Johanna Myllyharju

Johanna Myllyharju

Ph.D., Professor, Scientific director of Biocenter Oulu

Scientific director
Biocenter Oulu

115 peer reviewed publications in international journals. The publications have been cited 6020 times, of which the most cited publication accounts for 591 citations. The author has a h-index of 40.00. (Web of Science)


Professor Johanna Myllyharju is a scientific director of Biocenter Oulu, University of Oulu, 2010-2019.

Biocenter Oulu is the oldest of the Finnish biocenters, established in 1986. It is a focus institute within the Molecular and environmental basis of life-long health strategic research focus area of the University of Oulu. Biocenter Oulu coordinates together with the Medical Research Center the Fibrosis Diseasome PROFI3 theme of University of Oulu.

Research interests

  • Extracellular matrix biology
  • Hypoxia signaling

Ongoing outside research funding

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

For research infrastructure: Academy of Finland FIRI 2016 - Biocenter Finland, Academy of Finland FIRI2015 - Infrarontier ESFRI Finland


Hans Kupczyk Guest Professorship Award, University of Ulm, Germany, 2015

Other scientific expert positions and activities

Memberships in scientific societies: Finnish Connective Tissue Research Society 1994- (Chair 2006-2007), International Society for Matrix Biology 1999- (Member of the Council 2006-2014), American Society for Biochemistry and Molecular Biology, 2002-

Selected activities in scientific meetings: >70 invited keynote lectures, other invited talks or organisational tasks since 1991

Memberships in editorial boards of scientific journals: Journal of Biological Chemistry - Member of the Editorial Board 2008-2013, Hypoxia - Member of the Honorary Editorial Board, 2013- 


Scientific merits

  • Professor, University of Oulu, Finland, 2007
  • Docent, University of Oulu, Finland, 2000
  • PhD, University of Turku, Finland, 1998

Professional and community activities

  • European Molecular Biology Conference (EMBC) council: Finnish delegate 2016-
  • European Molecular Biology Laboratory (EMBL) council: Finnish delegate 2016
  • Biocenter Finland: Member of the Board 2010- (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
  • Northern Finland Biobank Borealis: Member of the Scientific Committee 2016-, Member of the Steering Group 2012-2014

Research groups

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

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 18 PhD or MD-PhD theses to completion.


Hypoxia Response Pathway as a Target for Novel Therapeutics

Strategic research project of the University of OuluFocus institute: Biocenter Oulu