Tine Engberg Thingholm

Ph.d. i Biokemi og Molekylær Biologi

20052019
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Personlig profil

CV

Personal Data:Name: Tine Engberg Thingholm Nationality: Danish Date of Birth: 13th of February 1974 Address: Sanderumvej 166A, 5250 Odense SV Phone: + 45 28 25 12 77 E-mail: tthingholm@health.sdu.dk ot tine@thingholm.com

Academic positions:Postdoctoral Fellow at The Molecular Endocrinology Unit (KMEB), Odense University Hospital (OUH), Denmark 2 years of funding from Research Foundation of Odense University Hospital (from October 1st 2008)Research Assistant at The Molecular Endocrinology Unit (KMEB), Odense University Hospital (OUH), Denmark Funded by Professor Henning Beck-Nielsen (from June 1st 2008)Academic Education: Ph. D. in Biochemistry and Molecular Biology “Phosphoproteomic analysis of human cells by mass spectrometry- setting up strategies”. University of Southern Denmark, Odense, Denmark (Examination on September 11th 2008)M.Sc. Degree Molecular and Cell Biology “Identification of phosphoproteins from plasma membranes of the human TERT20 stem cells using mass spectrometric techniques”. University of Southern Denmark and Odense University Hospital, Denmark. Grade: 11, A (Examination on May 18th 2005)Student Project in the Biomedical Research and Molecular Cell Biology Group “The role of Protein Kinase CK2 in the stability and activity of the Tumor Suppressor gene, PTEN”. University of Southern Denmark (September 2002 - June 2003)B.Sc. Molecular and Cell Biology “Cloning, purification, expression and characterization of recombinant human protein MAPK1/ERK1”. University of Southern Denmark Grade: 13, A+ (Evaluation in November 2001)Awards/Grants:

HUPO Young Investigator Award, 2009 Given at the HUPO 8th Annual World Congress, Sep 26-30 2009, Toronto, Canada Mogens Rendals Grant, 2009 Received in May 2009

EuPA Young Investigator Prize, 2008 Given at the HUPO 7th Annual World Congress, Aug 18-20 2008, Amsterdam, Holland HUPO Best Poster Award Given at the HUPO 6th Annual World Congress, Oct 6-10 2007, Seoul, South Korea European Calcified Tissue Society Young Investigator Award Given at the 34th European Symposium on Calcified Tissues, May 5-9 2007, Copenhagen, Denmark HUPO Poster Award, First prize Given at the HUPO 5th Annual World Congress, Oct 28-Nov 1 2006, Long Beach, California, USA Experience in Peer Reviewing:Frequently reviewing manuscripts for editors of journals such as Proteomics, Journal of Proteome Research, Electrophoresis and Expert Review of Proteomics Courses attended during Ph.D.:Practical course in communication and presentation (at SDU, 3 ECTS points) PhD Summer School 2006 in Functional Genomics and Proteomics as Applied to Metabolic Diseases (Nyborg, 3 ECTS points) Mass Spectrometry Applications in Protein Research (at Utrecht University, Holland, 1.5 ECTS points) PhD Summer School 2007 in Functional Genomics and Proteomics as Applied to Metabolic Diseases (Gl. Avernæs, 3 ECTS points) BM/BMP59 Differentiation and Cancer (at SDU, 9 ECTS points) Proteomics Basic (Opatija, Croatia, 4 ECTS points) PhD Summer School 2008 in Functional Genomics and Proteomics as Applied to Metabolic Diseases (Gl. Avernæs, 1 ECTS point) Biological Sequence Analysis (at DTU, 10 ECTS points) BM/BMP31 Advanced methods in Protein Chemistry (at SDU, 3 ECTS points) Research & Work Experience:Member of Protein Research Group, University of Southern Denmark (June 2003 to present)• Membrane protein purification • Fractionation of complex samples (gel electrophoresis, SAX, SCX) • Enrichments for phosphopeptides (IMAC, TiO2, SIMAC) • Quantitation (iTRAQ, 4plex/8plex) • Mass Spectrometry; MALDI- and LC-MS/MS • Bioinformatics

