{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,26]],"date-time":"2026-04-26T13:16:32Z","timestamp":1777209392609,"version":"3.51.4"},"reference-count":69,"publisher":"MDPI AG","issue":"23","license":[{"start":{"date-parts":[[2024,12,9]],"date-time":"2024-12-09T00:00:00Z","timestamp":1733702400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32170378"],"award-info":[{"award-number":["32170378"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Danshen (Salvia miltiorrhiza Bunge) is a perennial herbaceous plant of the Salvia genus in the family Lamiaceae. Its dry root is one of the important traditional Chinese herbal medicines with a long officinal history. The yield and quality of S. miltiorrhiza are influenced by various factors, among which drought is one of the most significant types of abiotic stress. Based on the transcriptome database of S. miltiorrhiza, our research group discovered a carotenoid cleavage dioxygenase gene, SmCCD4, belonging to the carotenoid cleavage oxygenase (CCO) gene family which is highly responsive to drought stress on the basis of our preceding work. Here, we identified 26 CCO genes according to the whole-genome database of S. miltiorrhiza. The expression pattern of SmCCD4 showed that this gene is strongly overexpressed in the aboveground tissue of S. miltiorrhiza. And by constructing SmCCD4 overexpression strains, it was shown that the overexpression of SmCCD4 not only promotes the synthesis of abscisic acid and increases plant antioxidant activity but also regulates the synthesis of the secondary metabolites tanshinone and phenolic acids in S. miltiorrhiza. In summary, this study is the first in-depth and systematic identification and investigation of the CCO gene family in S. miltiorrhiza. The results provide useful information for further systematic research on the function of CCO genes and provide a theoretical basis for improving the yield and quality of S. miltiorrhiza.<\/jats:p>","DOI":"10.3390\/ijms252313223","type":"journal-article","created":{"date-parts":[[2024,12,9]],"date-time":"2024-12-09T10:11:47Z","timestamp":1733739107000},"page":"13223","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Carotenoid Cleavage Dioxygenase Gene CCD4 Enhances Tanshinone Accumulation and Drought Resistance in Salvia miltiorrhiza"],"prefix":"10.3390","volume":"25","author":[{"given":"Qian","family":"Tian","sequence":"first","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wei","family":"Han","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shuai","family":"Zhou","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Liu","family":"Yang","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3437-7386","authenticated-orcid":false,"given":"Donghao","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wen","family":"Zhou","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6111-1551","authenticated-orcid":false,"given":"Zhezhi","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of the Ministry of Education for Medicinal Resources and Natural Pharmaceutical Chemistry, National Engineering Laboratory for Resource Development of Endangered Crude Drugs in Northwest of China, Shaanxi Normal University, Xi\u2019an 710062, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2024,12,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1016\/j.ijcard.2005.10.026","article-title":"Danshen: A versatile Chinese herbal drug for the treatment of coronary heart disease","volume":"113","author":"Cheng","year":"2006","journal-title":"Int. J. Cardiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Wang, Z., and Peters, R.J. (2022). Tanshinones: Leading the way into Lamiaceae labdane-related diterpenoid biosynthesis. Curr. Opin. Plant Biol., 66.","DOI":"10.1016\/j.pbi.2022.102189"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1016\/j.fct.2018.09.063","article-title":"Tanshinone IIA attenuates TNF-\u03b1 induced PTX3 expression and monocyte adhesion to endothelial cells through the p38\/NF-\u03baB pathway","volume":"121","author":"Fang","year":"2018","journal-title":"Food Chem. Toxicol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1016\/j.jep.2013.11.025","article-title":"Renoprotection of Danshen Injection on streptozotocin-induced diabetic rats, associated with tubular function and structure","volume":"151","author":"Yin","year":"2014","journal-title":"J. Ethnopharmacol."},{"key":"ref_5","first-page":"2721413","article-title":"A Review of Danshen Combined with Clopidogrel in the Treatment of Coronary Heart Disease","volume":"2019","author":"Zhang","year":"2019","journal-title":"Evid.-Based Complement. Altern. Med."