{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T04:37:40Z","timestamp":1787027860599,"version":"build-2736575974"},"reference-count":128,"publisher":"MDPI AG","issue":"4","license":[{"start":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T00:00:00Z","timestamp":1775001600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Natural Science Foundation of Qinghai Province","award":["2025-ZJ-717"],"award-info":[{"award-number":["2025-ZJ-717"]}]},{"name":"Qinghai Province Kunlun Talent \u201cYoung and Middle-aged Scientific and Technological Talent\u201d Support Project","award":["2023QHSKXRCTJ28"],"award-info":[{"award-number":["2023QHSKXRCTJ28"]}]},{"name":"Major Science and Technology Project of Qinghai Province","award":["2023-SF-A5"],"award-info":[{"award-number":["2023-SF-A5"]}]},{"name":"2025 Graduate Student Innovation Project of Qinghai Minzu University","award":["54M2025006"],"award-info":[{"award-number":["54M2025006"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["CIMB"],"abstract":"<jats:p>Medicago ruthenica L., a superior forage crop within the genus Medicago (Fabaceae), is endowed with remarkable stress tolerance and an abundance of bioactive compounds, conferring significant ecological and forage value. Existing reviews primarily focus on a single research direction, and the most recent findings are dated, failing to cover breakthroughs at the molecular level. This paper systematically synthesizes the latest research progress in five key areas: genetic diversity and genomic studies, biotic stress responses, abiotic stress tolerance mechanisms (drought, salinity, and low temperature, etc.), utilization (including genetic breeding, ecological restoration, and forage development), and future research prospects. This review addresses critical gaps in existing literature, particularly regarding advances in genomic sequencing, biotic stresses, and research on stress-associated microorganisms. Research indicates that M. ruthenica exhibits extensive genetic diversity, and its genome contains numerous positive selection signals associated with stress resistance. It can tolerate multiple abiotic and biotic stresses through morphoplasticity, physiological metabolic regulation, and transcriptional regulation. Furthermore, its symbiosis with microorganisms such as rhizobia significantly enhances its stress tolerance. M. ruthenica demonstrates outstanding application potential in degraded grassland restoration and high-quality forage production. Future research should focus on mining stress-resistant genes, optimizing molecular breeding techniques, and integrating artificial intelligence into breeding practices. That will facilitate its transformation from a regional endemic resource to a commercially viable functional species, thereby providing robust support for ecological security and the sustainable development of grassland-based livestock husbandry in cold and arid regions.<\/jats:p>","DOI":"10.3390\/cimb48040365","type":"journal-article","created":{"date-parts":[[2026,4,1]],"date-time":"2026-04-01T08:08:34Z","timestamp":1775030914000},"page":"365","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["From \u201cOmics\u201d to Field: Deciphering the Stress Adaptation Networks and Breeding Potential of Medicago ruthenica L."],"prefix":"10.3390","volume":"48","author":[{"given":"Chen","family":"Zhang","sequence":"first","affiliation":[{"name":"College of Ecology Environment and Resources, Qinghai Minzu University, Xining 810007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingfang","family":"Shen","sequence":"additional","affiliation":[{"name":"College of Ecology Environment and Resources, Qinghai Minzu University, Xining 810007, China"},{"name":"Key Laboratory of Resources Chemistry and Ecological Environment Protection on the Qinghai-Tibetan Plateau, State Ethnic Affairs Commission, Xining 810007, China"},{"name":"Key Laboratory of High-Value Utilization of Characteristic Economic Plants in Qinghai Province, Xining 810007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leping","family":"Qi","sequence":"additional","affiliation":[{"name":"College of Ecology Environment and Resources, Qinghai Minzu University, Xining 810007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinxin","family":"Sun","sequence":"additional","affiliation":[{"name":"College of Ecology Environment and Resources, Qinghai Minzu University, Xining 810007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,4,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Shi, S., Guo, Y., and Zhi, H. (Plant Cell Environ., 2025). Exploring the Sustainable Utilization of Alfalfa Within the Phytomedicine Domain, Plant Cell Environ., Epub ahead of printing.","DOI":"10.1111\/pce.70014"},{"key":"ref_2","first-page":"1845","article-title":"Research Progress and Potential Applications for the Abiotic Stress-Resistant Forage Plant Medicago ruthenica","volume":"37","author":"Wu","year":"2020","journal-title":"Pratac. Sci."