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TAKAYA Tomohide

Academic OrganizationInstitute of Agriculture, School of Science and Technology, Academic AssemblyTEL
Education and Research OrganizationDivision of Animal Science, Department of Agricultural and Life Science, Faculty of AgricultureFAX
PositionAssociate ProfessorMail Addressttakaya@shinshu-u.ac.jp
Address8304 Minami-minowa, Kami-ina, Nagano 399-4598, JapanWeb sitehttps://t-takaya.net/

Modified:02/14/2024

Profile

Assigned Class
Cellular Systems Unit, Integrated Bioscience Division, Department of Agriculture, Graduate School of Science and Technology
Integrated Bioscience Unit, Division of Biological and Agricultural Sciences, Department of Science and Technology, Graduate School of Medicine, Science and Technology
Research Field
Stem cell biology
Keywords:Stem cell , Skeletal muscle , Regeneration
Current Subject
Nucleic acid drugs for motor disorders
Keywords:Motor disorders , Muscle loss , Osteoporosis , Nucleic acid drug , Aptamer
Academic Societies
Academic Societies
Japanese Society of Anti-Aging Medicine
Nucleic Acids Therapeutics Society of Japan
Japan Poultry Science Association
Japan Society for Bioscience, Biotechnology, and Agrochemistry
Japanese Society of Animal Science
Japan Cardiovascular Association
Academic Background
Graduate School
Kyoto University , (医学専攻 , Graduate School of Medicine) , 2011
Okayama University , (分子・生物科学専攻 , Graduate School of Natural Science and Technology) , 2005

College
Okayama University , (Faculty of Science) , 2003

Degree
PhD (Medical Science) , Kyoto University
Awards
2020 , Young Scientist Award for Anti-Aging Medicine
2018 , Poster Presentation Award (Nakamura S)
2018 , Innovative Research Program Award
Research Career
Research Career
2023- , Associate Professor, Faculty of Agriculture, Shinshu University
2015-2023 , Assistant Professor, Faculty of Agriculture, Shinshu University
2014-2015 , Post-doctroral Fellow for Research Abroad, Japan Society for the Promotion of Science
2013-2015 , Post-doctoral Associate, Stem Cell Institute, University of Minnesota Medical School
2011-2012 , Research Fellow, Department of Vascular Physiology, National Cerebral and Cardiovascular Center Research Institute
2009-2011 , Research Fellow for Doctoral Course Students (DC2), Japan Society for the Promotion of Science
2005-2009 , Research Fellow, Division of Translational Research, National Hospital Organization Kyoto Medical Center

Research

Books, Articles, etc.
Articles
Myogenetic oligodeoxynucleotide induces myocardial differentiation of murine pluripotent stem cells.
Int J Mol Sci,24(18):14380 2023
Author:Ishioka M, Nihashi Y, Sunagawa Y, Umezawa K, Shimosato T, Kagami H, Morimoto T, Takaya T*.


Anti-nucleolin aptamer, iSN04, inhibits the inflammatory responses in C2C12 myoblasts by modulating the β-catenin/NF-κB signaling pathway.
Biochem Biophys Res Commun,664:1-8 2023
Author:Yamamoto M, Miyoshi M, Morioka K, Mitani T, Takaya T*.


Myogenetic oligodeoxynucleotides as anti-nucleolin aptamers inhibit the growth of embryonal rhabdomyosarcoma cells.
Biomedicines,10(11):2691 2022
Author:Nohira N, Shinji S, Nakamura S, Nihashi Y, Shimosato T, Takaya T*.


Myogenetic oligodeoxynucleotide restores differentiation and reverses inflammation of myoblasts aggravated by cancer-conditioned medium.
Muscles,1(2):111-120 2022
Author:Nihashi Y, Yamamoto M, Shimosato T, Takaya T*.


Identification of a novel osteogenetic oligodeoxynucleotide (osteoDN) that promotes osteoblast differentiation in a TLR9-independent manner.
Nanomaterials,12(10):1680 2022
Author:Nihashi Y, Miyoshi M, Umezawa K, Shimosato T, Takaya T*.


Three-dimensional structural analysis of mitochondria composing each subtype of fast-twitch muscle fibers in chicken.
J Vet Med Sci,84(6):809-816 2022
Author:Makida S, Kametani K, Hosotani M, Takahashi N, Iwasaki T, Hasegawa Y, Takaya T, Ueda H, Watanabe T.


