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Publications AG Doehlemann


Depotter JRD, Misas Villamil JC, Doehlemann G (2022) Maize immune signaling peptide ZIP1 evolved de novo from a retrotransposon. bioRxiv 2022.05.18.492421; doi:


2023 - 2024

Werner J, Zuo W, Doehlemann G (2024) CRISPR/Cas9 Ribonucleoprotein-Mediated Mutagenesis in Sporisorium reilianum. Bio-protocol 14(8): e4978.

Yuan W, Chen X, Du K, Jiang T, Li M, Cao Y, Li X, Doehlemann G, Fan Z, Zhou T (2024) NIa-Pro of sugarcane mosaic virus targets Corn Cysteine Protease 1 (CCP1) to undermine salicylic acid-mediated defense in maize. PloS Pathogens 14;20(3):e1012086.

Huang L, Ökmen B, Stolze SC, Kastl M, Khan M, Hilbig D, Nakagami H, Djamei A, Doehlemann G (2024) The fungal pathogen Ustilago maydis targets the maize corepressor TPL2 to modulate host transcription for tumorigenesis. New Phytologist 241(4):1747-1762.

Zuo W, Depotter JRL, Stolze SC, Nakagami H, Doehlemann G (2023)  A transcriptional activator effector of Ustilago maydis regulates hyperplasia in maize during pathogen-induced tumor formation. Nature Communications 23;14(1):6722 

Ökmen B, Katzy P, Huang L, Wemhöner R, Doehlemann G (2023) A conserved extracellular ribonuclease with broad-spectrum cytotoxic activity enables smut fungi to compete with host-associated bacteria. New Phytologist Sep 7.doi: 10.1111/nph.19244.; preprint available @ bioRxiv 2023.04.19.537453; doi:

Barghahn S, Saridis G, Mantz M, Meyer U, Mellüh JC, Misas Villamil JC, Huesgen PF, Doehlemann G (2023) Combination of transcriptomic, proteomic and degradomic profiling reveals common and distinct patterns of pathogen-induced cell death in maize. Plant Journal 2023 Jun 20. doi: 10.1111/tpj.16356.

Shi W, Stolze SC, Nakagami H, Misas Villamil JC, Saur IML, Doehlemann G (2023) Combination of in-vivo proximity labeling and co-immunoprecipitation identifies the host target network of a tumor-inducing effector in the fungal maize pathogen Ustilago maydis. Journal of Experimental Botany 2023 May 24;erad188.; doi: 10.1093/jxb/erad188.

Koenig M, Moser D, Leusner J, Depotter J, Doehlemann G, Misas Villamil JC (2023) Maize phytocytokines and microbial-patterns trigger antagonistic features in co-incidence with wounding and fungal pathogens. Mol Plant Microbe Interact. 27 Apr 2023.

Djamei A, Depotter J, Saridis G, Prokchorchik M, Barghahn S, de Sousa Teixeira E. Silva N, Zuo W, Misas Villamil J, Doehlemann G (2023) Modulation of Host Immunity and Development by Ustilago maydis. In: Scott, B., Mesarich, C. (eds) Plant Relationships. The Mycota, vol 5. Springer, Cham.


2021 -2022

Ökmen B, Jaeger E, Schilling L, Finke N, Lee YJ, Wemhöner R, Pauly M, Neumann U, Doehlemann G (2022) A conserved enzyme of smut fungi facilitates cell-to-cell extension in the plant bundle sheath. Nature Communications 12;13(1):6003.

Depotter JRL, Ökmen B, Ebert MK, Beckers J, Kruse J, Thines M, Doehlemann G (2022) High nucleotide substitution rates associated with retrotransposon proliferation drive dynamic secretome evolution in smut pathogens. Microbiology Spectrum 10(5):e0034922.

Bindics J, Khan M, Uhse S, Kogelmann B, Baggely L, Reumann D, Ingole KD, Stirnberg A, Rybecky A, Darino M, Navarrete F, Doehlemann G, Djamei A (2022) Many ways to TOPLESS – manipulation of plant auxin signalling by a cluster of fungal effectors. New Phytologist 6(4):1455-1470.

