Johanna Myllyharju

Johanna Myllyharju

Ph.D., Professor

Biocenter Oulu and Faculty of Biochemistry and Molecular Medicine

134 peer reviewed publications in international journals. The publications have been cited 7891 times, of which the most cited publication accounts for 735 citations. The author has a h-index of 47.00. (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, 2020-2024, Sigrid Juselius Foundation 2017-2022, FibroGen Inc. 2002-

For research infrastructure: EU Infrafrontier2020, Academy of Finland FIRI2015 and FIRI2016 for Biocenter Finland, FIRI2018 for Infrarontier FI


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

  • Byts, Nadiya; Sharma, Subodh; Laurila, Jenny; Paudel, Prodeep; Miinalainen, Ilkka; Ronkainen, Veli-Pekka; Hinttala, Reeta; Törnquist, Kid; Koivunen, Peppi; Myllyharju, Johanna (2020) Transmembrane prolyl 4-Hydroxylase is a novel regulator of calcium signaling in astrocytes. - eNeuro Online ahead of print, Online ahead of print . [Original]
  • Salo, Antti; Myllyharju, Johanna (2021) Prolyl and lysyl hydroxylases in collagen synthesis. - Experimental dermatology 30 (1), 38-49 . [Original] [Self-archived]
  • Tolonen, Jussi-Pekka; Heikkilä, Minna; Malinen, Marjo; Lee, Hang-Mao; Palvimo, Jorma J.; Wei, Gong-Hong; Myllyharju, Johanna (2019) A long hypoxia-inducible factor 3 isoform 2 is a transcription activator that regulates erythropoietin. - Cellular and molecular life sciences 77 (18), 3627-3642 . [Original] [Self-archived]
  • Cockman, Matthew E.; Lippl, Kerstin; Tian, Ya-Min; Pegg, Hamish B.; Figg, William D., Jnr; Abboud, Martine I.; Heilig, Raphael; Fischer, Roman; Myllyharju, Johanna; Schofield, Christopher J.; Ratcliffe, Peter J. (2019) Lack of activity of recombinant HIF prolyl hydroxylases (PHDs) on reported non-HIF substrates. - Elife 8, e46490 . [Original] [Self-archived]
  • Rappu, P.; Salo, A.M.; Myllyharju, J.; Heino, J. (2019) Role of prolyl hydroxylation in the molecular interactions of collagens. - Essays in biochemistry 63 (3), 325-335 . [Original] [Self-archived]
  • Leinonen, Henri; Koivisto, Hennariikka; Lipponen, Henna-Riikka; Matilainen, Anna; Salo, Antti M.; Dimova, Elitsa Y.; Hämäläinen, Elina; Staven, Saara; Miettinen, Pasi; Myllyharju, Johanna; Koivunen, Peppi; Tanila, Heikki (2019) Null mutation in P4h-tm leads to decreased fear and anxiety and increased social behavior in mice. - Neuropharmacology 153, 63-72 . [Original] [Self-archived]
  • Murthy, Abhinandan V.; Sulu, Ramita; Koski, M. Kristian; Tu, Hongmin; Anantharajan, Jothi; Sah-Teli, Shiv K.; Myllyharju, Johanna; Wierenga, Rik K. (2018) Structural enzymology binding studies of the peptide-substrate-binding domain of human collagen prolyl 4-hydroxylase (type-II): High affinity peptides have a PxGP sequence motif. - Protein Science 27 (9), 1692-1703 . [Original]
  • Raykhel, Irina; Moafi, Fazeh; Myllymäki, Satu M.; Greciano, Patricia G.; Matlin, Karl S.; Moyano, Jose V.; Manninen, Aki; Myllyharju, Johanna (2018) BAMBI is a novel HIF1-dependent modulator of TGF beta-mediated disruption of cell polarity during hypoxia. - Journal of Cell Science 131 (10), UNSP jcs210906 . [Original]
  • Sipilä, Kalle H.; Drushinin, Kati; Rappu, Pekka; Jokinen, Johanna; Salminen, Tiina A.; Salo, Antti M.; Käpylä, Jarmo; Myllyharju, Johanna; Heino, Jyrki (2018) Proline hydroxylation in collagen supports integrin binding by two distinct mechanisms. - Journal of biological chemistry 293 (20), 7645-7658 . [Original]
  • Ullah, Karim; Rosendahl, Ann-Helen; Izzi, Valerio; Bergmann, Ulrich; Pihlajaniemi, Taina; Mäki, Joni M.; Myllyharju, Johanna (2017) Hypoxia-inducible factor prolyl-4-hydroxylase-1 is a convergent point in the reciprocal negative regulation of NF-κB and p53 signaling pathways. - Scientific Reports 7 (1), 17220 . [Original]
  • 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]
  • Sharma, Urvashi; Carrique, Loïc; Vadon-Le Goff, Sandrine; Mariano, Natacha; Georges, Rainier-Numa; Delolme, Frederic; Koivunen, Peppi; Myllyharju, Johanna; Moali, Catherine; Aghajari, Nushin; Hulmes, David J. S. (2017) Structural basis of homo- and heterotrimerization of collagen I. - Nature Communications 8, 14671 . [Original] [Self-archived]

Thesis supervisions

Supervised 23 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