},{"key":"ref_3","first-page":"1669","article-title":"Observation and Comparison of the Pollen Morphology in Three Relative Leguminosae Forage Species","volume":"24","author":"Zhu","year":"2023","journal-title":"J. Plant Genet. Resour."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1007\/s11032-018-0905-z","article-title":"Development of Simple Sequence Repeat (SSR) Markers in Medicago ruthenica and Their Application for Evaluating Outcrossing Fertility under Open-Pollination Conditions","volume":"38","author":"Li","year":"2018","journal-title":"Mol. Breed."},{"key":"ref_5","first-page":"2005","article-title":"Genetic Variation of Allozymes in Medicago ruthenica","volume":"28","author":"Cao","year":"2008","journal-title":"Acta Bot. Boreal.-Occid. Sin."},{"key":"ref_6","first-page":"6","article-title":"Principal Component and Cluster Analysis of Phenotypic Diversity of Wild Medicago ruthenica Germplasm Resources in Three Ecological Regions of Inner Mongolia","volume":"45","author":"Hao","year":"2017","journal-title":"J. North. Agric."},{"key":"ref_7","first-page":"663","article-title":"Fine-Scale Genetic Structure of Medicago ruthenica Trautv","volume":"38","author":"Liu","year":"2020","journal-title":"Plant Sci. J."},{"key":"ref_8","first-page":"229","article-title":"Identification of Medicago ruthenica Hybrids and Heterosis Analysis of Main Agronomic Traits in F1 and F2","volume":"32","author":"Dong","year":"2023","journal-title":"Acta Pratacult. Sin."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"3118","DOI":"10.1111\/mec.16925","article-title":"Genomic Divergence between Two Sister Medicago Species Triggered by the Quaternary Climatic Oscillations on the Qinghai\u2013Tibet Plateau and Northern China","volume":"32","author":"Guo","year":"2023","journal-title":"Mol. Ecol."},{"key":"ref_10","first-page":"125","article-title":"Influence of Ageing on Genetic Integrity of Trigonella ruthenica L","volume":"20","author":"Su","year":"2012","journal-title":"Acta Agrestia Sin."},{"key":"ref_11","first-page":"84","article-title":"Effect of Aging Treatment on Changes of Genetic Integrity in Medicago ruthenica Germplasm","volume":"30","author":"Wang","year":"2010","journal-title":"Grassl. Turf."},{"key":"ref_12","first-page":"11","article-title":"Genetic Difference and Genetic Diversity Analysis of Melilotoides ruthenica Strains Based on ISSR Markers","volume":"32","author":"Wang","year":"2010","journal-title":"Chin. J. Grassl."},{"key":"ref_13","first-page":"1136","article-title":"Analysis of Genetic Diversity and Population Genetic Structure of Medicago archiducis-nicolai and Medicago ruthenica Populations Based on cpDNA trnL-trnF Sequences","volume":"33","author":"Wu","year":"2016","journal-title":"Pratac. Sci."},{"key":"ref_14","first-page":"124","article-title":"Comparative Analysis of Genetic Diversity of Medicago ruthenica (L.) in China Using AFLP and SSR Markers","volume":"32","author":"Li","year":"2017","journal-title":"Acta Agric. Boreal.-Sin."},{"key":"ref_15","unstructured":"Li, H.Y. (2008). Genetic Diversity Research of Germplasm Resources in Medicago ruthenica. [Ph.D. Thesis, Inner Mongolia Agricultural University]. (In Chinese)."},{"key":"ref_16","first-page":"34","article-title":"Genetic diversity of Medicago ruthenica based on SSR markers","volume":"30","author":"Li","year":"2008","journal-title":"Chin. J. Grassl."},{"key":"ref_17","first-page":"2415","article-title":"Genetic Diversity of Medicago ruthenica Strains Based on RAPD Markers","volume":"29","author":"Zhao","year":"2009","journal-title":"Acta Bot. Boreal.-Occid. Sin."},{"key":"ref_18","first-page":"65","article-title":"Analtsis of the Genetic Diversity of Medicago ruthenica by Phenotype and SSR Marker in China","volume":"39","author":"Li","year":"2011","journal-title":"J. Northwest A F Univ.-Nat. Sci. Ed."},{"key":"ref_19","unstructured":"Li, Z.Y. (2011). The Mechanism Studies on Genetic Diversity in Medicago ruthenica Germplasm Resources. [Ph.D Thesis, Chinese Academy of Agricultural Sciences]. (In Chinese)."},{"key":"ref_20","first-page":"52","article-title":"Genetic Diversity Studies of Medicago ruthenica in Inner Mongolia by ISSR","volume":"31","author":"Li","year":"2011","journal-title":"Acta Bot. Boreal.-Occid. Sin."},{"key":"ref_21","first-page":"107","article-title":"Genetic Diversity of Three Ecological Types of Wild Medicago ruthenica Germplasm Revealed by ISSR and SSR","volume":"21","author":"Li","year":"2012","journal-title":"Acta Pratacult. Sin."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1007\/s10722-012-9950-3","article-title":"Analysis of Genetic Diversity of Ruthenia Medic (Medicago ruthenica (L.) Trautv) in Inner Mongolia Using ISSR and SSR Markers","volume":"60","author":"Li","year":"2013","journal-title":"Genet. Resour. Crop. Evol."