Passarge A, Doehlemann G, Misas Villamil JC (2022) Detection of apoplastic protease inhibitors using convolution activity-based protein profiling.  Methods Mol Biol. 2447:95-104.

Bradley EL, Ökmen B, Doehlemann G, Henrissat B, Bradshaw R, Mesarich CH. (2022) Secreted glycoside hydrolase (GH) proteins as effectors and invasion patterns of plant-associated fungi and oomycetes.  Front Plant Sci.13:853106

Zuo W, Gupta DK, Depotter JRL, Thines M, Doehlemann G. (2021) Cross-species analysis between the maize smut fungi Ustilago maydis and Sporisorium reilianum highlights the role of transcriptional plasticity of effector orthologs for virulence and disease. New Phytologist 232(2):719-733

Passarge A, Demir F, Green J, Depotter JRL, Scott B, Huesgen PF, Doehlemann G, Misas Villamil J. (2021) Host apoplastic cysteine protease activity is supressed during the mutualistic association of Lolium perenne and Epichloë festucae. Journal of Experimental Botany [in press]

Schurack S, Depotter JRL, Gupta M, Thines M, Doehlemann G. (2021) Transcriptome analysis in the maize – Ustilago maydis interaction identifies maize-line specific activity of fungal effectors. Plant J DOI: 10.1111/tpj.15195

Eitzen K, Sengupta P, Kroll S, Kemen E, Doehlemann G. (2021) A fungal member of the Arabidopsis thaliana phyllosphere antagonizes Albugo laibachii via a GH25 lysozyme. eLife 10:e65306

Ökmen B, Schwammbach D, Bakkeren G, Neumann U, Doehlemann G. (2021) The Ustilago hordei-barley interaction is a versatile system to characterize fungal effectors. Journal of Fungi 27;7(2):86

Chaudry V, Runge P, Sengupta P, Doehlemann G, Parker JE, Kemen E. (2021) Shaping the leaf microbiota: plant-microbe-microbe interactions. Journal of Experimental Botany 20;72(1):36-56.


2019 – 2020

Depotter JRL, Zuo W, Hansen M, Zhang B, Xu M, Doehlemann G. (2020) Effectors with different gears: divergence of Ustilago maydis effector genes is associated with their temporal expression pattern during plant infection. Journal of Fungi 7(1), 16

Zuo W, Depotter JRL, Doehlemann G. (2020) Cas9HF1 enhanced specificity in Ustilago maydis. Fungal Biology 124 (3-4) 228-234

Depotter JRL, Doehlemann G (2020) Target the core: durable plant resistance against filamentous plant pathogens through effector recognition. Pest Man Sci 76 (2) 426-431

Zuo W, Ökmen B, Depotter JRL, Ebert MK, Redkar A, Misas Villamil JM, Doehlemann G. (2019) Molecular interactions between smut fungi and their host plants. Annual Reviews of Phytopathology. Annual Reviews of Phytopathology 57, 411-430

Villajuana-Bonequi M, Matei A, Ernst C, Hallab A, Usadel B, Doehlemann G (2019) Cell type specific transcriptional reprogramming of maize leaves during Ustilago maydis induced tumor formation. Scientific Reports 10227

Eitzen K, Schulze Hüynck J, Schurack S, Doehlemann G. (2019) Krieg und Frieden: molekulares Ping-Pong zwischen Pilzen und Pflanzen. BIOspektrum 25(4) 378-381

Schulze Hüynck J, Kaschani F, van der Linde K, Ziemann S, Müller AN, Kaiser M, Misas Villamil J, Doehlemann G. (2019) Proteases underground: analysis of the maize root apoplast identifies organ specific papain-like cysteine protease activity. Frontiers in Plant Science 10:473

Misas Villamil J, Mueller A, Demir F, Meyer U, Ökmen B, Schulze Hüynck J,  Breuer M, Dauben H, Win J,  Huesgen P, Doehlemann G. (2019) A fungal substrate mimicking molecule suppresses plant immunity via an inter-kingdom conserved motif. Nature Communications 10(1):1576

Sharma R, Ökmen B, Doehlemann G, Thines M. (2019) Saprotrophic yeasts formerly classified as Pseudozyma have retained a large effector arsenal, including functional Pep1 orthologs. Mycological Progress 18:763-768


