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Group Leader

M. Cristina Cardoso graduated in biology at the University of Lisbon, Portugal (1986). After finishing her doctoral thesis on phage molecular genetics (1990), she went on to her post-doctoral training in the group of Dr. B. Nadal-Ginard at the Harvard Medical School (Boston, USA) where she worked on inducing retro-differentiation in terminally differentiated cells. During this time, she started to explore the interplay between the mammalian cell nucleus landscape and (epi)genome metabolism. In 1995, she became a group leader at the Franz Volhard Clinic (Charité-Berlin, Germany) and from 1997 to 2008 at the Max Delbrück Center for Molecular Medicine (Berlin). Since 2008, she has established her group at the Technische Universität Darmstadt.

Link to publications page



Katrin Schallert is responsible to manage the secretariat of the work group. She is in charge for accounting and secretary matters. Accounting includes in particular the facts and figures of the research projects and in administrative matters. In the secretary, she is responsible among other things, to purchase lab and office material, doing the bills and taking care about personnel matters.



Dianca Imblan

Diana Imblan graduated in environment protection at the School of Applied Sciences in Bingen. She is working as a technical assistant in the Cardoso lab and takes care of several databases (Library of papers, Inventory lists, Peptides) and is supporting our PhD`s in molecular cloning.


Anne Lehmkuhl takes the best care of our cell cultures and is therefore absolutely not replaceable. She is all ears for special wishes and keeps the lab running with all the time freshly prepared solutions and sterilized lab materials.

Rausch, C.#, Weber, P.#, Prorok, P., Hörl, D., Maiser, A., Lehmkuhl, A., Chagin, V. O., Casas-Delucchi, C. S., Leonhardt, H. and Cardoso, M. C. (2020). Developmental differences in genome replication program and origin activation. Nucleic Acids Res. 48: 12751-12777.  

Ludwig, A. K.#, Zhang, P.#, Hastert, F. D., Meyer, S., Rausch, C., Herce, H. D., Muller, U., Lehmkuhl, A., Hellmann, I., Trummer, C., Storm, C., Leonhardt, H. and Cardoso, M. C. (2017). Binding of MBD proteins to DNA blocks Tet1 function thereby modulating transcriptional noise. Nucleic Acids Res 45: 2438-2457.

Zhang, P.#, Ludwig, A. K.#, Hastert, F. D., Rausch, C., Lehmkuhl, A., Hellmann, I., Smets, M., Leonhardt, H. and Cardoso, M. C. (2017). L1 retrotransposition is activated by Ten-eleven-translocation protein 1 and repressed by methyl-CpG binding proteins. Nucleus 8: 548-562.

Becker, A., Allmann, L., Hofstatter, M., Casa, V., Weber, P., Lehmkuhl, A., Herce, H. D. and Cardoso, M. C. (2013). Direct homo- and hetero-interactions of MeCP2 and MBD2. PLoS One 8: e53730.

Bertulat, B., De Bonis, M. L., Della Ragione, F., Lehmkuhl, A., Milden, M., Storm, C., Jost, K. L., Scala, S., Hendrich, B., D'esposito, M. and Cardoso, M. C. (2012). MeCP2 dependent heterochromatin reorganization during neural differentiation of a novel Mecp2-deficient embryonic stem cell reporter line. PLoS One 7: e47848.



Manuela Milden studied Biotechnology at the University of Applied Sciences Darmstadt, Germany. Her diploma thesis was finished in 2008 in the department of molecular biology. Now she takes care of the biochemistry lab, the antibodies and our Äkta system.

Bertulat, B., De Bonis, M. L., Della Ragione, F., Lehmkuhl, A., Milden, M., Storm, C., Jost, K. L., Scala, S., Hendrich, B., D'esposito, M. and Cardoso, M. C. (2012). MeCP2 dependent heterochromatin reorganization during neural differentiation of a novel Mecp2-deficient embryonic stem cell reporter line. PLoS One 7: e47848.