},{"key":"ref_23","first-page":"94","article-title":"Genetic Diversity of ISSR Markers and Environment Factors Correlation Analysis of Medicago ruthenica in China","volume":"29","author":"Li","year":"2014","journal-title":"Acta Agric. Boreal.-Sin."},{"key":"ref_24","first-page":"26","article-title":"Study on Taxonomy of Medicago ruthenica L. Using ISSR","volume":"37","author":"Liu","year":"2015","journal-title":"Chin. J. Grassl."},{"key":"ref_25","first-page":"20","article-title":"AFLP-Based Genetic Diversity Analysis of 49 Wild Medicago ruthenica Germplasm Resources","volume":"38","author":"Li","year":"2016","journal-title":"Chin. J. Grassl."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1186\/s12915-021-01033-0","article-title":"The Genome of a Wild Medicago Species Provides Insights into the Tolerant Mechanisms of Legume Forage to Environmental Stress","volume":"19","author":"Wang","year":"2021","journal-title":"BMC Biol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1641","DOI":"10.1111\/1755-0998.13363","article-title":"Genomic Analysis of Medicago ruthenica Provides Insights into Its Tolerance to Abiotic Stress and Demographic History","volume":"21","author":"Yin","year":"2021","journal-title":"Mol. Ecol. Resour."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1080\/23802359.2020.1845578","article-title":"Complete Chloroplast Genome of a High-Quality Forage in North China, Medicago ruthenica (Fabaceae: Trifolieae)","volume":"6","author":"Xie","year":"2021","journal-title":"Mitochondrial DNA B"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2688","DOI":"10.1080\/23802359.2021.1966328","article-title":"The Complete Chloroplast Genome of Medicago ruthenica Cv. \u2018Taihang\u2019 (Fabaceae) from Shanxi, China","volume":"6","author":"Xu","year":"2021","journal-title":"Mitochondrial DNA B"},{"key":"ref_30","unstructured":"Yan, W.H. (2021). Characterization of Chloroplast Genomes of Alfalfa (Medicago sativa L.) and Medicago ruthenica, Institute of Grassland Research, Chinese Academy of Agricultural Sciences. (In Chinese)."},{"key":"ref_31","first-page":"2323","article-title":"Screening of DNA Barcodes for Medicago archiducis-nicolai and Its Relative Medicago ruthenica","volume":"31","author":"Wang","year":"2023","journal-title":"Acta Agrestia Sin."},{"key":"ref_32","first-page":"10","article-title":"Analysis of Chloroplast Genome and Phylogeny of Yellow Flowered Type Medicago ruthenica","volume":"46","author":"Wen","year":"2024","journal-title":"Chin. J. Grassl."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Zhang, T., Li, M., Zhu, X., Li, S., Guo, M., Guo, C., and Shu, Y. (2024). Comparative Chloroplast Genomes Analysis Provided Adaptive Evolution Insights in Medicago ruthenica. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25168689"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Tian, Y., Wu, Z., Tian, C., Yang, Y., and Li, Z. (2025). Phylogenetic Classification and Genetic Insights from the Complete Mitochondrial Genome of Medicago ruthenica. Front. Plant Sci., 16.","DOI":"10.3389\/fpls.2025.1648505"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"406","DOI":"10.1111\/jipb.70098","article-title":"Structural Variation Drives Rhizome Innovation and Adaptive Divergence in Sister Medicago Species","volume":"68","author":"Hu","year":"2025","journal-title":"J. Integr. Plant Biol."},{"key":"ref_36","first-page":"43","article-title":"Occurrence and Control Strategies of Alfalfa Diseases and Pests in China","volume":"46","author":"Li","year":"2019","journal-title":"Tianjin Agric. For. Sci. Technol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2432","DOI":"10.1094\/PDIS-11-24-2341-PDN","article-title":"First Report of Powdery Mildew Caused by Erysiphe pisi on Medicago ruthenica in Inner Mongolia, China","volume":"109","author":"Tong","year":"2025","journal-title":"Plant Dis."},{"key":"ref_38","first-page":"3733","article-title":"Genetic Diversity Analysis of Medicago ruthenica L. Based on Phenotypic Traits","volume":"32","author":"Du","year":"2024","journal-title":"Acta Agrestia Sin."},{"key":"ref_39","first-page":"90","article-title":"Isolation, Identification and Weed Control Effect Study of Alternaria alternata Strain TL-3 from Cenchrus longispinus","volume":"47","author":"Wang","year":"2025","journal-title":"Chin. J. Grassl."},{"key":"ref_40","first-page":"2765","article-title":"Investigation and Research on Plant Fungal Diseases of Natural Grassland in Western China","volume":"33","author":"Xu","year":"2025","journal-title":"Acta Agrestia Sin."},{"key":"ref_41","first-page":"775","article-title":"Analysis of Volatiles from Forage Legume Medicago ruthenica under Different Conditions and Electroantennographic Response of Beet Webworm Loxostege sticticalis","volume":"49","author":"Bai","year":"2022","journal-title":"J. Plant Prot."