Myogenetic oligodeoxynucleotide complexed with berberine promotes differentiation of chicken myoblasts.
Anim Sci J,92(1):e13597 2021
Author:Nihashi Y, Shinji S, Umezawa K, Shimosato T, Ono T, Kagami H, Takaya T*.


Myogenetic oligodeoxynucleotide (myoDN) recovers the differentiation of skeletal muscle myoblasts deteriorated by diabetes mellitus.
Front Physiol,12:679152 2021
Author:Nakamura S, Yonekura S, Shimosato T, Takaya T*.


Transcription of endogenous retrovirus group K members and their neighboring genes in chicken skeletal muscle myoblasts.
J Poult Sci,58(2):79-87 2021
Author:Takaya T*, Nihashi Y, Ono T, Kagami H.


Identification of the myogenetic oligodeoxynucleotides (myoDNs) that promote differentiation of skeletal muscle myoblasts by targeting nucleolin.
Front Cell Dev Biol,8:616706 2021
Author:Shinji S, Umezawa K, Nihashi Y, Nakamura S, Shimosato T, Takaya T*.


Physiological and pathological mitochondrial clearance is related to pectoralis major muscle pathogenesis in broilers with wooden breast syndrome.
Front Physiol,11:579 2020
Author:Hosotani M, Kawasaki T, Hasegawa Y, Wakasa Y, Hoshino M, Takahashi N, Ueda H, Takaya T, Iwasaki T, Watanabe T.


Berberine and palmatine inhibit the growth of human rhabdomyosarcoma cells.
Biosci Biotechnol Biochem,84(1):63-75 2020
Author:Shinji S, Nakamura S, Nihashi Y, Umezawa K, Takaya T*.


Distinct cell proliferation, myogenic differentiation, and gene expression in skeletal muscle myoblasts of layer and broiler chickens.
Sci Rep,9:16527 2019
Author:Nihashi Y, Umezawa K, Shinji S, Hamaguchi Y, Kobayashi H, Kono T, Ono T, Kagami H, Takaya T*.


Toll-like receptor ligand-dependent inflammatory responses in chick skeletal muscle myoblasts.
Dev Comp Immunol,91:115-122 2019
Author:Nihashi Y, Ono T, Kagami H, Takaya T*.


Theophylline suppresses interleukin-6 expression by inhibiting glucocorticoid receptor signaling in pre-adipocytes.
Arch Biochem Biophys,646:98-106 2018
Author:Mitani T, Takaya T, Harada N, Katayama S, Yamaji R, Nakamura S, Ashida H.


Autonomous xenogenic cell fusion of murine and chick skeletal muscle myoblasts.
Anim Sci J,88(11):1880-1885 2017
Author:Takaya T*, Nihashi Y, Kojima S, Ono T, Kagami H.


Lectin-like oxidized low-density lipoprotein receptor-1 plays an important role in vascular inflammation in current smokers.
J Atheroscler Thromb,20(6):585-590 2013
Author:Takanabe-Mori R, Ono K, Wada H, Takaya T, Ura S, Yamakage H, Satoh-Asahara N, Shimatsu A, Takahashi Y, Fujita M, Fujita Y, Sawamura T, Hasegawa K.


Roles of microRNAs and myocardial cell differentiation.
Prog Mol Biol Transl Sci,111:139-152 2012
Author:Takaya T, Nishi H, Horie T, Ono K, Hasegawa K.


Highly purified eicosapentaenoic acid increases interleukin-10 levels of peripheral blood monocytes in obese patients with dyslipidemia.
Diabetes Care,35(12):2631-2639 2012
Author:Satoh-Asahara N, Shimatsu A, Sasaki Y, Nakaoka H, Himeno A, Tochiya M, Kono S, Takaya T, Ono K, Wada H, Suganami T, Hasegawa K, Ogawa Y.


Distinct characteristics of circulating vascular endothelial growth factor-A and C levels in human subjects.
PLoS One,6(12):e29351 2011
Author:Wada H, Ura S, Kitaoka S, Satoh-Asahara N, Horie T, Ono K, Takaya T, Takanabe-Mori R, Akao M, Abe M, Morimoto T, Murayama T, Yokode M, Fujita M, Shimatsu A, Hasegawa K.