Member of Clinic for Molecular Endocrinology Treatment, Odense University Hospital (June 2003 to present) • Cell Culture (human cell lines) • Stem cell: Osteoblast Differentiation • Stem Cell: Treatment with Phosphatase inhibitors and Analysis • Human myotubes: Insulin stimulation and analysis Member of Children´s Medical Research Institute (CMRI), Westmead, Australia (November 2007 to February 2008)• Expression of GST-constructs in E.coli • HeLa Cells: Cell Culture and Synchronization • Pull downs in HeLa Cells and Rat tissues using GST constructs • Cell Culture (HeLa cells and bacteria) Kinase Detect A/S (August 2001 - May 2002)Research Assistant Biomedical Research and Molecular Cell Biology Group, University of Southern Denmark (February 2001 – June 2003)• RNA, DNA and protein purification • Primer design and RT-PCR • Cloning (transformation and expression) • Western Blotting • Activity test (using 32P-ATP) • DNA Sequencing • Transfection • Cell Culture (bacteria) Extracurricular Activities: Tour guide for new students at the Faculty of Science (August 2003) Study guidance for new members of the Dept. of Biochemistry and Molecular Biology (BMB) (August 2003 to January 2004)Member of the Society of Cell Biology (August 1997 – present) List of publications:“Analytical strategies for phosphoproteomics” Tine E. Thingholm, Ole N. Jensen, Martin R. Larsen Proteomics. 2009 Feb 20. [Epub aheah of print] PMID: 19235172 [PubMed - indexed for MEDLINE] “TiO2-based phosphoproteomic analysis of the plasma membrane and the effects of phosphatase inhibitor treatment” Tine E. Thingholm; Martin R. Larsen; Christian R. Ingrell; Moustapha Kassem and Ole N. Jensen J Proteome Res. 2008 Aug 1;7(8):3304-3313. Epub 2008 Jun 26. PMID: 18578522 [PubMed - indexed for MEDLINE] “SIMAC – A phosphoproteomic strategy for the rapid separation of mono-phosphorylated from multiply phosphorylated peptides” Tine E. Thingholm, Ole N. Jensen, Phillip J. Robinson and Martin R. Larsen Mol Cell Proteomics. 2008 Apr;7(4):661-71. Epub 2007 Nov 26. PMID: 18039691 [PubMed - indexed for MEDLINE] “Highly selective enrichment of phosphorylated peptides using titanium dioxide” Tine E. Thingholm, Thomas J. D. Jørgensen, Ole N. Jensen and Martin R. Larsen Nat Protoc. 2006;1(4):1929-35. PMID: 17487178 [PubMed - indexed for MEDLINE] “Analysis of posttranslational modifications of proteins by tandem mass spectrometry” Martin R. Larsen, Morten B. Trelle, Tine E. Thingholm, Ole N. Jensen Biotechniques. 2006 Jun;40(6):790-8. Review. PMID: 16774123 [PubMed - indexed for MEDLINE] “Highly selective enrichment of phosphorylated peptides from complex mixtures using titanium dioxide microcolumns” Martin R. Larsen, Tine E. Thingholm, Ole N. Jensen, Peter Roepstorff and Thomas J.D. Jørgensen Mol Cell Proteomics. 2005 Jul;4(7):873-86. Epub 2005 Apr 27. PMID: 15858219 [PubMed - indexed for MEDLINE] “Enrichment and Characterization of Phosphopeptides by Immobilized Metal Affinity Chromatography (IMAC) and Mass Spectrometry” Tine E. Thingholm and Ole N. Jensen Methods Mol Biol. 2009;527:47-56 PMID: 19241004 [PubMed - indexed for MEDLINE] “The use of titanium dioxide micro-columns to selectively isolate phosphopeptides from proteolytic digests” Tine E. Thingholm and Martin R. Larsen Methods Mol Biol. 2009;527:57-66 PMID: 19241005 [PubMed - indexed for MEDLINE] “Enrichment and separation of mono- and multi-phosphorylated peptides using Sequential elution from IMAC prior to mass spectrometric analysis” Tine E. Thingholm, Ole N. Jensen and Martin R. Larsen Methods Mol Biol. 2009;527:67-78 PMID: 19241006 [PubMed - indexed for MEDLINE] Technologies for plasma membrane proteomics” Stuart J. Cordwell and Tine E. Thingholm Proteomics, 2009 Oct 15. PMID: 19834916 [PubMed - as supplied by publisher] “Methods and strategies for Quantitation in Phosphoproteomics” Tine E. Thingholm Submitted to Expert Review of Proteomics on Nov. 16th 