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Tong, C., Zhang, Y., and Shi, F. (2023). Genome-Wide Identification and Analysis of the NLR Gene Family in Medicago ruthenica. Front. Genet., 13.","DOI":"10.3389\/fgene.2022.1088763"},{"key":"ref_43","first-page":"381","article-title":"The Ecological Characteristics of Food Habits for Four Herbivores in Gannan Meadow","volume":"52","author":"Su","year":"2017","journal-title":"Chin. J. Zool."},{"key":"ref_44","first-page":"43","article-title":"Short-Term Photosynthetic Responses of Medicago ruthenica Var. Inschanicus to Simulated Yak and Tibetan Sheep Trampling and Rainfall","volume":"26","author":"Xiao","year":"2017","journal-title":"Acta Pratacult. Sin."},{"key":"ref_45","first-page":"348","article-title":"Effects of Simulated Yak and Tibetan Sheep Trampling and Rainfall on Root Characteristics of Medicago ruthenica Var. inschanica in Alpine Meadow","volume":"26","author":"Xiao","year":"2018","journal-title":"Acta Agrestia Sin."},{"key":"ref_46","first-page":"1976","article-title":"Effects of Simulated Yak and Tibetan Sheep Trampling and Rainfall on Sexual Reproduction Characteristics of Medicago ruthenica var. inschanica in Alpine Meadow","volume":"37","author":"Xiao","year":"2018","journal-title":"Chin. J. Ecol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1007\/s10661-018-6896-8","article-title":"Yak and Tibetan Sheep Trampling Inhibit Reproductive and Photosynthetic Traits of Medicago ruthenica var. Inschanica","volume":"190","author":"Xiao","year":"2018","journal-title":"Environ. Monit. Assess."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"108522","DOI":"10.1016\/j.agee.2023.108522","article-title":"Different Grazers and Grazing Practices Alter the Growth, Soil Properties, and Rhizosphere Soil Bacterial Communities of Medicago ruthenica in the Qinghai-Tibetan Plateau Grassland","volume":"352","author":"Zhang","year":"2023","journal-title":"Agric. Ecosyst. Environ."},{"key":"ref_49","first-page":"1248","article-title":"Spatial Pattern of Degraded Alpine Communities Invaded by Stellera chamaejasme and Environmental Interpretation in the Central Qilian Mountain","volume":"51","author":"Liu","year":"2024","journal-title":"J. Plant Prot."},{"key":"ref_50","unstructured":"Xu, D.Q., Li, Y.Y., Hao, L.F., Tian, T., Guo, L.Z., Qi, D.J.Y., Lin, K.J., Ji, Y., and Su, H.H. (2025). Allelopathy Effects of Aqueous Extracts from Solanum Rostratum on Seed Germination of Four Excellent Forages. Pratac. Sci., (In Chinese)."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"864","DOI":"10.3390\/plants14060864","article-title":"Sustainable Weed Management: The Effects of Applying Pre- and Post-Emergence Herbicides to Medicago ruthenica","volume":"14","author":"Li","year":"2025","journal-title":"Plants"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1007\/s00284-025-04410-y","article-title":"Correlation Analysis of Soil Microbial Communities and Physicochemical Properties with Growth Characteristics of Medicago ruthenica in Different Habitats","volume":"82","author":"Wan","year":"2025","journal-title":"Curr. Microbiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1002\/glr2.70021","article-title":"Cuticular Waxes in Alpine Grassland Plants: Chemical Diversity, Biosynthesis, and Ecological Adaptation with Biotechnological Insights","volume":"4","author":"Xu","year":"2025","journal-title":"Grassl. Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1590\/1678-4685-gmb-2017-0284","article-title":"Transcriptome Sequencing and Expression Profiling of Genes Involved in the Response to Abiotic Stress in Medicago ruthenica","volume":"41","author":"Shu","year":"2018","journal-title":"Genet. Mol. Biol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Wu, R., Xu, B., and Shi, F. (2022). MrERF, MrbZIP, and MrSURNod of Medicago ruthenica Are Involved in Plant Growth and Abiotic Stress Response. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.907674"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"3498","DOI":"10.1002\/ldr.3930","article-title":"Root System Plays an Important Role in Responses of Plant to Drought in the Steppe of China","volume":"32","author":"Jiang","year":"2021","journal-title":"Land Degrad. Dev."},{"key":"ref_57","first-page":"489","article-title":"Evaluation on Drought Resistance of 10 Wild Medicago ruthenica L. Materials from Different Provenances at Germination Period","volume":"32","author":"Hou","year":"2024","journal-title":"Acta Agrestia Sin."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"109644","DOI":"10.1016\/j.ecolind.2022.109644","article-title":"Coarsened Soil Reduces Drought Resistance of Fibrous-Rooted Species on Degraded Steppe","volume":"145","author":"Jiang","year":"2022","journal-title":"Ecol. Indic."