The discovery of LOX-1, its ligands and clinical significance.
Cardiovasc Drugs Ther,25(5):379-391 2011
Author:Yoshimoto R, Fujita Y, Kakino A, Iwamoto S, Takaya T, Sawamura T.


A natural p300-specific histone acetyltransferase inhibitor, curcumin, in addition to angiotensin-converting enzyme inhibitor, exerts beneficial effects on left ventricular systolic function after myocardial infarction in rats.
Circ J,75(9):2151-2159 2011
Author:Sunagawa Y, Morimoto T, Wada H, Takaya T, Katanasaka Y, Kawamura T, Yanagi S, Marui A, Sakata R, Shimatsu A, Kimura T, Kakeya H, Fujita M, Hasegawa K.


MicroRNA-27a regulates beta cardiac myosin heavy chain gene expression by targeting thyroid hormone receptor beta1 in neonatal rat ventricular myocytes.
Mol Cell Biol,31(4):744-755 2011
Author:Nishi H, Ono K, Horie T, Nagao K, Kinoshita M, Kuwabara Y, Watanabe S, Takaya T, Tamaki Y, Takanabe-Mori R, Wada H, Hasegawa K, Iwanaga Y, Kawamura T, Kita T, Kimura T.


Cyclin-dependent kinase 9 forms a complex with GATA4 and is involved in the differentiation of mouse ES cells into cardiomyocytes.
J Cell Physiol,226(1):248-254 2011
Author:Kaichi S, Takaya T, Morimoto T, Sunagawa Y, Kawamura T, Ono K, Shimatsu A, Baba S, Heike T, Nakahata T, Hasegawa K.


Lectin-like oxidized low-density lipoprotein receptor-1 is required for the adipose tissue expression of proinflammatory cytokines in high-fat diet-induced obese mice.
Biochem Biophys Res Commun,398(3):576-580 2010
Author:Takanabe-Mori R, Ono K, Sowa N, Wada H, Takaya T, Horie T, Satoh-Asahara N, Shimatsu A, Fujita M, Sawamura T, Hasegawa K.


Cell line-dependent differentiation of induced pluripotent stem cells into cardiomyocytes in mice.
Cardiovasc Res,88(2):314-323 2010
Author:Kaichi S, Hasegawa K, Takaya T, Yokoo N, Mima T, Kawamura T, Morimoto T, Ono K, Baba S, Doi H, Yamanaka S, Nakahata T, Heike T.


Left ventricular expression of lectin-like oxidized low-density lipoprotein receptor-1 in failing rat hearts.
Circ J,74(4):723-729 2010
Author:Takaya T, Wada H, Morimoto T, Sunagawa Y, Kawamura T, Takanabe-Mori R, Shimatsu A, Fujita Y, Sato Y, Fujita M, Kimura T, Sawamura T, Hasegawa K.


Cyclin-dependent kinase-9 is a component of the p300/GATA4 complex required for phenylephrine-induced hypertrophy in cardiomyocytes.
J Biol Chem,285(13):9556-9568 2010
Author:Sunagawa Y, Morimoto T, Takaya T, Kaichi S, Wada H, Kawamura T, Fujita M, Shimatsu A, Kita T, Hasegawa K.


Aldosterone signaling associates with p300/GATA4 transcriptional pathway during the hypertrophic response of cardiomyocytes.
Circ J,74(1):156-162 2010
Author:Yoshida Y, Morimoto T, Takaya T, Kawamura T, Sunagawa Y, Wada H, Fujita M, Shimatsu A, Kita T, Hasegawa K.


MicroRNA-1 and microRNA-133 in spontaneous myocardial differentiation of mouse embryonic stem cells.
Circ J,73(8):1492-1497 2009
Author:Takaya T, Ono K, Kawamura T, Takanabe R, Kaichi S, Morimoto T, Wada H, Kita T, Shimatsu A, Hasegawa K.


Up-regulated expression of microRNA-143 in association with obesity in adipose tissue of mice fed high-fat diet.
Biochem Biophys Res Commun,376(4):728-732 2008
Author:Takanabe R, Ono K, Abe Y, Takaya T, Horie T, Wada H, Kita T, Satoh N, Shimatsu A, Hasegawa K.