2017 - 2018

Ökmen B, Mathow D, Hof A, Lahrmann U, Aßmann D, Doehlemann G. (2018) Mining the effector repertoire of the biotrophic fungal pathogen Ustilago hordei during host and non-host infection. Molecular Plant Pathology 19(12)_2603-2622

Ökmen B, Kemmerich B, Hilbig D, Wemhöner R, Aschenbroich J, Perrar A, Huesgen PF, Schipper K, Doehlemann G. (2018) Dual function of a secreted fungalysin metalloprotease in Ustilago maydis. New Phytologist  220(1):249-261

van der Linde K, Timofejeva L, Egger RL, Ilau B, Hammond R, Teng C, Meyers B, Doehlemann G, Walbot V. (2018) Pathogen trojan horse delivers bioactive host protein to alter maize (Zea mays) anther cell behavior in situ. Plant Cell 30(3):528-542

Ziemann S, van der Linde K, Lahrmann U, Acar B, Kaschani F, Colby T, Kaiser M, Ding Y, Schmelz E, Huffaker A, Holton N, Zipfel C, Doehlemann G. (2018) An apoplastic peptide activates salicylic acid signalling in maize. Nature Plants 4(3): 172-180

Matei  A, Ernst C, Günl M, Thiele B, Altmüller J, Walbot V, Usadel B, Doehlemann G. (2018) How to make a tumour: cell type specific dissection of Ustilago maydis-induced tumour development in maize leaves. New Phytologist 217:1681–1695.
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Redkar A, Matei A, Doehleman G. (2017) Insights into host cell modulation and induction of new celly by the corn smut Ustilago maydis. Frontiers in Plant Science  29;8:899.

Kruse J, Doehlemann  G , Kemen  E , Thines M. (2017). Asexual and sexual morphs of Moesziomyces revisited. IMA Fungus 8, 117-129.

Doehlemann G, Ökmen B, Zuh W, Sharon A. (2017) Plant Pathogenic Fungi. Microbiol Spectr. 5(1) doi: 10.1128/microbiolspec.FUNK-0023-2016.


2015 - 2016

Ökmen, B. and Doehlemann, G. (2016) Clash between the borders: spotlight on apoplastic processes in plant-microbe interactions. New Phytologist 212;4:799-801 Link

Matei, A. and Doehlemann, G. (2016) Cell Biology of Corn Smut Disease - Ustilago maydis as a Model for Biotrophic Interactions. Curr Opin Microbiol 6,34:60-66 Pubmed

Misas-Villamil, JC., van der Hoorn, RAL., Doehlemann, G. (2016) Papain-like Cys proteases as hubs in plat immunity. New Phytologist 212;4:902-907 Pubmed

Doehlemann G., Ökmen B., Zuh W., Sharon A. (2016) Plant Pathogenic Fungi. The Fungal Kindom.  [in press]

Hampel, M., Jakobi, M., Schmitz, L., Meyer, U., Finkernagel, F., Doehlemann, G., Heimel, K. (2016) Unfolded protein response regulator Cib1 mediates expression of secreted virulence factors in Ustilago maydis. PloS One 11(4):e0153861 Pubmed

Redkar, A. and Doehlemann, G. (2016). EdU Based DNA Synthesis and Cell Proliferation Assay in Maize Infected by the Smut Fungus Ustilago maydis. Bio-protocol 6(6): e1761 PDF

Redkar, A. and Doehlemann, G. (2016). Ustilago maydis Virulence Assays in Maize. Bio-protocol 6(6):e1760

Brych, A., Mascarenhas, J., Jaeger, E., Charkiewicz, E., Pokorny, R., Bölker, M., Doehlemann, G., Batschauer, A. (2015) White collar 1-induced photolyase expression contributes to UV-tolerance of Ustilago maydis. Microbiology open 5(2)_224-43 Pubmed

Redkar, A., Villajuana-Bonequi, Doehlemann, G. (2015) Conservation of the Ustilago maydis effector See1 in related smuts. Sign Plant Behav 10(12):e1086855 Pubmed