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.plaphy.2021.09.033","article-title":"Genome-Wide Identification of microRNAs Associated with Osmotic Stress and Elucidation of the Role of miR319 in Medicago ruthenica Seedlings","volume":"168","author":"Guo","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_60","first-page":"1901","article-title":"Medicago ruthenica (L.) Sojak. Cv. Zhilixing Response and Adaptation Strategy to Drought Stress and Rehydration","volume":"28","author":"Wu","year":"2020","journal-title":"Chin. J. Eco-Agric."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Wu, R., Xu, B., and Shi, F. (2022). Leaf Transcriptome Analysis of Medicago ruthenica Revealed Its Response and Adaptive Strategy to Drought and Drought Recovery. BMC Plant Biol., 22.","DOI":"10.1186\/s12870-022-03918-w"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Shi, R., Jiao, W., Yun, L., Zhang, Z., Zhang, X., Wang, Q., Li, Y., and Mi, F. (2021). Utilization of Transcriptome, Small RNA, and Degradome Sequencing to Provide Insights Into Drought Stress and Rewatering Treatment in Medicago ruthenica. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.675903"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Zi, N., Ren, W., Guo, H., Yuan, F., Liu, Y., and Fry, E. (2024). DNA Methylation Participates in Drought Stress Memory and Response to Drought in Medicago ruthenica. Genes, 15.","DOI":"10.3390\/genes15101286"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Wang, X., Dong, X., Li, P., Li, M., Wang, Z., Zhou, Q., Liu, Z., and Yan, L. (2025). Genome-Wide Identification of the GRAS Transcription Factor Family in Medicago ruthenica and Expression Analysis Under Drought Stress. Agronomy, 15.","DOI":"10.3390\/agronomy15020306"},{"key":"ref_65","first-page":"4528","article-title":"Identification of Key Drought-Responsive Genes in Upright Medicago ruthenica Sojak Cv. Zhilixing Based on Transcriptome Sequencing and WGCNA","volume":"58","author":"Mu","year":"2025","journal-title":"Sci. Agric. Sin."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Guo, Y., Zhao, X., Li, Y., Li, Z., Xiao, Q., Wang, Y., Zhang, X., and Ni, Y. (2021). Environment-Driven Adaptations of Leaf Cuticular Waxes Are Inheritable for Medicago ruthenica. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.620245"},{"key":"ref_67","unstructured":"Li, J.C., Han, B.C., Luo, D., Wang, Z.M., Liu, Z.P., and Ma, W.X. (2025). Identification and Expression Pattern Analysis of Medicago ruthenica SBP Gene Family. Pratac. Sci., (In Chinese)."},{"key":"ref_68","first-page":"179","article-title":"Identification of TCP Gene Family in Medicago ruthenica and Their Expression Pattern Analysis under Drought Stress","volume":"41","author":"Wang","year":"2025","journal-title":"Biotechnol. Bull."},{"key":"ref_69","first-page":"1","article-title":"Identification of the Dof Family Gene in Medicago ruthenica and Expression Analysis Pattern under Drought Stress","volume":"47","author":"Yuan","year":"2025","journal-title":"Chin. J. Grassl."},{"key":"ref_70","unstructured":"Zhang, L., Yan, W., Luo, D., Wang, Z., Liu, Z., and Zhou, Q. (2025). Genome-Wide Identification and Expression Analysis of the CIPK Gene Family in Medicago ruthenica under Drought Stress. Pratac. Sci., (In Chinese)."},{"key":"ref_71","first-page":"1","article-title":"Identification of HSF Gene Family in Medicago ruthenica and Analysis of Their Response to Drought Stress","volume":"47","author":"He","year":"2025","journal-title":"Chin. J. Grassl."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1007\/s00122-021-04002-x","article-title":"Genome-Wide Analysis of the Glutathione S-Transferase Family in Wild Medicago ruthenica and Drought-Tolerant Breeding Application of MruGSTU39 Gene in Cultivated Alfalfa","volume":"135","author":"Wang","year":"2022","journal-title":"Theor. Appl. Genet."},{"key":"ref_73","first-page":"564","article-title":"Effects of Different NaCl Concentrations on the Growth and Physiological Indices of Melilotoides ruthenica Seedling","volume":"22","author":"Yao","year":"2014","journal-title":"Acta Agrestia Sin."},{"key":"ref_74","first-page":"91","article-title":"Effects of NaCl Stress on Substances Linked to Osmotic Adjustment and on Photosynthetic Physiology of Melilotoides ruthenica in the Seedling Stage","volume":"24","author":"Yao","year":"2015","journal-title":"Acta Pratacult. Sin."},{"key":"ref_75","first-page":"189","article-title":"Effect of Salt Stress on Proline Accumulation and the Activities of the Key Enzymes Involved in Proline Metabolism in Medicago ruthenica Seedlings","volume":"27","author":"Zhong","year":"2018","journal-title":"Acta Pratacult. Sin."