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Publication list of Thomas Schmülling

>145 peer-reviewed publications with ~13.100 citations, h-index 62 (Clarivate), 30 April 2023.
>185 invited talks at conferences, research institutes and universities around the world


HÖNIG, M., ROEBER, V.M., SCHMÜLLING, T., CORTLEVEN, A. (2023) Chemical priming of plant defense responses to pathogen attacks. Front. Plant Sci. 14:1146577. doi: 10.3389/fpls.2023.1146577

TALESKI, M., CHAPMAN, K., NOVÁK, O., SCHMÜLLING, T., FRANK, M., DJORDJEVIC, M.A. (2023) CEP peptide and cytokinin pathways converge on CEPD glutaredoxins to inhibit root growth. Nature Communications 14, 1683; doi: 10.1038/s41467-023-37282-6.


RIEFLER, M., BRÜGMANN, T., FLADUNG M., SCHMÜLLING, T. (2022) A constitutively active cytokinin receptor variant increases cambial activity and stem growth in poplar. International Journal of Molecular Sciences 23, 8321; https://doi.org/10.3390/ijms23158321

CORTLEVEN, A., ROEBER, V.M., FRANK, M., BERTELS, J., LORTZING, V., BEEMSTER, G., SCHMÜLLING, T. (2022) The transcriptomic landscape of the photoperiodic stress response in Arabidopsis thaliana resembles the response to pathogen infection. Frontiers in Plant Science 13, 838284. https://doi.org/10.3389/fpls.2022.838284

FRANK, M., CORTLEVEN, A., SCHMÜLLING, T. (2022) The response to photoperiod stress in Arabidopsis thaliana depends on auxin acting as an antagonist to the protectant cytokinin. International Journal of Molecular Sciences 23, 2936; https://doi.org/10.3390/ijms23062936

PINO, L.E., LIMA, J.E., VICENTE, M.H., DE SÁ, A.F.L., PÉREZ-ALFOCEA, F., ALBACETE, A., COSTA, J.L., WERNER, T., SCHMÜLLING, T., FRESCHI, L., FIGUEIRA, A., ZSÖGÖ, A., PERES, L.E.P. (2022) Increased branching independent of strigolactone in cytokinin oxidase2-overexpressing tomato is mediated by reduced auxin transport. Molecular Horticulture 2, 12. https://doi.org/10.1186/s43897-022-00032-1

ROMANOV G.A., SCHMÜLLING, T. (2022) On the biological activity of cytokinin free bases and their ribosides. Planta 255, 27. https://doi.org/10.1007/s00425-021-03810-1

ROEBER, V.M., SCHMÜLLING, T., CORTLEVEN, A. (2022) The photoperiod: handling and causing stress in plants. Frontiers in Plant Science 12, 781998. https://doi.org/10.3389/fpls.2021.781988


WERNER, T., SCHMÜLLING, T. (2021) Moonlighting PPKL1 reveals a role of cytokinin in regulating rice grain size. Molecular Plant 15, 216-218. https://doi.org/10.1016/j.molp.2021.11.014

WERNER S., BARTRINA, I., SCHMÜLLING T. (2021) Cytokinin regulates vegetative phase change in Arabidopsis thaliana through the miR172/TOE1-TOE2 module. Nature Communications 12, 5816. https://doi.org/10.1038/s41467-021-26088-z

LEUENDORF, J.E., SCHMÜLLING, T. (2021) Meeting at the DNA: Specifying cytokinin responses by modulating the transcriptional output. Plants 10, 1458. https://doi.org/10.3390/plants10071458

RAMIREDDY, E., NELISSEN, H., LEUENDORF, J.E., VAN LIJSEBETTENS, M., INZÉ, D., SCHMÜLLING, T. (2021) Root engineering in maize by increasing cytokinin degradation causes enhanced root growth and leaf mineral enrichment. Plant Mol. Biol. 106, 555–567. https://doi.org/10.1007/s11103-021-01173-5

ROMANOV, G., SCHMÜLLING, T.(2021) Opening doors for cytokinin at the ER membrane. Trends Plant Sci. 26, 305-308. https://doi.org/10.1016/j.tplants.2021.02.006

WERNER, S., BARTRINA, I., NOVÁK, O., STRNAD, M., WERNER, T. SCHMÜLLING, T.(2021) The cytokinin status of the epidermis regulates aspects of vegetative and reproductive development in Arabidopsis thaliana. Front. Plant Sci. 12, 613488. https://doi.org/10.3389/fpls.2021.613488

HALAWA, M., CORTLEVEN, A. SCHMÜLLING, T. HEYL, A. (2021) Characterization of CHARK, a novel type cytokinin receptor from Oryza sativa. Scientific Rep. 11, 1722. https://doi.org/10.1038/s41598-020-80223-2

ROEBER, V.M., BAJAJ, I., ROHDE, M., SCHMÜLLING, T., CORTLEVEN, A.(2021) Light acts as a stressor and influences abiotic and biotic stress responses in plants. Plant Cell Environ. 44, 645-664. https://doi.org/10.1111/pce.13948


SCHWARZ, I., OTTO, E., MARIE-THERESE SCHEIRLINCK, M.-T., BARTRINA, I. SCHMIDT, R., SCHMÜLLING, T. (2020) Cytokinin regulates the activity of the inflorescence meristem and components of seed yield in oilseed rape. J. Exp. Bot. 71, 7146-7159. https://doi.org/10.1093/jxb/eraa419

This article has been highlighted: JAMESON, P.E., SONG., J. (2020) Will cytokinins underpin the second ‘Green Revolution’? J. Exp. Bot. 71, 6872–6875. https://doi.org/10.1093/jxb/eraa447