Myocardial regulation of p300 and p53 by doxorubicin involves ubiquitin pathways.
Circ J,72(9):1506-1511 2008
Author:Morimoto T, Fujita M, Kawamura T, Sunagawa Y, Takaya T, Wada H, Shimatsu A, Kita T, Hasegawa K.


Statins activate GATA-6 and induce differentiated vascular smooth muscle cells.
Biochem Biophys Res Commun,374(4):731-736 2008
Author:Wada H, Abe M, Ono K, Morimoto T, Kawamura T, Takaya T, Satoh N, Fujita M, Kita T, Shimatsu A, Hasegawa K.


The dietary compound curcumin inhibits p300 histone acetyltransferase activity and prevents heart failure in rats.
J Clin Invest,118(3):868-878 2008
Author:Morimoto T, Sunagawa Y, Kawamura T, Takaya T, Wada H, Nagasawa A, Komeda M, Fujita M, Shimatsu A, Kita T, Hasegawa K.


Identification of p300-targeted acetylated residues in GATA4 during hypertrophic responses in cardiac myocytes.
J Biol Chem,283(15):9828-9835 2008
Author:Takaya T, Kawamura T, Morimoto T, Ono K, Kita T, Shimatsu A, Hasegawa K.


Spaceflight results in increase of thick filament but not thin filament proteins in the paramyosin mutant of Caenorhabditis elegans.
Adv Space Res,41(5):816-823 2008
Author:Adachi R, Takaya T, Kuriyama K, Higashibata A, Ishioka N, Kagawa H.


Trichostatin A induces myocardial differentiation of monkey ES cells.
Biochem Biophys Res Commun,356(2):386-391 2007
Author:Hosseinkhani M, Hasegawa K, Ono K, Kawamura T, Takaya T, Morimoto T, Wada H, Shimatsu A, Prat SG, Suemori H, Nakatsuji N, Kita T.


C. elegans model for studying tropomyosin and troponin regulations of muscle contraction and animal behavior.
Adv Exp Med Biol,592:153-161 2007
Author:Kagawa H, Takaya T, Ruksana R, Anokye-Danso F, Amin MZ, Terami H.


Tissue expression of four troponin I genes and their molecular interactions with two troponin C isoforms in Caenorhabditis elegans.
Genes Cells,10(3):261-276 2005
Author:Ruksana R, Kuroda K, Terami H, Bando T, Kitaoka S, Takaya T, Sakube Y, Kagawa H.


Presentations
Identification of a candidate drug for the treatment of facioscapulohumeral muscular dystrophy.
Human Genetics Asia 2023 2023(Oct. 13)
Presenter:Yoshizawa T, Tomiyama Y, Shiba N, Takaya T, Miyazaki D, Nakada T, Shiba Y, Nakamura A, Kosho T.


Possible chicken meat production by ex-ovo embryo culture system and the application for poultry biotechnology.
The 68th International Congress of Meat Science and Technology 2022(Aug. 24)
Presenter:Shindo H, Saito S, Takizawa S, Murai K, Takaya T, Hiramatsu K, Kawahara T, Kagami H.


Application of a novel chick embryo culture system for developmental studies and chicken meat production.
The 68th International Congress of Meat Science and Technology 2022(Aug. 24)
Presenter:Murai K, Takizawa S, Saito S, Shindo H, Takaya T, Hiramatsu K, Kawahara T, Kagami H.


Avian stem cell technologies for cellular agriculture.
The 7th International Scientific Conference on Cultured Meat 2021(Nov. 30)
Presenter:Kagami H, Murai K, Takizawa S, Shindo H, Saito S, Takaya T, Hiramatsu K, Sheng G.


Discovery and application of myogenetic oligodeoxynucleotides (myoDNs).
International Joint Seminar 2019 Sejong University-Shinshu University-Kasetsart University 2019(Sep. 24)
Presenter:Takaya T.


Action mechanism of myogenetic oligodeoxynucleotides (myoDNs).
International Joint Seminar 2019 Sejong University-Shinshu University-Kasetsart University 2019(Sep. 24)
Presenter:Shinji S, Takaya T.


Expression of helix-loop-helix transcription factor Ascl4 induces myogenic program in embryonic stem cells.
Frontiers in Myogenesis Meeting 2019(Sep. 24)
Presenter:Asakura A, Kodaka Y, Watanabe S, Ito H, Verma M, Takaya T, Asakura Y, Kyba M.