2009 Manuscripts in preparation:“Characterization of the Type 2 Diabetic proteome baseline using quantitative proteomics” Tine E. Thingholm and Michael Gaster In preparation, for submission in December 2009 This manuscript describes a quantitative proteomic study of human myotubes in order to characterize differences at the proteome baseline between myotubes established from type 2 diabetic subjects and two healthy control groups (healthy lean subjects and healthy obese subjects). The data shows clear differences between the three groups. Specific groups of proteins are clearly regulated when comparing the diabetic myotubes to healthy myotubes, but also myotubes established from obese subjects (both diabetic and healthy obese subjects) show distinc changes when compared to myotubes established from healthy lean subjects. The data has been prepared, the manuscript is in preparation. “Proteomic changes in Type 2 Diabetic myotubes upon insulin stimulation” Tine E. Thingholm and Michael Gaster In preparation This manuscript characterizes the protein changes occurring as a result of insulin stimulation. Two comprehensive experiments were performed in order to compare the protein profiles of myotubes from healthy obese, healthy lean and obese T2D patients during insulin stimulation for 0, 30 or 60 minutes. The data has been prepared, the manuscript is in preparation. “Phosphoproteomic study of Type 2 Diabetic myotubes using iTRAQ and SIMAC” Tine E. Thingholm and Michael Gaster In preparation, October 2009 This manuscript describes a quantitative phosphoproteomic study of human myotubes in order to identify the mechanisms causing insulin resistance in Type 2 Diabetes patients. As several proteins are expected to be involved in insulin resistance, it is important also to obtain an overall picture of the phosphorylation taking place during insulin stimulation in a time-dependent manner. Four comprehensive experiments were performed in order to compare the phosphorylation profiles of myotubes from obese, lean and obese T2D patients during insulin stimulation at six time points. The phosphorylation profiles were characterized and compared using quantitative phosphoproteomics. The data has been prepared. We are now in the process of validating the data. Presentations at scientific meetings:I have been very fortunate as to get the opportunity to present my work at various international (and national) conferences. As a Ph.D. student you have limited funding for attending international conferences. However, at the beginning of my Ph.D., I presented some of my work in which I had been using the ProteinCenter program from Proxeon at the ASMS conference. After this I was contacted by Proxeon, who offered to sponsor my attendance at all the conferences in which I were giving oral presentations. As I have been appointed oral presentations at many of the subsequent conferences and also have been an invited speaker at many of the conferences/meetings, I have given 28 oral presentations at international scientific meetings (incl. 15 lectures as invited speaker, 1 plenary lecture and 1 presentation presented by a co-author). In addition, I have 10 poster presentations at international scientific meetings (incl. 4 posters presented by co-authors). Invited speaker at:Scientific Meeting: TGen, The Translational Genomics Research Institute, Phoenix, Arizona Title of the talk: “Phosphoproteomics – Method development and applications” Date: November 20th, 2009 Scientific Meeting: Thermo: Nordic High Resolution tour, Gothenburg, Sweden Title of the talk: “Proteomic and Phosphoproteomic characterization of T2D using iTRAQ and LTQ-orbitrap XL/Velos” Date: November 12th , 2009 Scientific Meeting: Thermo: Nordic High Resolution tour, Lund, Sweden Title of the talk: “Proteomic and Phosphoproteomic characterization of T2D using iTRAQ and LTQ-orbitrap XL/Velos” Date: November 11th, 2009 Scientific