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Wei, X., Wang, J., Xu, C., Zhao, Y., Pu, X., Wang, W., and Lu, G. (2025). Analysis of Germination Characteristics and Metabolome of Medicago ruthenica in Response to Saline-Alkali Stress. Front. Plant Sci., 16.","DOI":"10.3389\/fpls.2025.1592555"},{"key":"ref_77","unstructured":"Hou, R.H. (2024). Salt Tolerance Evaluation and Gene Mining of 10 Medicago ruthenica L. Germplasms. [Master\u2019s Thesis, Inner Mongolia Agricultural University]. (In Chinese)."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1007\/s00299-024-03395-5","article-title":"Identification of Dehydrin Family Genes in Three Medicago Species and Insights into Their Tolerant Mechanism to Salt Stress","volume":"44","author":"Zhang","year":"2025","journal-title":"Plant Cell Rep."},{"key":"ref_79","first-page":"118","article-title":"Heterologous Expression of an SK2-Type Dehydrin Gene (MrDHN3) from Medicago ruthenica Enhances Escherichia coli Tolerance under Salt and High Temperature Stress","volume":"25","author":"Shen","year":"2016","journal-title":"Acta Pratacult. Sin."},{"key":"ref_80","first-page":"1944","article-title":"Cloning and Prokaryotic Expression of MrLEA2 from Medicago ruthenica","volume":"34","author":"Ma","year":"2014","journal-title":"Acta Bot. Boreal.-Occid. Sin."},{"key":"ref_81","first-page":"85","article-title":"Effects of Different Temperature Treatment on Seed Germination of Medicago ruthenica","volume":"39","author":"Yang","year":"2020","journal-title":"Seed"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1111\/grs.12384","article-title":"Effects of Low Temperature on Distribution and Metabolism Enzyme Activity of Carbon and Nitrogen of Medicago ruthenica (L.)","volume":"69","author":"Yang","year":"2023","journal-title":"Grassl. Sci."},{"key":"ref_83","unstructured":"Yang, Y.T. (2022). Carbon and Nitrogen Metabolism and Response to Low Temperature Stress of Medicago ruthenica. [Ph.D. Thesis, Inner Mongolia Agricultural University]. (In Chinese)."},{"key":"ref_84","unstructured":"Zhang, Y. (2013). Cloning and Expression Analysis of the Cold Resistance Related Genes of Medicago ruthenica. [Ph.D. Thesis, Inner Mongolia Agricultural University]. (In Chinese)."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1834","DOI":"10.1111\/pce.14845","article-title":"MrERF039 Transcription Factor Plays an Active Role in the Cold Response of Medicago ruthenica as a Sugar Molecular Switch","volume":"47","author":"Fu","year":"2024","journal-title":"Plant Cell Environ."},{"key":"ref_86","first-page":"3251","article-title":"Cloning and Expression Analysis of DNA Methyltransferase Gene MrMET1 in Medicago ruthenica","volume":"19","author":"Shen","year":"2021","journal-title":"Mol. Plant Breed."},{"key":"ref_87","first-page":"4657","article-title":"Cloning and Bioinformatics Analysis of a DNA Methyltransferase Gene MrDMR2 in Medicago ruthenica","volume":"19","author":"Shen","year":"2021","journal-title":"Mol. Plant Breed."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"e71895","DOI":"10.1002\/ece3.71895","article-title":"Combining Niche Breadth to Predict the Current and Future Distribution of Leguminosae Under Climate Change on the Qinghai-Xizang Plateau","volume":"15","author":"Chai","year":"2025","journal-title":"Ecol. Evol."},{"key":"ref_89","first-page":"793","article-title":"Effects of Heavy Metals: Copper, Cadmium and Lead on the Seed Germination and Seedling Growth of Leguminous Forage","volume":"23","author":"Yu","year":"2015","journal-title":"Acta Agrestia Sin."},{"key":"ref_90","unstructured":"Sai, N.G. (2023). Research on Soil Heavy Metal Pollution and Phytoremediation Potential\u2014Taking a Certain Industrial Park as an Example. [Master\u2019s Thesis, Gansu Agricultural University]. (In Chinese)."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"153696","DOI":"10.1016\/j.scitotenv.2022.153696","article-title":"Soil Amendment with a Humic Substance and Arbuscular Mycorrhizal Fungi Enhance Coal Mine Reclamation","volume":"823","author":"Zhao","year":"2022","journal-title":"Sci. Total Environ."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1007\/s42994-024-00193-1","article-title":"Enacting Partner Specificity in Legume\u2013Rhizobia Symbioses","volume":"6","author":"Yu","year":"2025","journal-title":"aBIOTECH"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1002\/ldr.5369","article-title":"Responses of Legume-Associated Rhizobacterial Communities to Plant Diversity and Soil Traits in Alpine Grassland","volume":"36","author":"Zhang","year":"2025","journal-title":"Land Degrad. Dev."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1787","DOI":"10.1111\/1365-2435.14343","article-title":"Effects of Long-Term Mowing on Leaf- and Root-Associated Bacterial Community Structures Are Linked to Functional Traits in 11 Plant Species from a Temperate Steppe","volume":"37","author":"Bai","year":"2023","journal-title":"Funct. Ecol."