163508","article-title":"The genetics of lignin biosynthesis: Connecting genotype to phenotype","volume":"44","author":"Bonawitz","year":"2010","journal-title":"Ann. Rev. Genet."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1105\/tpc.106.044495","article-title":"Flavonoid accumulation in Arabidopsis repressed in lignin synthesis affects auxin transport and plant growth","volume":"19","author":"Besseau","year":"2007","journal-title":"Plant Cell"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1104\/pp.19.00533","article-title":"Ectopic defense gene expression is associated with growth defects in medicago truncatula lignin pathway mutants","volume":"181","author":"Chan","year":"2019","journal-title":"Plant Physiol."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Dong, D., Zhao, Y., Teng, K., Tan, P., Liu, Z., Yang, Z., Han, L., and Chao, Y. (2022). Expression of ZjPSY, a Phytoene Synthase Gene from Zoysia Japonica Affects Plant Height and Photosynthetic Pigment Contents. Plants, 11.","DOI":"10.3390\/plants11030395"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1046\/j.1365-313x.1998.00343.x","article-title":"Floral Dip: A Simplified Method for Agrobacterium-Mediated Transformation of Arabidopsis Thaliana","volume":"16","author":"Clough","year":"2010","journal-title":"Plant J."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1007\/s00299-003-0734-2","article-title":"Agrobacterium Tumefaciens-Mediated Creeping Bentgrass (Agrostis Stolonifera L.) Transformation Using Phosphinothricin Selection Results in a High Frequency of Single-Copy Transgene Integration","volume":"22","author":"Luo","year":"2004","journal-title":"Plant Cell Rep."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1093\/nar\/30.1.325","article-title":"PlantCARE, a Database of Plant Cis-Acting Regulatory Elements and a Portal to Tools for in Silico Analysis of Promoter Sequences","volume":"30","author":"Lescot","year":"2002","journal-title":"Nucleic Acids Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1093\/molbev\/msy096","article-title":"MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms","volume":"35","author":"Kumar","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.envexpbot.2017.10.010","article-title":"Interactive Effects of Melatonin and Cytokinin on Alleviating Drought-Induced Leaf Senescence in Creeping Bentgrass (Agrostis Stolonifera)","volume":"145","author":"Ma","year":"2018","journal-title":"Environ. Exp. Bot."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1038\/nprot.2008.73","article-title":"Analyzing Real-Time PCR Data by the Comparative CT Method","volume":"3","author":"Schmittgen","year":"2008","journal-title":"Nat. Protoc."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.aca.2006.04.060","article-title":"Comparison between Conventional Indirect Competitive Enzyme-Linked Immunosorbent Assay (IcELISA) and Simplified IcELISA for Small Molecules","volume":"571","author":"Zhao","year":"2006","journal-title":"Anal. Chim. Acta"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1105\/tpc.13.1.73","article-title":"Downregulation of Caffeic Acid 3-O-Methyltransferase and Caffeoyl CoA 3-O-Methyltransferase in Transgenic Alfalfa: Impacts on Lignin Structure and Implications for the Biosynthesis of G and S Lignin","volume":"13","author":"Guo","year":"2001","journal-title":"Plant Cell"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/16\/9500\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T00:13:42Z","timestamp":1760141622000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/23\/16\/9500"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,8,22]]},"references-count":40,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2022,8]]}},"alternative-id":["ijms23169500"],"URL":"https:\/\/doi.org\/10.3390\/ijms23169500","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,8,22]]}}}