Meeting: Diabetes Meeting, University Hospital, Odense, Denmark Title of the talk: “Quantitative phosphoproteomics - Studying insulin resistance in T2D patients” Date: December 15th, 2008 Scientific Meeting: The 9th London Biological Mass Spectrometry Discussion Group (LBMSDG) Meeting, London, England Title of the talk: Is total phosphoproteome analysis obtainable? (Plenary Lecture) Date: December 11th, 2008 Scientific Meeting: LTQ Orbetrap User Meeting, Odense, Denmark Title of the talk: “Proteome Discoverer in action - Quantitative phosphoproteomics in diabetes research” Date: December 3rd, 2008 Scientific Meeting: The 4th Symposium of the Danish Proteomics Society (DAPSOC), Odense, Denmark Title of the talk: “Quantitative phosphoproteomics in diabetes research” Date: December 2th, 2008 Scientific Meeting: The 2nd European Summer School "Proteomic Basics" 2008, Brixen, Italy Title of the talk: “Looking into the proteomics toolbox” Date: July 13th-19th, 2008 Scientific Meeting: Pre-Lorne Symposium, Melbourne, Australia Title of the talk: “Interpretation and validation of MS/MS data” Date: February 6th, 2008 Scientific Meeting: Pre-Lorne Symposium, Melbourne, Australia Title of the talk: “Phosphoproteomic studies of plasma membranes in human mesenchymal stem cells” Date: February 6th, 2008 Scientific Meeting: Educational Session at the 6th Annual Human Proteome Organisation (HUPO) World Congress, Seoul, South Korea Title of the talk: “Protein Identification by MS and MS/MS” Date: October 6-10, 2007 Scientific Meeting: Invited to give a Phosphoproteomic seminar at Plataforma de Proteòmica, University of Barcelona, Barcelona, Spain Title of the talk: “Functional Phosphoproteomics” Date: June, 2007 Scientific Meeting: The Dionex Meeting “MS Applications and Instruments”, Odense, Denmark Title of the talk: “Functional phosphoproteomics on plasma membranes from human stem cells” Date: January, 2007 Scientific Meeting: Educational Session at the 5th Annual Human Proteome Organisation (HUPO) World Congress, Long Beach, USA Title of the talk: “Peptide Mass Fingerprinting” and “Protein Identification with MALDI and LC-ESI MS/MS” Date: October-November 2006 Scientific Meeting: The University of Newcastle, Newcastle, Australia Title of the talk: “Identification of plasma membrane phosphoproteins from human mesenchymal stem cells using titanium dioxide (TiO2) purification and mass spectrometry” Date: September 2005 Other oral presentations:Scientific Meeting: The 7th Annual Human Proteome Organisation (HUPO) World Congress, Amsterdam, Holland Title of the talk: “Dlk1 function in stem cells - studied by quantitative phosphoproteomics by iTRAQ and SIMAC/TiO2” Date: August 16-20, 2008 Award: EuPA Young Investigator Award Scientific Meeting: The 13th Annual Proteomics Symposium, Lorne, Australia Title of the talk: “SIMAC - A phosphoproteomic strategy for Separation of mono- from multi-phosphorylated peptides” Date: February 7-10, 2008 Scientific Meeting: The 55th American Society for Mass Spectrometry (ASMS) Conference, Indianapolis, IN, USA Title of the talk: “A new phosphoproteomic strategy - Separation of mono- from multi-phosphorylated peptides” Date: June 3-7 2007 Scientific Meeting: The 34th European Symposium on Calcified Tissues (ECTS), Copenhagen, Denmark Title of the talk: “Identification of novel Phosphoproteins on Plasma Membranes of Human Mesenchymal Stem Cells using TiO2 and Mass spectrometry” Date: May 5-9 2007 Award: ECTS Young Investigator Award Scientific Meeting: The European Proteomics Association (EuPA) Conference, Valencia, Spain Title of the talk: “Phosphoproteomics on plasma membranes from human stem cells using titanium dioxide chromatography and mass spectrometry” Date: February 2007 Scientific Meeting: Nordic Network on Proteomics Meeting (Discussion Group) – “Quantitation MS”, Odense, Denmark Title of the talk: “Phosphoproteomics on human