Application of myogenetic oligodeoxynucleotide to diabetic skeletal muscle myoblasts.
日本核酸医薬学会第5回年会 2019(Jul. 11)
Presenter:Nakamura S, Umezawa K, Yonekura S, Shimosato T, Takaya T.


Induction of cardiac differentiation of pluripotent stem cells by myogenetic oligodeoxynucleotide.
日本核酸医薬学会第5回年会 2019(Jul. 10)
Presenter:Ishioka M, Nihashi Y, Umezawa K, Kagami H, Shimosato T, Takaya T.


Conformational difference between two types of biologically active oligodeoxynucleotides.
日本核酸医薬学会第5回年会 2019(Jul. 10)
Presenter:Umezawa K, Yamamoto Y, Shimosato T, Takaya T.


RNA sequencing for global gene expression associated with proliferation and differentiation in chicken skeletal muscle myoblasts.
The 2nd International Conference on Tropical Animal Science and Production 2019(Jul. 10)
Presenter:Nihashi Y, Umezawa K, Hamaguchi Y, Kobayashi H, Kono T, Ono T, Kagami H, Takaya T.


Myogenic differentiation-inducing oligodeoxynucleotide recovers the myoblast differentiation impaired by abnormal glucose concentration.
The 4th International Conference on Pharma-Food 2018(Nov. 15)
Presenter:Nakamura S, Yonekura S, Shimosato T, Takaya T.


Toll-like receptor ligand-dependent inflammatory responses in chick skeletal muscle myoblasts.
2018 International Symposium on Animal Science and Technology 2018(Aug. 10)
Presenter:Nihashi Y, Ono T, Kagami H, Takaya T.


Functional structure analyses of the DNA-alkaloid complex promoting myogenic differentiation.
2018 International Symposium on Animal Science and Technology 2018(Aug. 10)
Presenter:Shinji S, Umezawa K, Shimosato T, Takaya T.


Myogenic oligodeoxynucleotide, myoDN, induces myocardial differentiation of murine iPS cells.
2018 International Symposium on Animal Science and Technology 2018(Aug. 10)
Presenter:Ishioka M, Nihashi Y, Kagami H, Shimosato T, Takaya T.


Intermittent glucose stimulation impairs proliferation and differentiation of C2C12 myoblasts.
2018 International Symposium on Animal Science and Technology 2018(Aug. 10)
Presenter:Nakamura S, Yonekura S, Takaya T.


Developmental analysis of the donor mice ES cells in the recipient chick embryo.
2018 International Symposium on Animal Science and Technology 2018(Aug. 10)
Presenter:Asai M, Kusudo T, Kobayashi K, Watanabe K, Ono T, Takaya T, Tomioka I, Kagami H.


Comprehensive gene expression analyses between broiler and layer chick skeletal muscle myoblasts.
2018 International Symposium on Animal Science and Technology 2018(Aug. 09)
Presenter:Takaya T.


Distinct expression of TLR genes between broiler and layer chick skeletal muscle myoblasts.
The 11th Asian Pacific Poulty Conference 2018(Mar. 26)
Presenter:Nihashi Y, Ono T, Kagami H, Takaya T.


Novel DNA aptamer activating skeletal muscle stem cells.
Symposium on Advanced Materials and 3D Printing for Biomedical Applications 2017(Oct. 20)
Presenter:Takaya T.


Regulation of skeletal muscle stem cells for meat production.
Japan-Thailand International Symposium on Animal Biotechnology 2017(Jul. 05)
Presenter:Takaya T.


Autonomous xenogenic cell fusion of murine and chick skeletal muscle myoblasts.
Japan-Thailand International Symposium on Animal Biotechnology 2017(Jul. 05)
Presenter:Takaya T, Nihashi Y, Kojima S, Ono T, Kagami H.


Distinct cell characteristics between broiler and layer chick skeletal muscle myoblasts.
Japan-Thailand International Symposium on Animal Biotechnology 2017(Jul. 05)
Presenter:Nihashi Y, Ono T, Kagami H, Takaya T.


Application of the oligodeoxynucleotide from lactic acid bacteria that promotes myogenic differentiation.
Japan-Thailand International Symposium on Animal Biotechnology 2017(Jul. 05)
Presenter:Shinji S, Nihashi Y, Umezawa K, Shimosato T, Takaya T.