Christensen, SA., Huffaker, A., Kaplan, F., Sims, J., Ziemann, S., Doehlemann, G., Ji L., Schmitz, RJ., Kolomiets, MV., Alborn, HT., Mori, N., Ni, X., Byers, S., Abdo, Z., Schmelz, EA. (2015) Maize death acids, 9-lipoxygenase derived cyclopentenones and derivatives, display activity as cytotoxic phytoalexins and transcriptional mediators. PNAS 8;112:11407-12 Pubmed

Redkar, A., Hoser, R., Schilling, L., Zechmann, B., Krzymowska, M., Walbot, V., Doehlemann, G. (2015) A Secreted effector protein of Ustilago maydis guides maize leaf cells to form tumors. Plant Cell 27(4):1332-51 Pubmed

Hemetsberger, C., Mueller, A., Matei, A., Herrberger, C., Hensel, G., Kumlehn, J., Mishra, B., Sharma, R., Thines, M., Hückelhoven, R., Doehlemann, G. (2015) The fungal core effector Pep1 is conserved across smuts of dicots and monocots. New Phytologist 206(3):1116-26 Pubmed


2013 - 2014

Doehlemann, G., Requena, N., Schaefer, P., Brunner, F., O’Connel, R., Parker JE. (2014) Reprogramming of plant cells by filamentous plant-colonizing microbes. New Phytologist 204(4):803-14 PDF

Schilling, L., Matei, A., Redkar, A., Walbot, V., Doehlemann, G. (2014) Virulence of the maize smut Ustilago maydis is shaped by organ-specific effectors. Molecular Plant Pathology 15:780-9 Pubmed

Brefort, T., Tanaka, S., Neidig, N., Doehlemann, G., Vincon, V., Kahmann, R. (2014) Characterization of the largest effector gene cluster of Ustilago maydis. PloS Pathogens 10(7):e1003866 PDF

Ökmen, B., Doehlemann, G. (2014) Inside plant: biotrophic strategies to modulate host immunity and metabolism. Curr Opin Plant Biol. 20:19-25 PDF

Doehlemann, G., Schirawski, J., Kämper, J. (2014) Functional genomics of smut fungi - from genome sequencing to protein function. Advances in Botanical Research. Genomics of Plant-Related Fungi. 70: 143-172 PDF

Hof, A., Zechmann, B., Schwammbach, D., Hückelhoven, R., Doehlemann, G. (2014) Alternative cell death mechanisms determine epidermal resistance in incompatible barley-Ustilago interactions. Molecular Plant-Microbe Interactions 27(5):403-14 Pubmed

Rabe, F., Ajami-Rashidi, Z., Doehlemann, G., Kahmann, R., Djamei, A. (2013) Degradation of the plan defence hormone salicylic acid by the biotrophic fungus Ustilago maydis. Molecular Microbiology 89:179-188 Pubmed

Doehlemann, G. and Hemetsberger, C. (2013) Apoplastic immunity and its suppression by filamentous microbial pathogens. New Phytologist 198:1001-16 Pubmed

van der Linde, K., Mueller, A., Redkar, A., Schilling, L., Doehlemann, G. (2013) Brandpilz mit Pep: Ustilago maydis und das pflanzliche Immunsystem. Biospektrum 19:140-142 PDF

Mueller, AN., Ziemann, S., Treitschke, S., Aßmann, D., Doehlemann, G. (2013) Compatibility in the Ustilago maydis – maize interaction requires inhibition of host cysteine proteases by the fungal effector Pit2. PloS Pathogens 9(2): e1003177 Highlighted by: De Wit P: F1000Prime Recommendation of [Mueller, AN. et al., PLoS Pathog 2013, 9(2):e1003177]. In F1000Prime, 25 Feb 2013 PDF

van der Linde, K. and Doehlemann, G. (2013) Utilizing virus induced gene silencing for the functional characterization of maize genes during infection with the fungal pathogen Ustilago maydis. Methods in Molecular Biology 975:47-60 Pubmed


2011 - 2012

van der Linde, K., Mueller, A., Hemetsberger, C., Kaschani, F., van der Hoorn, RAL.,  Doehlemann, G. (2012) The maize cystatin CC9 interacts with apoplastic cysteine proteases. Plant Signal Behav. 7(11):1397-401 Pubmed

Doehlemann, G. (2012b) The maize cystatin CC9 interacts with apoplastic cysteine proteases. Plant Signaling & Behavior 7:1397-1401 PDF