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1111\/jcpt.13283","article-title":"Prospective study of the effect of sulfotanshinone sodium combined with tirofiban on vascular endothelial function and indicators of plaque stability in elderly patients with acute coronary syndrome","volume":"46","author":"Lin","year":"2021","journal-title":"J. Clin. Pharm. Ther."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"160726","DOI":"10.1155\/2012\/160726","article-title":"Salvianolic Acid B Attenuates Rat Hepatic Fibrosis via Downregulating Angiotensin II Signaling","volume":"2012","author":"Li","year":"2012","journal-title":"Evid.-Based Complement. Altern. Med."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1941","DOI":"10.1016\/j.chroma.2008.12.032","article-title":"Advancement in analysis of Salviae miltiorrhizae Radix et Rhizoma (Danshen)","volume":"1216","author":"Li","year":"2009","journal-title":"J. Chromatogr. A"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1002\/med.20077","article-title":"New developments in the chemistry and biology of the bioactive constituents of Tanshen","volume":"27","author":"Wang","year":"2007","journal-title":"Med. Res. Rev."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/S1360-1385(03)00053-0","article-title":"Carotenoid oxygenases: Cleave it or leave it","volume":"8","author":"Giuliano","year":"2003","journal-title":"Trends Plant Sci."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Yu, J., Wang, Y., Bai, H., Zhang, X., and Wang, R. (2024). Genome-wide identification and expressional analysis of carotenoid cleavage oxygenase (CCO) gene family in Betula platyphylla under abiotic stress. BMC Genom., 25.","DOI":"10.1186\/s12864-024-10777-2"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1007\/s00775-018-1586-0","article-title":"Preparation and characterization of metal-substituted carotenoid cleavage oxygenases","volume":"23","author":"Sui","year":"2018","journal-title":"J. Biol. Inorg. Chem."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"982","DOI":"10.1111\/j.1365-313X.2006.02666.x","article-title":"Characterization of three members of the Arabidopsis carotenoid cleavage dioxygenase family demonstrates the divergent roles of this multifunctional enzyme family","volume":"45","author":"Auldridge","year":"2006","journal-title":"Plant J."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"2514","DOI":"10.1271\/bbb.130626","article-title":"Mutations in the CCD4 Carotenoid Cleavage Dioxygenase Gene of Yellow-Flesh Peaches","volume":"77","author":"Fukamatsu","year":"2013","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e29376","DOI":"10.4161\/psb.29376","article-title":"Gene silencing of CCD7 and CCD8 in Phelipanche aegyptiaca by tobacco rattle virus system retarded the parasite development on the host","volume":"9","author":"Aly","year":"2014","journal-title":"Plant Signal. Behav."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1111\/j.1365-313X.2005.02622.x","article-title":"Functional analysis of Arabidopsis NCED6 and NCED9 genes indicates that ABA synthesized in the endosperm is involved in the induction of seed dormancy","volume":"45","author":"Lefebvre","year":"2006","journal-title":"Plant J."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2189","DOI":"10.1093\/jxb\/erx476","article-title":"From carotenoids to strigolactones","volume":"69","author":"Jia","year":"2018","journal-title":"J. Exp. Bot."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1872","DOI":"10.1126\/science.276.5320.1872","article-title":"Specific oxidative cleavage of carotenoids by VP14 of maize","volume":"276","author":"Schwartz","year":"1997","journal-title":"Science"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1046\/j.1365-313X.2003.01786.x","article-title":"Molecular characterization of the Arabidopsis 9-cis epoxycarotenoid dioxygenase gene family","volume":"35","author":"Tan","year":"2003","journal-title":"Plant J."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"4737","DOI":"10.1007\/s11033-013-2570-y","article-title":"Genome-wide identification and transcription analysis of soybean carotenoid oxygenase genes during abiotic stress treatments","volume":"40","author":"Wang","year":"2013","journal-title":"Mol. Biol. Rep."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1007\/s11105-015-0943-1","article-title":"A Comprehensive Analysis of Carotenoid Cleavage Dioxygenases Genes in Solanum Lycopersicum","volume":"34","author":"Wei","year":"2016","journal-title":"Plant Mol. Biol. Report."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1111\/j.1742-4658.2008.06820.x","article-title":"Characterization of the rice carotenoid cleavage dioxygenase 1 reveals a novel route for geranial biosynthesis","volume":"276","author":"Ilg","year":"2009","journal-title":"FEBS J."