Jost, K. L., Rottach, A., Milden, M., Bertulat, B., Becker, A., Wolf, P., Sandoval, J., Petazzi, P., Huertas, D., Esteller, M., Kremmer, E., Leonhardt, H. and Cardoso, M. C. (2011). Generation and Characterization of Rat and Mouse Monoclonal Antibodies Specific for MeCP2 and Their Use in X-Inactivation Studies. PLoS One 6: e26499.






María C Arroyo Lopez (Postdoc) graduated in Biological Sciences, with a Master in Biotechnology and Biomedicine (2008-2013) and Molecular and Cellular Biology PhD at the University of Jaen, Spain (2014-2018). Her research work was focused in MCPH1 gene, functional analysis in chromosome condensation and cell cycle control (Human Genetics Department). She also worked during an internship at Dr. Duncan Clarke lab (Department of Genetics, Cell Biology and Development, University of Minnesota) investigating the functional interplay between TopoisomeraseII and MCPH1 regulating the G2 Decatenation Checkpoint. In July 2019 she joined Prof. Cristina Cardoso´s group to continue its scientific calling investigating epigenetic regulation. Specifically, she focuses on DNA modifications and the role of TET enzymes maintaining 5mC oxidation products. In her spare time, Maria enjoys climbing and drawing.

Duan, N., Arroyo, M., Deng, W., Cardoso, M. C. and Leonhardt, H. (2021). Visualization and characterization of RNA-protein interactions inliving cells. Nucleic Acids Res. 49: e107.

Arroyo, M., Sánchez, A., Cañuelo, A., Heredia-Molina, R. F., Martínez-Molina, E., Clarke, D. J., and Marchal, J. A. (2020). MCPH1 Lack of Function Enhances Mitotic Cell Sensitivity Caused by Catalytic Inhibitors of Topo II. Genes 11: 406.

Arroyo, M., Canuelo, A,. Calahorra, J., Hastert, F. D., Sanchez, A., Clarke, D. J. and Marchal, J. A. (2020). Mitotic entry upon Topo II catalytic inhibition is controlled by Chk1 and Plk1. FEBS J. 287: 4933.

Arroyo, M., Kuriyama, R., Guerrero, I., Keifenheim, D., Cañuelo, A., Calahorra, J., Marchal, J. A. (2019). MCPH1 is essential for cellular adaptation to the G 2 -phase decatenation checkpoint. The FASEB Journal 33: 8363-8374.

Arroyo, M., Kuriyama, R., Trimborn, M., Keifenheim, D., Cañuelo, A., Sánchez, A., Marchal, J. A. (2017). MCPH1, mutated in primary microcephaly, is required for efficient chromosome alignment during mitosis. Scientific Reports 7.

Arroyo, M., Trimborn, M., Sánchez, A., Hirano, T., Neitzel, H., & Marchal, J. A. (2015). Chromosome structure deficiencies in MCPH1 syndrome. Chromosoma 124: 491–501.


Ksenia Kolobynina joined Cardoso group as a PhD student in July 2017 and is currently focusing on discovering novel regulators of chromatin response to DNA damage with means of ubiquitinome wide screening and fluorescent microscopy. She studied biology at the Kazan Federal University, Russia, to obtain her bachelor degree with a focus on genetics in 2015. Meanwhile she got research experience in the lab of Prof. Dr. Barbara Schmidt in University of Regensburg, Germany, being a part of the project aiming to test HIV mimetic peptides. She did her master thesis at the Institute of Fundamental Medicine and Biology, Kazan, investigating potential role of human CHP3 protein in stem cell differentiation process under supervision of Prof. Albert Rizvanov.

Schwach, J., Kolobynina, K., Brandstetter, K., Gerlach, M., Ochtrop, P., Helma, J., Hackenberger, C. P. R., Harz, H., Cardoso, M. C. Leonhardt, H. and Stengl, A. (2021). Site-specific antibody fragment conjugates for reversible staining in fluorescence microscopy. ChemBioChem 22: 1205-1209.

Kolobynina, K. G., Solovyeva, V. V., Levay, K., Rizvanov, A. A. and Slepak, V. Z. (2016). Emerging roles of the single EF-hand Ca2+ sensor tescalcin in the regulation of gene expression, cell growth and differentiation. J Cell Sci 129: 3533-3540.