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Xing, J., Fan, W., Wang, J., and Shi, F. (2023). Variety-Driven Effect of Rhizosphere Microbial-Specific Recruitment on Drought Tolerance of Medicago ruthenica (L.). Microorganisms, 11.","DOI":"10.3390\/microorganisms11122851"},{"key":"ref_96","unstructured":"Xing, J. (2025). Correlation of Rhizosphere Bacteria in Medicago ruthenica (L.) with Drought Resistance. [Master\u2019s Thesis, Inner Mongolia Agricultural University]. (In Chinese)."},{"key":"ref_97","unstructured":"Li, M.Y. (2022). Characterization and Plant Growth-Promoting Mechanism of Cold-Adapted PGPB from Dominant Herbage in the Alpine Grassland of the Qilian Mountains. [Ph.D. Thesis, Gansu Agricultural University]. (In Chinese)."},{"key":"ref_98","unstructured":"Li, X.M. (2025). Screening of Rhizobia from Main Legume Forages in Gansu Province and Their Symbiotic Effects on Host Plants. [Master\u2019s Thesis, Gansu Agricultural University]. (In Chinese)."},{"key":"ref_99","first-page":"113","article-title":"Diversity and Community Structures of the Endophytic Actinomycetes Isolated from Medicago ruthenica","volume":"22","author":"Zhang","year":"2013","journal-title":"Acta Pratacult. Sin."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1002\/glr2.70019","article-title":"Characterization and Application of Highly Effective Rhizobia Isolated from Medicago ruthenica in Alpine Grassland","volume":"4","author":"Zhang","year":"2025","journal-title":"Grassl. Res."},{"key":"ref_101","first-page":"102","article-title":"Progress in Medicago ruthenica Breeding Study","volume":"40","author":"Zhang","year":"2018","journal-title":"Chin. J. Grassl."},{"key":"ref_102","first-page":"41","article-title":"Construction of Medicago ruthenica L. Core Germplasm Based on Phenotypic Traits","volume":"33","author":"Huang","year":"2025","journal-title":"Acta Agrestia Sin."},{"key":"ref_103","doi-asserted-by":"crossref","unstructured":"Zhu, L., Li, H., Wu, Z., Li, Z., Guo, M., Ning, B., Liu, W., Liu, Q., Liu, L., and Wang, Z. (2022). Targeted Metabolomics Provide Chemotaxonomic Insights of Medicago ruthenica, with Coupled Transcriptomics Elucidating the Mechanism Underlying Floral Coloration. Plants, 11.","DOI":"10.3390\/plants11182408"},{"key":"ref_104","first-page":"268","article-title":"Agrobacterium Rhizogenes-Mediated Transformation of Hairy Roots in Medicago ruthenica","volume":"28","author":"Wang","year":"2020","journal-title":"Acta Agrestia Sin."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"3167","DOI":"10.1002\/ldr.3882","article-title":"Plant Community Distribution Induced by Microtopography Due to Soil Cracks Developed in Overgrazed Alpine Meadows on the Tibetan Plateau","volume":"32","author":"Niu","year":"2021","journal-title":"Land Degrad. Dev."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"2821","DOI":"10.1111\/1365-2435.13934","article-title":"Divergent Contributions of Living Roots to Turnover of Different Soil Organic Carbon Pools and Their Links to Plant Traits","volume":"35","author":"Huang","year":"2021","journal-title":"Funct. Ecol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1235","DOI":"10.1002\/ldr.4529","article-title":"Species-Specific Soil Legacy Effects on Ecosystem Multifunctionality via Regulating Plant Overyielding","volume":"34","author":"Yan","year":"2023","journal-title":"Land Degrad. Dev."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Li, M., Qi, J., Lu, X., Zhang, T., and Yuan, Q. (2025). Soil Nitrogen Prevails in Controlling Alpine Meadow Productivity Despite Medicago ruthenica Reseeding and Phosphorus Application. Agronomy, 15.","DOI":"10.3390\/agronomy15081988"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"114587","DOI":"10.1016\/j.phytochem.2025.114587","article-title":"Unraveling Allelopathic Potential of Medicago ruthenica (L.) Trautv.: Insights from Allelopathic Effects and Untargeted Metabolomics","volume":"239","author":"Hua","year":"2025","journal-title":"Phytochemistry"},{"key":"ref_110","first-page":"1","article-title":"Research Progress for Seed Production Technology of Herbage and Turfgrass in China","volume":"45","author":"Mao","year":"2023","journal-title":"Chin. J. Grassl."},{"key":"ref_111","first-page":"1948","article-title":"Agronomic Traits of Different Medicago ruthenica Germplasms in the Tianzhu Alpine Region of Gansu Province","volume":"35","author":"Zhang","year":"2018","journal-title":"Pratac. Sci."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/S2095-3119(15)61036-X","article-title":"Rumen Methane Output and Fermentation Characteristics of Gramineous Forage and Leguminous Forage at Differing Harvest Dates Determined Using an In Vitro Gas Production Technique","volume":"15","author":"Zhong","year":"2016","journal-title":"J. Integr. Agric."