mesenchymal stem cells” Date: November 27-28th, 2006 Scientific Meeting: The PhD Summer School 2006, Functional Genomics and Proteomics as Applied to Metabolic Diseases, Gl. Avernæs, Denmark Title of the talk: “Important Tools in Phosphoproteomics” Date: September 2006 Scientific Meeting: The University of Utrecht, Utrecht, Holland Title of the talk: “Studying Phosphorylation in Stem Cells by TiO2 purification and Mass Spectrometry” Date: April 2006 Scientific Meeting: The Federal University of Rio de Janeiro, Rio de Janeiro, Brasil Title of the talk: “Enrichment and identification of phosphopeptides from human stem cells using IMAC/TiO2 and multistage Mass Spectrometry” Date: February 6th, 2006 Scientific Meeting: The PhD Summer School 2005, Functional Genomics and Proteomics as Applied to Metabolic Diseases, Nyborg, Denmark Title of the talk: “Identification of plasma membrane phosphoproteins from human stem cells using titanium dioxide (TiO2) purification and mass spectrometry” Date: September 2005 Scientific Meeting: Dansk Infektions Medicinsk Workshop, Kertemide, Danmark Title of the talk: “BILF1 regulation of signal transducers identified by phosphoproteomics” Co-author: R. Lyngaa, S. J. Paulsen, T. E. Thingholm, B. S. Andresen, M. M. Rosenkilde, O. N. Jensen and T. N. Kledal Date: September 2005 Scientific Meeting: The University Hospital of Odense, Odense, Denmark Title of the talk: “Multidimensional Separation Technology (SAX-IMAC) and Mass Spectrometry for Identification of Membrane Phosphoproteins” Date: June 2005 Scientific Meeting: The 2nd Italian Proteome Society (Ipso) Congress, Viterbo, Italy Title of the talk: “Titanium dioxide and Mass Spectrometry for Identification of Plasma Membrane Phosphoproteins from Human Stem Cells” Date: May 2005 Poster presentations:Scientific Meeting: The 8th Annual Human Proteome Organisation (HUPO) World Congress, Toronto, Canada Title of the talk: “Proteomic and Phosphoproteomic characterization of T2D using iTRAQ and LTQ-orbitrap XL/Velos” Date: September - October 2009 Award: HUPO Young Investigator Award Scientific Meeting: The 57th American Society for Mass Spectrometry (ASMS) Conference, Philadelphia, PA, USA Title of the talk: “Phosphoproteomic characterization of insulin resistance in T2D using iTRAQ and SIMAC combined with Multistage Activation and Higher-energy Collision Dissociation MS/MS” Date: May 31st –June 4th 2009 Scientific Meeting: The 31st International Herpesvirus Workshop, Seattle, USA Title of the talk: “Viral G protein coupled receptors induces phosphorylation of the microtubule destabilizing oncoprotein OP18/stathmin” Co-author: R. Lyngaa, T. Carlsen, T. E. Thingholm, O. N. Jensen and T. N. Kledal Date: July 2006 Scientific Meeting: The 54th American Society for Mass Spectrometry (ASMS) Conference, Seattle, WA, USA Title of the talk: “Phosphoproteome analysis of human mesenchymal stem cells using TiO2 columns and mass spectrometry” Date: May-June 2006 Scientific Meeting: Educational Session at the 4th Annual Human Proteome Organisation (HUPO) World Congress, Munich, Germany Title of the talk: “Highly selective purification of phosphorylated peptides using Titanium Dioxide microcolumns” Co-author: M. R. Larsen, T. E. Thingholm, O. N. Jensen, P. Roepstorff and T. J. D. Jørgensen Date: August 2005 Award: Best Poster Award, HUPO 2005 Scientific Meeting: The 30th International Herpesvirus Workshop, Turku, Finland Title of the talk: “BILF1 regulation of signal transducers identified by phosphoproteomics” Co-author: R. Lyngaa, S. J. Paulsen, T. E. Thingholm, B. S. Andresen, M. M. Rosenkilde, O. N. Jensen and T. N. Kledal Date: July-Aug 2005 Scientific Meeting: The Danish Cancer Research Symposium, Copenhagen, Denmark Title of the talk: “Multidimensional Separation Technology(SAX-IMAC) and Mass Spectrometry for Identification of Membrane Phosphoproteins from Human Stem Cells” Date: August 2004