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1007\/s00425-008-0781-6","article-title":"Cloning and characterisation of a maize carotenoid cleavage dioxygenase (ZmCCD1) and its involvement in the biosynthesis of apocarotenoids with various roles in mutualistic and parasitic interactions","volume":"228","author":"Sun","year":"2008","journal-title":"Planta"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"882","DOI":"10.1111\/j.1365-313X.2004.02263.x","article-title":"The tomato carotenoid cleavage dioxygenase 1 genes contribute to the formation of the flavor volatiles \u03b2-ionone, pseudoionone, and geranylacetone","volume":"40","author":"Simkin","year":"2004","journal-title":"Plant J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1104\/pp.106.087130","article-title":"Carotenoid cleavage dioxygenase (CmCCD4a) contributes to white color formation in chrysanthemum petals","volume":"142","author":"Ohmiya","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1007\/s10681-011-0602-z","article-title":"The carotenoid cleavage dioxygenase 4 (CmCCD4a) gene family encodes a key regulator of petal color mutation in chrysanthemum","volume":"184","author":"Yoshioka","year":"2012","journal-title":"Euphytica"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1111\/j.1469-8137.2010.03394.x","article-title":"Modified CAROTENOID CLEAVAGE DIOXYGENASE8 expression correlates with altered branching in kiwifruit (Actinidia chinensis)","volume":"188","author":"Ledger","year":"2010","journal-title":"New Phytol."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Gao, J., Zhang, T., Xu, B., Jia, L., Xiao, B., Liu, H., Liu, L., Yan, H., and Xia, Q. (2018). CRISPR\/Cas9-Mediated Mutagenesis of Carotenoid Cleavage Dioxygenase 8 (CCD8) in Tobacco Affects Shoot and Root Architecture. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19041062"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"46940","DOI":"10.1074\/jbc.M409004200","article-title":"The biochemical characterization of two carotenoid cleavage enzymes from Arabidopsis indicates that a carotenoid-derived compound inhibits lateral branching","volume":"279","author":"Schwartz","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1007\/s00344-015-9520-y","article-title":"Genome-Wide Classification and Abiotic Stress-Responsive Expression Profiling of Carotenoid Oxygenase Genes in Brassica rapa and Brassica oleracea","volume":"35","author":"Kim","year":"2016","journal-title":"J. Plant Growth Regul."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1007\/s11103-014-0250-5","article-title":"New target carotenoids for CCD4 enzymes are revealed with the characterization of a novel stress-induced carotenoid cleavage dioxygenase gene from Crocus sativus","volume":"86","author":"Ahrazem","year":"2014","journal-title":"Plant Mol. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/S1360-1385(01)02187-2","article-title":"Complex regulation of ABA biosynthesis in plants","volume":"7","author":"Seo","year":"2002","journal-title":"Trends Plant Sci."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"3011","DOI":"10.1093\/jxb\/erp137","article-title":"Cloning and functional characterization of carotenoid cleavage dioxygenase 4 genes","volume":"60","author":"Huang","year":"2009","journal-title":"J. Exp. Bot."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1046\/j.1365-313x.2000.00789.x","article-title":"Ectopic expression of a tomato 9-cis-epoxycarotenoid dioxygenase gene causes over-production of abscisic acid","volume":"23","author":"Thompson","year":"2000","journal-title":"Plant J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1046\/j.1365-313X.2003.01708.x","article-title":"Interaction between two cis-acting elements, ABRE and DRE, in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydration and high-salinity stresses","volume":"34","author":"Narusaka","year":"2003","journal-title":"Plant J."},{"key":"ref_36","first-page":"277","article-title":"Cloning genes of Arabidopsis thaliana by chromosome walking","volume":"82","author":"Leung","year":"1998","journal-title":"Methods Mol. Biol."},{"key":"ref_37","first-page":"251","article-title":"A novel cis-acting element in an Arabidopsis gene is involved in responsiveness to drought, low-temperature, or high-salt stress","volume":"6","author":"Shinozaki","year":"1994","journal-title":"Plant Cell"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"9781715","DOI":"10.1155\/2016\/9781715","article-title":"Danshen (Salvia miltiorrhiza) Compounds Improve the Biochemical Indices of the Patients with Coronary Heart Disease","volume":"2016","author":"Liu","year":"2016","journal-title":"Evid.