Efficient production of chicks by ex ovo culture system with novel avian assisted reproductive technology (AART).
Japan-Thailand International Symposium on Animal Biotechnology 2017(Jul. 05)
Presenter:Sato S, Sato T, Miki S, Ono T, Takaya T, Kagami H.

Research Grants
Other
2021 - 2021 , 脂肪細胞における筋形成型オリゴDNAの抗炎症作用の解明, 長野県科学振興会 令和3年度科学研究費助成金(学生採択:森岡一乃)
Research Activities
Journal Reviewer
2023- ,
Biomolecules
Epigenetics
Food Bioscience
Genes
International Journal of Biological Macromolecules
Pharmaceuticals
2022- ,
Bioengineering
Biomedicines
BMC Endocrine Disorders
Frontiers in Nutrition
Life
Journal of Functional Biomaterials
Metabolites
2021- ,
Advanced Science
BIOCELL
Cellular Signalling
Differentiation
Frontiers in Bioengineering and Biotechnology
Frontiers in Cell and Developmental Biology
Frontiers in Endocrinology
Frontiers in Molecular Biosciences
Frontiers in Veterinary Science
Heliyon
International Journal of Biological Sciences
Journal of Animal Science and Biotechnology
Journal of Poultry Science
Nutrients
Onco
Regenerative Therapy
Stem Cells
2020- ,
Animals
Antioxidants
Cells
Frontiers in Physiology
Histochemistry and Cell Biology
International Journal of Molecular Sciences
Journal of Medical Virology
Methods and Protocols
Poultry Science
2019- ,
Animal Science Journal
Current Pharmaceutical Design
Journal of Diabetes Research
Journal of Integrative Neuroscience
2018- ,
Frontiers in Immunology
Journal of Oncology
Journal of Veterinary Medical Science
Scientific Reports
2017- ,
Bioscience, Biotechnology, and Biochemistry
2015- ,
BMC Genomics
2014- ,
Stem Cells International
2013- ,
Circulation
2013- ,
PLoS One

Education

Lectures
Seminar for Freshmen (2017-2021, 2023-)
Reading of Foreign Articles I (2016)
Biochemistry (2017-2022)
Bioanalytical Chemistry
Special Lecture for Cutting Edge (2024-)
Career Design (2018, 2023)
Studies on Domestic Animals (2021)
Animal Science (2018)
Animal Cell Biology (2022-)
Hands-on Seminar on Farm Practices (2017, 2021, 2023)
Presentation II (2018-)
Selected Seminar (2017-)
Thesis Research II (2018-)
Advanced Lecture in Food, Agricultural and Environmental Sciences (2020, 2023)
Advanced Lecture in Integrated Bioscience (2016-)
Animal Life Science (2016-)
Compendium of Animal Life Science (2017-)
Animal Genetics and Breeding (2019)
Biostatistics for Animal Experiments (2016-)
Experiments in Animal Bioscience
Scientific English II (2017-)
Presentation I (2017-)
Thesis Research I (2017-)
Thesis Research (2018-)
Advanced Seminar for Integrated Bioscience (2016, 2023)
Advanced Lecture in Animal Biotechnology (2018-)
Volunteer I (2018, 2023)
Volunteer II (2023)
Volunteer III (2023)
Volunteer IV (2023)
Internship I (2018, 2023)
Internship II (2018, 2023)
Internship III (2018-2019, 2023)
Advanced Internship (2016, 2023)
Special Research (2019-)
Advanced Seminar for Cellular Systems I (2018-)
Advanced Seminar for Cellular Systems II (2018-)
Advanced Experimental Course on Cellular Systems (2018-)
Molecular Biotechnology (2018-)
Special Seminar (2020-)
Special Seminar on Research Subject

Social Activities

Social Activitie
Media Appearances
20231010 , EurekAlert! , DNA aptamer finds novel application in regulating cell differentiation
20220523 , EurekAlert! , Identification of osteogenetic oligodeoxynucleotide that promotes bone differentiation
20210526 , EurekAlert! , Skeletal muscle loss exacerbated by diabetes improved with oligo DNA
20210125 , EurekAlert! , Identification of oligo-DNA that promotes skeletal muscle differentiation