},{"key":"ref_113","doi-asserted-by":"crossref","unstructured":"Wei, K., Xiang, H., Liu, Y., Zhang, X., and Yu, X. (2024). Mixed Cropping of Medicago ruthenica-Bromus inermis Exhibits Higher Yield and Quality Advantages in the Longxi Loess Plateau Region of Northwest China. Front. Sustain. Food Syst., 8.","DOI":"10.3389\/fsufs.2024.1411687"},{"key":"ref_114","first-page":"115","article-title":"The Study on Pod Dehiscence in Medicago ruthenica","volume":"43","author":"Wu","year":"2021","journal-title":"Chin. J. Grassl."},{"key":"ref_115","first-page":"303","article-title":"A Study on the Relationship between the Pod, Seed Development and the Occurrence of Seed Hardness of Melissitus ruthenicus","volume":"21","author":"Wang","year":"2012","journal-title":"Acta Pratacult. Sin."},{"key":"ref_116","first-page":"2992","article-title":"Study on Seed Features and Germination Characteristics of Wild Medicago ruthenica L. from Different Provenances","volume":"31","author":"Tang","year":"2023","journal-title":"Acta Agrestia Sin."},{"key":"ref_117","first-page":"132","article-title":"Study on the Method of Breaking Hardness of Wild Medicago ruthenica Seeds","volume":"41","author":"Yi","year":"2022","journal-title":"Seed"},{"key":"ref_118","first-page":"37","article-title":"Study on Breaking Hard Seed Effect of Different Treat Methods for Melilotoides ruthenica Strain 90-36","volume":"30","author":"Du","year":"2011","journal-title":"Seed"},{"key":"ref_119","first-page":"69","article-title":"Study on the Effect of Seed Treatment on Germination Rate of Melilotoides ruthenica in Medium of 1\/2 MS","volume":"31","author":"Tao","year":"2011","journal-title":"Grassl. Turf."},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"Zhang, Y., Shan, X., Zhao, Q., and Shi, F. (2022). The MicroRNA397a-LACCASE17 Module Regulates Lignin Biosynthesis in Medicago ruthenica (L.). Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.978515"},{"key":"ref_121","first-page":"27","article-title":"Effects of Endogenous Hormones on Pod Cracking of Medicago ruthenica","volume":"46","author":"Liu","year":"2024","journal-title":"Chin. J. Grassl."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1016\/j.plaphy.2022.06.013","article-title":"Mechanism of Pod Shattering in the Forage Legume Medicago ruthenica","volume":"185","author":"Guo","year":"2022","journal-title":"Plant Physiol. Biochem."},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"Zhu, L., Guo, M., Li, Z., Li, J., Li, H., Wu, Z., Tian, Y., and Zhao, C. (2025). Anatomical, Physiological, and Transcriptome Analyses Revealing Pod Shattering of Medicago ruthenica Associated with Pericarp Lignin Biosynthesis. Biomolecules, 15.","DOI":"10.3390\/biom15091269"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"894","DOI":"10.1007\/s40333-022-0026-8","article-title":"Effects of Mulch and Planting Methods on Medicago ruthenica Seed Yield and Soil Physical-Chemical Properties","volume":"14","author":"Wang","year":"2022","journal-title":"J. Arid. Land"},{"key":"ref_125","first-page":"60","article-title":"Effects of Planting Method on Seed Yield and Its Components in Medicago ruthenica in an Arid Area of Longzhong","volume":"30","author":"Wang","year":"2021","journal-title":"Acta Pratacult. Sin."},{"key":"ref_126","first-page":"54","article-title":"The Effects of Combined Nitrogen and Phosphorus Fertilization on Seed Yield and Quality of Medicago ruthenica in the Wuwei Irrigation District, Gansu Province","volume":"34","author":"Fan","year":"2025","journal-title":"Acta Pratacult. Sin."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"2868","DOI":"10.1007\/s42729-023-01242-2","article-title":"Arbuscular Mycorrhizal Fungi Alter the Interaction Effects Between Bacillus and Rhizobium on Root Morphological Traits of Medicago ruthenica L","volume":"23","author":"Yu","year":"2023","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_128","first-page":"113","article-title":"Determination of the Appropriate Harvest Time of Seed Production in Melilotoides ruthenicus","volume":"5","author":"Wang","year":"2015","journal-title":"Heilongjiang Anim. Sci. Vet. Med."}],"container-title":["Current Issues in Molecular Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1467-3045\/48\/4\/365\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,3]],"date-time":"2026-04-03T04:20:32Z","timestamp":1775190032000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1467-3045\/48\/4\/365"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,1]]},"references-count":128,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2026,4]]}},"alternative-id":["cimb48040365"],"URL":"https:\/\/doi.org\/10.3390\/cimb48040365","relation":{},"ISSN":["1467-3045"],"issn-type":[{"value":"1467-3045","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,1]]}}}