Maruthi Kumar Pabba started his bachelor from the Indian Institute of Technology Raipur, India, in 2013. He obtained his master's degree in biology from the Indian Institute of Technology Gandhinagar, India, in 2019. He joined the Cardoso group for his master thesis as a DAAD Exchange student in September 2018 where his research focus was to study the human Ubiquitin E3 Ligases and their role in DNA damage and repair. He joined the Cardoso lab as a Ph.D. to investigate the regulation of mammalian genome architecture and mobility. We will manipulate epigenetic states and subnuclear spatial localization using synthetic biology approaches. Subsequently, we will test how these impact genome architecture and mobility. His spare time activities involve traveling and painting.

Mamberti, S., Pabba, M. K., Rapp, A., Cardoso, M. C.# and Scholz, M.# (2022). The chromatin architectural protein CTCF is critical for cell survival upon irradiation-induced DNA damage. Int. J. Mol. Sci.  23: 3896.

# Corresponding authors    


Sunil Kumar Pradhan received his BS-MS dual degree in Biological Sciences from Indian Institute of Science Education and Research (IISER) Kolkata, India. His master thesis was focused on intraspecific competition of sibling colonies of Bacillus cereus. He also worked at the University of Zurich to study epigenetic regulated metabolism in brown adipose tissues. He started his Ph.D. in Cardoso lab to work on the (epi)genetic regulated replication progression and model of mammalian replication progression. In his spare time, he plays flute, goes trekking, and watches movies.


plays flute with the BiophoniX


Paulina Prorok (Postdoc) obtained her PhD in Biochemistry in 2013. Her thesis was consecrated to DNA repair mechanisms; she was specifically interested in describing novel and/or alternative repair pathways for highly mutagenic DNA lesions, like etheno-DNA adducts. Then she joined the laboratory of Dr Marcel Mechali at the Institute of Human Genetics in Montpellier, France. The DNA replication became her main scientific interest at that time and by using genome-wide approaches she was trying to depict factors determining the specification of DNA replication origins in metazoan cells. In 2019 she joined the laboratory of Prof. Cristina Cardoso at the TU Darmstadt, Germany to pursue her studies on molecular mechanisms regulating the DNA replication program as well as the response of the replisome when different types of DNA replication stress challenge the normal DNA replication progression.

Prorok, P., Grin, I. R., Matkarimov, B. T., Ishchenko, A. A., Laval, J., Zharkov, D. O., Saparbaev, M. (2021). Evolutionary Origins of DNA Repair Pathways: Role of Oxygen Catastrophe in the Emergence of DNA Glycosylases. Cells 10(7): 159.

Rausch, C.#, Weber, P.#, Prorok, P., Hörl, D., Maiser, A., Lehmkuhl, A., Chagin, V. O., Casas-Delucchi, C. S., Leonhardt, H. and Cardoso, M. C. (2020). Developmental differences in genome replication program and origin activation. Nucleic Acids Res. 48: 12751-12777.

Bazlekowa-Karaban, M.#, Prorok, P.#, Baconnais, S., Taipakova, S., Akishev, Z., Zembrzuska, D., Popov, A.V., Endutkin, A.V., Groisman, R., Ishchenko, A.A., Matkarimov, B.T., Bissenbaev, A., Le Cam, E., Zharkov, D.O., Tudek, B., Saparbaev, M. (2019). Mechanism of stimulation of DNA binding of the transcription factors by human apurinic/apyrimidinic endonuclease 1, APE1. DNA Repair (Amst) 82: 102698.

# First Authors

Prorok, P., Artufel, M., Aze, A., Coulombe, P., Peiffer, I., Lacroix, L., Guedin, A., Mergny, J.L., Damaschke, J., Schepers, A., Ballester, B. and Mechali, M. (2019). Involvement of G-quadruplex regions in mammalian replication origin activity. Nat Commun. 10: 3274.

Ganier, O.#, Prorok, P.#, Akerman, I. and Mechali, M. (2019). Metazoan DNA replication origins. Current Opinion in Cell Biology 58: 134-141.

