Johanna Myllyharju

Johanna Myllyharju

Ph.D., Professor

Biocenter Oulu and Faculty of Biochemistry and Molecular Medicine

128 peer reviewed publications in international journals. The publications have been cited 7415 times, of which the most cited publication accounts for 707 citations. The author has a h-index of 45.00. (Web of Science)


Professor Johanna Myllyharju is an expert on the key enzymes regulating hypoxia response and extracellular matrix synthesis. Oxygen and extracellular matrix homeostases are vital for normal functioning of cells and tissues, and disturbances in their balance are associated with many common diseases. Therefore, detailed information on these enzymes is essential for understanding their molecular mechanisms and biological effects in detail. This has already paved the way towards pharmacologic interventions, exemplified by a longterm industrial collaboration to develop enzyme inhibitors that activate the hypoxia response as therapeutic drugs, the first one being approved for treatment of severe anemia of chronic kidney diseases in 2018.

Research interests

  • Extracellular matrix biology
  • Hypoxia signaling

Research funding

For own research: Academy of Finland Academy Project 2016-2020, Academy of Finland Center of Excellence 2012-2017, Jane and Aatos Erkko Foundation 2015-2019, Sigrid Juselius Foundation 2017-2020, FibroGen Inc. 2002-

For research infrastructure: EU Infrafrontier2020, Academy of Finland FIRI2015 - Biocenter Finland, FIRI2018 - Infrarontier FI, FIRI 2016 - Biocenter Finland


Scientific merits

Professional and community activities

  • Academy of Finland: Chair of the Board 2019-2021
  • European Molecular Biology Conference (EMBC) council: Finnish delegate 2016 onwards
  • European Molecular Biology Laboratory (EMBL) council: Finnish delegate 2016 onwards
  • Biocenter Oulu: Director 2010-2018
  • Biocenter Finland: Member of the Board 2010-2018 (Chair 2016-2017, Vice Chair 2014-2015)
  • ESFRI Health and Food Strategic Working Group: Finnish representative 2015-2016
  • Academy of Finland: Member and Vice Chair of the Research Council for Health 2010-2015
  • COST Action TD0901 Hypoxia sensing, signalling and adaptation: Member of the Management Committee and Working Group Leader, 2009-2013

Research groups

  • Group Leader, principal investigator, Hypoxia & collagens
  • Vice Director, principal investigator, Finnish Center of Excellence in Cell-Extracellular Matrix Research 2012-2017

Selected publications

  • Myllyharju, Johanna & Kivirikko, Kari (2004) Collagens, modifying enzymes and their mutations in humans, flies and worms. - Trends in Genetics 20 (1), 33-43
  • Myllyharju, Johanna (2008) Prolyl 4-hydroxylases, key enzymes in the synthesis of collagens and regulation of the response to hypoxia, and their roles as treatment targets. - Annals of medicine 40, 402-417
  • Zou, Yaqun; Donkervoort, Sandra; Salo, Antti M.; Foley, A. Reghan; Barnes, Aileen M.; Hu, Ying; Makareeva, Elena; Leach, Meganne E.; Mohassel, Payam; Dastgir, Jahannaz; Deardorff, Matthew A.; Cohn, Ronald D.; DiNonno, Wendy O.; Malfait, Fransiska; Lek, Monkol; Leikin, Sergey; Marini, Joan C.; Myllyharju, Johanna; Bonnemann, Carsten G. (2017) P4HA1 mutations cause a unique congenital disorder of connective tissue involving tendon, bone, muscle and the eye. - Human Molecular Genetics 26 (12), 2207-2217 . [Original]
  • Hirsilä, Maija; Koivunen, Peppi; Gunzler, V.; Kivirikko, Kari & Myllyharju, Johanna (2003) Characterization of the human prolyl 4-hydroxylases that modify the hypoxia-inducible factor. - Journal of biological chemistry 278 (33), 30772-30780
  • Mäki, Joni; Sormunen, Raija; Lippo, Sari; Kaarteenaho-Wiik, Riitta; Soininen, Raija & Myllyharju, Johanna (2005) Lysyl oxidase is essential for normal development and function of the respiratory system and for the integrity of elastic and collagen fibers in various tissues. - American journal of pathology 167 (4), 927-936
  • Holster, Tiina; Pakkanen, Outi; Soininen, Raija; Sormunen, Raija; Nokelainen, Minna; Kivirikko, Kari & Myllyharju, Johanna (2007) Loss of assembly of the main basement membrane collagen, type IV, but not fibril-forming collagens and embryonic death in collagen prolyl 4-hydroxylase I null mice. - Journal of biological chemistry 282, 2512-2519
  • Koivunen, Peppi; Hirsilä, Maija; Remes, Anne; Hassinen, Ilmo; Kivirikko, Kari & Myllyharju, Johanna (2007) Inhibition of hypoxia-inducible factor (HIF) hydroxylases by citric acid cycle intermediates. Possible links between cell metabolism and stabilization of HIF. - Journal of biological chemistry 282, 4524-4532
  • Qi, H. H.; Ongusaha, P. P.; Myllyharju, Johanna; Cheng, D., Pakkanen, Outi; Shi, Y.; Lee, S. W.; Peng, J. & Shi, Y. (2008) Prolyl 4-hydroxylation regulates Argonaute 2 stability. - Nature 455, 421-424
  • Hyvärinen Jaana,Hassinen Ilmo,Sormunen Raija,Mäki Joni,Kivirikko Kari,Koivunen Peppi,Myllyharju Johanna (2010) Hearts of hypoxia-inducible factor prolyl 4-hydroxylase-2 hypomorphic mice show protection against acute ischemia-reperfusion injury.. - Journal of biological chemistry 285, 13646-13657
  • Laitala, Anu; Aro, Ellinoora; Walkinshaw, Gail; Mäki, Joni M; Rossi, Maarit; Heikkilä, Minna; Savolainen, Eeva-Riitta; Arend, Michael; Kivirikko, Kari I; Koivunen, Peppi; Myllyharju, Johanna (2012) Transmembrane prolyl 4-hydroxylase is a fourth prolyl 4-hydroxylase regulating EPO production and erythropoiesis. - Blood 120 (16), 3336-3344 . [Original]
  • Aro, Ellinoora; Salo, Antti M.; Khatri, Richa; Finnilä, Mikko; Miinalainen, Ilkka; Sormunen, Raija; Pakkanen, Outi; Holster, Tiina; Soininen, Raija; Prein, Carina; Clausen-Schaumann, Hauke; Aszodi, Attila; Tuukkanen, Juha; Kivirikko, Kari I.; Schipani, Ernestina; Myllyharju, Johanna (2015) Severe Extracellular Matrix Abnormalities and Chondrodysplasia in Mice Lacking Collagen Prolyl 4-Hydroxylase Isoenzyme II in Combination with a Reduced Amount of Isoenzyme I. - Journal of biological chemistry 290 (27), 16964-16978 . [Original]

Thesis supervisions

Supervised 22 PhD or MD-PhD theses to completion.

Research visits

  • Hans Kupczyk Guest Professor

    University of Ulm, Germany


Key enzyme regulators of the hypoxia response and collagen synthesis as therapeutic targets - Johanna Myllyharju

Maintenance of oxygen and extracellular matrix (ECM) homeostases is crucial for normal development and physiology, and several pathologies are asso