Hemetsberger, C., Herrberger, C., Zechmann, B., Hillmer, M., Doehlemann, G. (2012) The Ustilago maydis effector Pep1 suppresses plant immunity by inhibition of host peroxidase activity. PloS Pathogens 8(5): e1002684 Highlighted by: Jermy, A. (2012) Ustilago gives plants a Pep talk. Nature Reviews Microbiology 10, 444-445 Pubmed

van der Linde, K., Hemetsberger, C., Kastner, C., Kaschani, F., van der Hoorn, RAL., Kumlehn, J., Doehlemann, G. (2012a) A maize cystatin suppresses host immunity by inhibition of apoplastic cysteine proteases. Plant Cell 24:1285-1300 PDF

Meier, K., Mathieu, ELl., Reuter, LM,, Scharfe, M., Doehlemann, G., Jarek, M., Brehm, A. (2012) LINT - a novel dL(3)mbt containing complex represses malignant brain tumour signature genes. PloS Genetics 8(5):e1002676 PDF

Voll, LM., Horst, RJ., Voitsik, AM., Zajic, D., Samans, B., Pons-Kühnemann, J., Doehlemann, G., Münch, S., Wahl, R., Molitor, A., Hofmann, J., Schmiedl, A., Waller, F., Deising, HB., Kahmann, R., Kämper, J., Kogel HK., Sonnewald, U. (2011) Common motifs in the response of cereal primary metabolism to fungal pathogens are not based on similar transcriptional reprogramming. Frontiers in Plant Physiology 2:81-6 Pubmed

Doehlemann, G., Reissmann, S., Aßmann, D., Fleckenstein, M., Kahmann, R. (2011) Two linked genes encoding a secreted effector and a membrane protein are essential for Ustilago maydis induced tumor formation. Molecular Microbiology 81:751-766 PDF

Leroch, M., Mernke, D., Koppenhoefer, D., Schneider, P., Mosbach, A., Doehlemann, G., Hahn, M. (2011) Living colors in the gray mold pathogen: Codon-optimized genes encoding GFP and mCherry showing bright fluorescence in Botrytis cinerea. Applied Environmental Biology 77: 2887-97 PDF

van der Linke, K., Kastner, C., Kumlehn, J., Kahmann, R., Doehlemann, G. (2011) Systemic virus induced gene silencing allows functional characterization of maize genes during the biotrophic interaction with Ustilago maydis. New Phytologist 189: 471-483 Highlighted by: Benavente, L. and Scofield, R. (2011) A new tool for functional genomics in maize. New Phytologist 189: 363-365 Pubmed

de Vries, RP., Benoit, I., Doehlemann, G., Kobayashi, T., Magnuson, JK., Panisko, EA., Baker, SE., Lebrun, MH. (2011) Post-genomic approaches to understanding interactions between fungi and their environment. IMA Fungus 2:81-86 PDF

Schipper, K. and Doehlemann, G. (2011) Compatibility in biotrophic plant-fungal interactions: Ustilago maydis and friends. In Signaling and Communication on Plant Symbiosis. (Perotto S and Baluska F, eds.) Springer Verlag, Berlin, Germany, 213:238 PDF

Doehlemann, G., Schipper, K., Kahmann, R. (2011) The effectors of smut fungi. In Effectors in Plant-Microbe Interactions (Martin F and Kamoun S, eds), Wiley VHC:77-100 PDF


2009 - 2010

Schirawski, J., Mannhaupt, G., Münch, K., Brefort, T., Schipper, K., Doehlemann, G., Di Stasio, M., Rössel, N., Mendoza-Mendoza, A., Pester, D., Müller, O., Winterberg, B., Meyer, E., Ghareeb, H., Wollenberg, T., Münsterkötter, M., Wong, P., Walter, M., Stukenbrock, E., Güldener, U., Kahmann, R. (2010) Pathogenicity determinants in smut fungi revealed by genome comparison. Science 330: 1546-8 Pubmed

Horst, RJ., Doehlemann, G., Wahl, R., Hofmann, J., Schmiedl, A., Kahmann, R., Kämper, J., Voll, LM. (2010) A model of Ustilago maydis leaf tumor metabolism. Plant Signaling & Behavior 5: 11, 1-4 PDF