-Based Complement. Altern. Med."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Ma, X., Zhang, L., Gao, F., Jia, W., and Li, C. (2023). Salvia miltiorrhiza and Tanshinone IIA reduce endothelial inflammation and atherosclerotic plaque formation through inhibiting COX-2. Biomed. Pharmacother., 167.","DOI":"10.1016\/j.biopha.2023.115501"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"155521","DOI":"10.1016\/j.phymed.2024.155521","article-title":"Salvia miltiorrhiza Bunge (Danshen) based nano-delivery systems for anticancer therapeutics","volume":"128","author":"Jia","year":"2024","journal-title":"Phytomedicine"},{"key":"ref_41","first-page":"4832","article-title":"Variation in microbial community structure in the rhizosphere soil of Salvia miltiorrhiza Bunge under three cropping modes","volume":"39","author":"Wang","year":"2019","journal-title":"Acta Ecol. Sin."},{"key":"ref_42","first-page":"100032","article-title":"A review on drought stress in plants: Implications, mitigation and the role of plant growth promoting rhizobacteria","volume":"5","author":"Ahluwalia","year":"2021","journal-title":"Resour. Environ. Sustain."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.pbi.2006.03.005","article-title":"Plant carotenoid cleavage oxygenases and their apocarotenoid products","volume":"9","author":"Auldridge","year":"2006","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"24816","DOI":"10.1074\/jbc.M804000200","article-title":"Cytosolic and plastoglobule-targeted carotenoid dioxygenases from Crocus sativus are both involved in \u03b2-ionone release","volume":"283","author":"Rubio","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_45","first-page":"1099","article-title":"Salvia miltiorrhiza as medicinal model plant","volume":"48","author":"Song","year":"2013","journal-title":"Yao Xue Xue Bao = Acta Pharm. Sin."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1016\/j.indcrop.2010.09.006","article-title":"Effect of drought stress on growth and accumulation of active constituents in Salvia miltiorrhiza Bunge","volume":"33","author":"Liu","year":"2011","journal-title":"Ind. Crops Prod."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"124521","DOI":"10.1016\/j.envpol.2024.124521","article-title":"Physiological and molecular mechanisms of carbon quantum dots alleviating Cu2+ toxicity in Salvia miltiorrhiza bunge","volume":"358","author":"Zhong","year":"2024","journal-title":"Environ. Pollut."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"153862","DOI":"10.1016\/j.jplph.2022.153862","article-title":"Overexpression of SmMYC2 enhances salt resistance in Arabidopsis thaliana and Salvia miltiorrhiza hairy roots","volume":"280","author":"Deng","year":"2023","journal-title":"J. Plant Physiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.plaphy.2017.12.001","article-title":"Genome-wide analysis of carotenoid cleavage oxygenase genes and their responses to various phytohormones and abiotic stresses in apple (Malus domestica)","volume":"123","author":"Chen","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Haider, M.Z., Sami, A., Shafiq, M., Anwar, W., Ali, S., Ali, Q., Muhammad, S., Manzoor, I., Shahid, M.A., and Ali, D. (2023). Genome-wide identification and in-silico expression analysis of carotenoid cleavage oxygenases gene family in Oryza sativa (rice) in response to abiotic stress. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1269995"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Yao, Y., Jia, L., Cheng, Y., Ruan, M., Ye, Q., Wang, R., Yao, Z., Zhou, G., Liu, J., and Yu, J. (2022). Evolutionary Origin of the Carotenoid Cleavage Oxygenase Family in Plants and Expression of Pepper Genes in Response to Abiotic Stresses. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.792832"},{"key":"ref_52","first-page":"455","article-title":"Cloning and expression analysis of GmNCED1 from Glycine max","volume":"36","author":"Li","year":"2014","journal-title":"Chin. J. Oil Crop Sci."},{"key":"ref_53","first-page":"611","article-title":"Regulation of drought tolerance by gene manipulation of 9-cis-epoxycarotenoid dioxygenase, a key enzyme in abscisic acid biosynthesis in Arabidopsis (vol 27, pg 325, 2001)","volume":"30","author":"Iuchi","year":"2002","journal-title":"Plant J."