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  • 54 Lignende profiler
Phosphopeptides Medicin og biovidenskab
Peptides Medicin og biovidenskab
Phosphorylation Kemiske forbindelser
Proteins Medicin og biovidenskab
Affinity Chromatography Medicin og biovidenskab
Mass spectrometry Kemiske forbindelser
Phosphoproteins Medicin og biovidenskab
Metals Medicin og biovidenskab

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Publikationer 2005 2018

72 Downloads (Pure)

XK-related protein 5 (XKR5) is a novel negative regulator of KIT/D816V-mediated transformation

Sun, J., Thingholm, T., Højrup, P. & Rönnstrand, L., 18. jun. 2018, I : Oncogenesis. 7, 6, 11 s., 48.

Publikation: Bidrag til tidsskriftTidsskriftartikelForskningpeer review

Åben adgang
Fil
Proteins
Tyrosine
Down-Regulation

Phosphopeptide enrichment by immobilized metal affinity chromatography

Thingholm, T. E. & Larsen, M. R., 2016, Phospho-Proteomics: Methods and Protocols. von Stechow, L. (red.). 2. udg. Humana Press, s. 123-133 (Methods in Molecular Biology, Bind 1355).

Publikation: Bidrag til bog/antologi/rapport/konference-proceedingBidrag til bog/antologiForskningpeer review

Phosphopeptides
Affinity Chromatography
Metals
Peptides
Ions

Characterization of human myotubes from type 2 diabetic and non-diabetic subjects using complementary quantitative mass spectrometric methods

Thingholm, T. E., Bak, S., Beck-Nielsen, H., Jensen, O. N. & Gaster, M., 1. sep. 2011, I : Molecular and Cellular Proteomics. 10, 9, s. M110.006650

Publikation: Bidrag til tidsskriftTidsskriftartikelForskningpeer review

Muscle
Skeletal Muscle
Proteins
Adenosine Deaminase
Medical problems

Quantitative iTRAQ-Based Proteomic Identification of Candidate Biomarkers for Diabetic Nephropathy in Plasma of Type 1 Diabetic Patients

Overgaard, A. J., Thingholm, T. E., Larsen, M. R., Tarnow, L., Rossing, P., McGuire, J. N. & Pociot, F. M., 1. dec. 2010, I : Clinical Proteomics. 6, 4, s. 105-114 10 s.

Publikation: Bidrag til tidsskriftTidsskriftartikelForskningpeer review

Diabetic Nephropathies
Biomarkers
Apolipoprotein A-II
Plasmas
Immunoassay

Strategies for quantitation of phosphoproteomic data

Palmisano, G. & Thingholm, T. E., 1. jun. 2010, I : Expert Review of Proteomics. 7, 3, s. 439-56 18 s.

Publikation: Bidrag til tidsskriftTidsskriftartikelForskningpeer review

Phosphorylation
Phosphoproteins
Chemical modification
Labeling
Mass spectrometry

Aktiviteter 2005 2010

  • 28 Foredrag og præsentationer i privat eller offentlig virksomhed
  • 1 Organisering af eller deltagelse i konference
  • 1 Gæsteophold ved andre institutioner

“Quantitative characterization of human myotubes in type 2 diabetes - Selective Reaction Monitoring for validation of iTRAQ data”

Tine Engberg Thingholm (Foredragsholder)
21. sep. 2010

Aktivitet: Foredrag og mundtlige bidragForedrag og præsentationer i privat eller offentlig virksomhed

“Sequential elution from IMAC (SIMAC) - Separation of mono- and multi-phosphorylated peptides”

Tine Engberg Thingholm (Foredragsholder)
19. sep. 2010

Aktivitet: Foredrag og mundtlige bidragForedrag og præsentationer i privat eller offentlig virksomhed

“Quantitative characterization of a biological system - considerations and challenges”

Tine Engberg Thingholm (Foredragsholder)
19. apr. 2010

Aktivitet: Foredrag og mundtlige bidragForedrag og præsentationer i privat eller offentlig virksomhed

"Phosphoproteomics - Method development and applications"

Tine Engberg Thingholm (Foredragsholder)
20. nov. 2009

Aktivitet: Foredrag og mundtlige bidragForedrag og præsentationer i privat eller offentlig virksomhed

"Proteomic and Phosphoproteomic characterization of T2D using iTRAQ and LTQ-orbitrap XL/Velos"

Tine Engberg Thingholm (Foredragsholder)
12. nov. 2009

Aktivitet: Foredrag og mundtlige bidragForedrag og præsentationer i privat eller offentlig virksomhed

Projekter 2015 2019

Priser

Mogens Rendals Grant

Tine Engberg Thingholm (Modtager), maj 2011

Pris: Priser, stipendier, udnævnelser

“Young Gun” Early Career Investigator Award, 2010

Tine Engberg Thingholm (Modtager), 22. sep. 2010

Pris: Priser, stipendier, udnævnelser

EuPA Young Investigator Prize, 2008

Tine Engberg Thingholm (Modtager), 20. aug. 2008

Pris: Priser, stipendier, udnævnelser