Treitschke, S., Doehlemann, G., Schuster, M., Steinberg, G. (2010) The Myosin Motor Domain of Fungal Chitin Synthase V Is Dispensable for Vesicle Motility but Required for Virulence of the Maize Pathogen Ustilago maydis. Plant Cell 22: 2476-2494 PDF

Skibbe, D*., Doehlemann, G*., Fernandes, J., Walbot, V. (2010) Maize tumors caused by Ustilago maydis require organ-specific genes in host and pathogen. Science 328: 89-92 *contributed equally Highlighted by: Zahn, LM. (2010) Tailor made tumor. Science Signaling 3: 104, Villanueva, T. (2010) Tumorigenisis: Popping oncorngenes. Nature Reviews Cancer 10:314-315 Pubmed

Horst, R., Doehlemann, G., Wahl, R., Hofmann, J., Schmiedl, A., Kahmann, R., Kämper, J., Sonnewald, U., Voll L. (2010) Ustilago maydis infection strongly alters organic nitrogen allocation in maize and stimulates productivity of systemic source leaves. Plant Physiology 152: 293-30 PDF

Doehlemann, G., van der Linde, K., Aßmann, D., Schwammbach, D., Hof, A., Mohanty, A., Jackson, D., Kahmann, R. (2009) Pep1, a secreted effector protein of Ustilago maydis is required for successful invasion of plant cells. PLoS Pathogens 5: 2 e1000290 PDF

Brefort, T., Doehlemann, G., Mendoza-Mendoza, A., Reissmann, S., Djamei, A., Kahmann, R. (2009) Ustilago maydis as a pathogen. Annual Reviews of Phytopathology 47: 423-445 PDF

Schipper, K., Doehlemann, G., Brefort, T., Djamei, A., Liang, L., Khrunyk, Y., Schirawski, J., Kahmann, R. (2009) The effectors of smut fungi. Biology of Molecular Plant-Microbe Interactions, Vol. 7 (2009 IS-MPMI Symposium Proceedings) (H. Antoun et al., eds.)


2005 - 2008

Doehlemann, G.*, Wahl, R.*, Horst, R., Voll, L., Usadel, B., Poree, F., Stitt, M., Pons-Kühnemann, J., Sonnewald, U., Kahmann, R., Kämper, J. (2008b)  Reprogramming a maize plant: Transcriptional and metabolic changes induced by the fungal biotroph Ustilago maydis. Plant Journal, 56:181-195 *contributed equally Pubmed

Doehlemann, G., Wahl, R., Vranes, M., de Vries, RP., Kämper, J., Kahmann, R. (2008a) Establishment of compatibility in the Ustilago maydis/maize pathosystem. Journal of  Plant Physiology 165: 29-40 Pubmed

Brefort, T., Molina, L., Doehlemann, G., Schipper, K., Mendoza-Mendoza, A., Mueller, O., Schirawski, J. and Kahmann, R. (2008) The role of secreted proteins during establishment of biotrophy in the Ustilago maydis/maize pathosystem. Biology of Molecular Plant-Microbe Interactions, Vol. 6 (Lorito, M., Woo, S. and Scals, F., eds.)

Brefort, T., Schipper, K., Doehlemann, G. and Kahmann, R. (2008) The biotrophic phase of Ustilago maydis: novel determinants of compatibility. Genomics of Disease (P. Gustafson, ed.), Springer Verlag, Berlin, Germany PDF

Doehlemann, G., Berndt, P. and Hahn, M. (2006b) Trehalose metabolism is important for heat stress tolerance and spore germination of Botrytis cinerea. Microbiology 152: 2625-2634 Pubmed

Doehlemann, G., Berndt, P. and Hahn M. (2006a) Different signalling pathways involving a Gα protein, cAMP and a MAP kinase control germination of Botrytis cinerea conidia. Molecular Microbiology 59: 821-835 Pubmed

Doehlemann, G., Molitor, F and Hahn, M. (2005) Molecular and functional characterization of a fructose specific transporter from the gray mold fungus Botrytis cinerea. Fungal Genetics & Biology 42: 601-610 Pubmed