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2089","DOI":"10.1126\/science.1085162","article-title":"Biosynthesis of the food and cosmetic plant pigment bixin (annatto)","volume":"300","author":"Bouvier","year":"2003","journal-title":"Science"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1105\/tpc.006536","article-title":"Oxidative remodeling of chromoplast carotenoids: Identification of the carotenoid dioxygenase CsCCD and CsZCD genes involved in crocus secondary metabolite biogenesis","volume":"15","author":"Bouvier","year":"2003","journal-title":"Plant Cell"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1104\/pp.124.1.343","article-title":"Characterization of the 9-cis-epoxycarotenoid dioxygenase gene family and the regulation of abscisic acid biosynthesis in avocado","volume":"124","author":"Chernys","year":"2000","journal-title":"Plant Physiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/j.tplants.2010.04.006","article-title":"ABA perception and signalling","volume":"15","author":"Raghavendra","year":"2010","journal-title":"Trends Plant Sci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1007\/s11105-013-0628-6","article-title":"Identifying a Carotenoid Cleavage Dioxygenase (ccd4) Gene Controlling Yellow\/White Fruit Flesh Color of Peach","volume":"31","author":"Adami","year":"2013","journal-title":"Plant Mol. Biol. Report."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"117529","DOI":"10.1016\/j.indcrop.2023.117529","article-title":"Carotenoid cleavage dioxygenases 4 from woodiness plant and their relationships with herbaceous plants","volume":"205","author":"Jonathan","year":"2023","journal-title":"Ind. Crops Prod."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1007\/s11105-013-0650-8","article-title":"Inactivation of a Gene Encoding Carotenoid Cleavage Dioxygenase (CCD4) Leads to Carotenoid-Based Yellow Coloration of Fruit Flesh and Leaf Midvein in Peach","volume":"32","author":"Ma","year":"2014","journal-title":"Plant Mol. Biol. Report."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"535","DOI":"10.2503\/hortj.UTD-080","article-title":"Comparison of Expression CCD4 Gene Levels in Petals of Evergreen Azalea Species","volume":"88","author":"Ureshino","year":"2019","journal-title":"Hortic. J."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"19401","DOI":"10.1074\/jbc.M116.744912","article-title":"Key Residues for Catalytic Function and Metal Coordination in a Carotenoid Cleavage Dioxygenase","volume":"291","author":"Sui","year":"2016","journal-title":"J. Biol. Chem."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1229","DOI":"10.1093\/molbev\/mst012","article-title":"Building Phylogenetic Trees from Molecular Data with MEGA","volume":"30","author":"Hall","year":"2013","journal-title":"Mol. Biol. Evol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1007\/s11240-006-9187-y","article-title":"Genetic transformation of the medicinal plant Salvia miltiorrhiza by Agrobacterium tumefaciens-mediated method","volume":"88","author":"Yan","year":"2007","journal-title":"Plant Cell Tissue Organ Cult."},{"key":"ref_65","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_66","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1007\/s11033-009-9703-3","article-title":"Characterization of reference genes for quantitative real-time PCR analysis in various tissues of Salvia miltiorrhiza","volume":"37","author":"Yang","year":"2010","journal-title":"Mol. Biol. Rep."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"616","DOI":"10.1111\/j.1365-313X.2005.02617.x","article-title":"Gateway-compatible vectors for plant functional genomics and proteomics","volume":"45","author":"Earley","year":"2006","journal-title":"Plant J."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"14850","DOI":"10.1021\/acs.jafc.0c05902","article-title":"JA-Responsive Transcription Factor SmMYB97 Promotes Phenolic Acid and Tanshinone Accumulation in Salvia miltiorrhiza","volume":"68","author":"Li","year":"2020","journal-title":"J. Agric. Food Chem."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Wang, B., Niu, J., Li, B., Huang, Y., Han, L., Liu, Y., Zhou, W., Hu, S., Li, L., and Wang, D. (2018). Molecular Characterization and Overexpression of SmJMT Increases the Production of Phenolic Acids in Salvia miltiorrhiza. Int. J. Mol. Sci., 19.","DOI":"10.3390\/ijms19123788"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/25\/23\/13223\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T16:50:38Z","timestamp":1760115038000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/25\/23\/13223"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,12,9]]},"references-count":69,"journal-issue":{"issue":"23","published-online":{"date-parts":[[2024,12]]}},"alternative-id":["ijms252313223"],"URL":"https:\/\/doi.org\/10.3390\/ijms252313223","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,12,9]]}}}