# First Authors

Brustel, J., Kirstein, N., Izard, F., Grimaud, C., Prorok, P., Cayrou, C., Schotta, G., Abdelsamie, A.F., Dejardin, J., Mechali, M., Baldacci, G., Sardet, C., Cadoret, J.C., Schepers, A. and Julien, E. (2017). Histone H4K20 tri-methylation at late-firing origins ensures timely heterochromatin replication. EMBO J 36: 2726-2741.

Zdzalik, D.#, Domanska, A.#, Prorok, P.#, Kosicki, K., van den Born, E., Falnes, P., Rizzo, C.J.,  Guengerich, F.P. and Tudek, B. (2015). Differential repair of etheno-DNA adducts by bacterial and human AlkB proteins. DNA Repair 30: 1-10.

# First Authors

Joldybayeva, B.#, Prorok, P.#, Grin, I.R., Zharkov, D.O., Ishchenko, A.A., Tudek, B., Bissenbaev, A.K. and Saparbaev, M. (2014). Cloning and characterization of a Wheat homologue of apurinic/apyrimidinic endonuclease Ape1L. PLoS One 9: e92963.

# First Authors

Prorok, P., Alili, D., Saint-Pierre, Ch., Gasparutto, D., Zharkov, D., Ishchenko, A.A., Tudek, B. and Saparbaev, M. (2013). Uracil in duplex DNA is a substrate for the human nucleotide incision repair pathway. Proc Natl Acad Sci U S A 110: e3695-703.

Prorok, P., Saint-Pierre, Ch., Gasparutto, D., Fedorova, O.S., Ishchenko, A.A., Tudek, B. and Saparbaev, M.  (2012). Highly mutagenic exocyclic DNA adducts are substrates for the human nucleotide incision repair pathway. PLoS One 7: e51776.

Janowska, B., Kurpios-Piec, D., Prorok, P., Szparecki, G., Komisarski, M., Kowalczyk, P., Janion, C. and Tudek, B. (2012). Role of damage-specific DNA polymerases in M13 phage mutagenesis induced by a major lipid peroxidation product trans-4-hydroxy-2-nonenal. Mutat Res. 729: 41-51.

Janowska, B., Komisarski, M., Prorok, P., Sokolowska, B., Kusmierek, J.T., Janion, C. and Tudek, B. (2009). Nucleotide excision repair and recombination are engaged in repair of trans-4-hydroxy-2-nonenal adducts to DNA bases in Escherichia coli. Int.J.Biol.Sci. 5: 611-620.


Alexander Rapp (Postdoc) graduated in biochemistry at the Friedrich Schiller University in Jena, Germany, with a diploma thesis on assessing DNA damage after laser micro beams using the Comet-FISH assay.  During this time he got experience on different multi photon laser beams and their use for micromanipulation and damage induction.  After that he did his PhD in Molecular Biology at the Institute of Molecular Biotechnology (IMB) in Jena in the Group of K.O. Greulich (until 2004).  In 2005 he went to the Sir William Dunn School of Pathology, University of Oxford, UK as a Marie Curie Fellow in the lab of P. Cook.  During this time he studied the effects of radiation induced DNA damage in dependency on chromatin structure.  In 2008 he joined the Cardoso group at the TU Darmstadt for his habilitation in the field of DNA repair and chromatin.

Link to publications page


Hector Romero (Postdoc) graduated in biology in 2012 at the Complutense University of Madrid. During his master degree in biochemistry, obtained in Madrid in 2013, he studied the effect of HGF on the first steps of malignancy of oval cells treated with TGF-β. He did his PhD shared between the group of Juan Alonso at the National Centre of Biotechnology of Spain and Prof. Peter Graumann at Phillips-Universität Marburg, where he defend his thesis in 2018 based on the genetic and single-molecule characterization of two factors involved in DNA replication and repair in Bacillus subtilis. Since 2019, he is in Prof. Cristina Cardoso‘s group analyzing how chromatin compartimentalization is affecting protein dynamics and the factors involved in the process. In his spare time, Hector writes fantasy books and enjoys the nature together with his family.

Zhang, H. #, Romero, H. #, Schmidt, A., Gagova, K. Qin, W., Bertulat, B., Lehmkuhl, A., Milden, M., Eck, M., Meckel, T., Leonhardt, H. and Cardoso, M. C. (2022). MeCP2-induced heterochromatin organization is driven by oligomerization-based liquid-liquid phase separation and restricted by DNA methylation. Nucleus 13: 1-34

# First Authors

Romero, H., Serrano, E., Hernández-Tamayo, R., Carrasco, B., Cárdenas, PP., Ayora, S., Graumann, PL., Alonso, JC. (2020) Bacillus subtilis RarA Acts as a Positive RecA Accessory Protein. Front Microbiol. Feb 13 :11:92.

Romero, H., Torres, R., Hernandez-Tamayo, R., Carrasco, B., Ayora, S., Graumann, P. L., Alonso, J.C. (2019) Bacillus subtilis RarA acts at the interplay between replication and repair-by-recombination. DNA repair 78: 27-36. 

Torres, R., Romero, H., Rodriguez-Cerrato, V. and Alonso, J.C. (2017) Interplay between Bacillus subtilis RecD2 and the RecG and RuvAB helicase in recombinational repair. DNA repair 55: 44-46.

Romero, H., Rösch, T., Hernandez-Tamayo, R., Lucena, D., Ayora, S., Alonso, J.C. and Graumann, P. (2019) Single molecule tracking reveals function of RarA at replication forks but also independently from replication during DNA repair in Bacillus subtilis. Sci Reports 9: 1997.


Andreas Zhadan began his studies with Biology at the Technical University of Darmstadt in 2016, and obtained his Master degree in 2021. He joined the Cardoso Lab for his bachelor thesis, where he researched the influence of solar radiation on cells. For his Master thesis, he investigated the regulatory role of the zinc finger domain of Tet1 (Ten-eleven translocation protein 1), as well as the regulation of the cancer-related short isoform of Tet1 throughout the cell cycle. In 2021, he started his PhD at the Cardoso Lab, where he will be investigating the regulation of DNA modifications and modifiers, mainly focusing on the role of Tet1.





Hui Zhang graduated in Biology at Harbin Institute of Technology (HIT) in China as a Master student in 2017. His Master Thesis involved the functions of Marveld1 (MARVEL domain containing protein 1) during mouse placenta development and underlying molecular mechanisms. In 2017 he joined the Cardoso Lab for his PhD. Here he will focus on the underlying mechanism of Mecp2 (methyl-CpG binding protein 2) multiple functions during neural differentiation, highlighting the emerging hypothesis underlying various membraneless organelles within cells and particularly nucleus. More recently, the work expands to the MBD protein family.

Zhang, H. #, Romero, H. #, Schmidt, A., Gagova, K. Qin, W., Bertulat, B., Lehmkuhl, A., Milden, M., Eck, M., Meckel, T., Leonhardt, H. and Cardoso, M. C. (2022). MeCP2-induced heterochromatin organization is driven by oligomerization-based liquid-liquid phase separation and restricted by DNA methylation. Nucleus 13: 1-34

# First Authors

Schmidt, A.#, Zhang, H.# and Cardoso, M. C. (2020). MeCP2 and chromatin compartmentalization. Cells 9: 878.

Chen, Y., Zhang, H., Han, F., Yue, L., Qiao, C., Zhang, Y., Dou, P., Liu, W. and Li, Y. (2018). The depletion of MARVELD2 leads to murine placenta accreta via integrin β4-dependent trophoblast cell invasion. J. Cell. Physiol 233: 2257-2269.



Simon Berg is a Biology student supporting Cardoso Lab during the first two semesters of their Masters Degree. After finishing their bachelor thesis at Cardoso Lab about single molecule tracking of histone dynamics in living cells, they continued working in the area of single molecule / particle tracking to study the dynamics of chromatin and proteins involved in DNA replication. They also enjoy providing assistance to the lab through programming and bioinformatics, and are continually trying to widen their experience in anticipation of an eventual PhD. In their free time they enjoy creating things, be it through digital art, tinkering or carpentry. As a transgender person currently in the process of transitioning, they prefer being referred to as Veronika but are also fine being called by their legal name in professional settings.


Cardoso Lab  •  Technische Universität Darmstadt  •  Germany


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