FRANK, M., CORTLEVEN, A., NOVAK, O., SCHMÜLLING, T.(2020) Root-derived trans-zeatin cytokinin protects Arabidopsis plants against photoperiod stress. Plant Cell Environm. 43, 2637–2649. https://doi.org/10.1111/pce.13860

ABUELSOUD, W., CORTLEVEN, A., SCHMÜLLING, T. (2020) Photoperiod stress induces an oxidative burst-like response and is associated with increased apoplastic peroxidase and decreased catalase activities. J. Plant Physiol., 153252. https://doi.org/10.1016/j.jplph.2020.153252

ILLGEN, S., ZINTL, S., ZUTHER, E., HINCHA, D.K., SCHMÜLLING, T. (2020) Characterisation of the ERF102 to ERF105 genes of Arabidopsis thaliana and their role in the response to cold stress. Plant Mol. Biol. 103, 303–320. https://doi.org/10.1007/s11103-020-00993-1

LEUENDORF, J.E., FRANK, M., SCHMÜLLING, T. (2020) Acclimation, priming and memory of the cold stress response in Arabidopsis thaliana seedlings. Scientific Rep. 10, 689. https://doi.org/10.1038/s41598-019-56797-x


NEHNEVAJOVA, E., RAMIREDDY, E., STOLZ, A., GERDEMANN-KNÖRCK, M., NOVAK, O., STRNAD, M., SCHMÜLLING, T. (2019) Root enhancement in cytokinin-deficient oilseed rape causes leaf mineral enrichment, increases the chlorophyll concentration under nutrient limitation and enhances the phytoremediation capacity. BMC Plant Biol. 19, 83. https://doi.org/10.1186/s12870-019-1657-6

CORTLEVEN, A., EHRET, S., SCHMÜLLING, T., JOHANSSON, H. (2019) Ethylene-independent promotion of photomorphogenesis in the dark by cytokinin requires COP1 and the CDD complex. J. Exp. Bot. 70, 165-178. https://doi.org/10.1093/jxb/ery344

VOLLER, J., BÉRES, T., ZATLOUKAL, M., DŽUBÁK, P., HAJDÚCH, M., DOLEŽAL, K., SCHMÜLLING, T., STRNAD, M. (2019) Anti-cancer activities of cytokinin ribosides. Phytochem Rev. 18, 1101–1113. https://doi.org/10.1007/s11101-019-09620-4

HILKER, M., SCHMÜLLING, T. (2019) Stress priming, memory and signalling in plants. Plant Cell Environm. 42, 753-761. https://doi.org/10.1111/pce.13526

CORTLEVEN, A., LEUENDORF, J.E., FRANK, M., BOLT, S., PEZZETTA, D., SCHMÜLLING, T. (2019) Cytokinin action in response to abiotic and biotic stresses in plants. Plant Cell Environm. 42, 998-1018. https://doi.org/10.1111/pce.13494


RAMIREDDY, E., GALUSZKA, P., SCHMÜLLING, T. (2018) Zn-fortified cereal grains in field-grown barley by enhanced root cytokinin breakdown. Plant Signal. Behav. 13, 1-4. https://doi.org/10.1080/15592324.2018.1530023

RAMIREDDY, E., HOSSEINI, S.A., EGGERT, K., GILLANDT, S., GNAD, H., VON WIRÉN, N., SCHMÜLLING, T. (2018) Root engineering in barley by increasing cytokinin degradation causes mineral enrichment in leaves and seeds and enhanced drought tolerance. Plant Physiol. 177, 1078-1095. https://doi.org/10.1104/pp.18.00199

This article has been highlighted: JULKOWSKA, M: Releasing the cytokinin brakes on root growth. Plant Physiol. 177, 865-866, 2018. https://doi.org/10.1104/pp.18.00660

SAMAR, K., SCHMÜLLING, T., PAPON, N. (2018) CHASEing cytokinin receptors in plants, bacteria, fungi, and beyond. Trends Plant Sci. 23, 179-181. https://doi.org/10.1016/j.tplants.2018.01.001

ROMANOV, G.A., LOMIN, S., SCHMÜLLING, T. (2018) Cytokinin signaling: from the ER or from the PM? That is the question! New Phytol. 218, 41-53. (Tansley Review) https://doi.org/10.1111/nph.14991

WITZEL, K., MATROS, A., MØLLER, A.L.B., RAMIREDDY, E., FINNIE, C., PEUKERT, M., RUTTEN, T., KUNZE, G., MELZER, M., KASPAR, S., SCHMÜLLING, T., SVENSSON, B., MOCK, H.-P. (2018) Plasma membrane proteomic analysis identifies a role of barley Membrane Steroid Binding Protein in root architecture response to salinity. Plant Cell Environm. 41, 1311-1330. https://doi.org/10.1111/pce.13154

LOMIN, S.N., ARKHIPOV, D.V., MYAKUSHINA, Y.A., LEONOVA, O.G., POPENKO, V.L., SCHMÜLLING, T., ROMANOV, G.A. (2018) Studies of cytokinin receptor–phosphotransmitter interaction provide evidences for the initiation of cytokinin signaling in the endoplasmic reticulum. Funct. Plant Biol. 45, 192–202. https://doi.org/10.1071/FP16292


NITSCHKE, S., CORTLEVEN, A., SCHMÜLLING, T. (2017) Novel stress in plants by altering the photoperiod. Trends in Plant Sci. 22, 913-916. https://doi.org/10.1016/j.tplants.2017.09.005

Dowd, C.D., Chronis, D., Radakovic, Z.S., Siddique, S., Schmülling, T., Werner, T., Kakimoto, T., Grundler, F.M.W., Mitchum, M.G. (2017) Divergent expression of cytokinin biosynthesis, signaling and catabolism genes underlying differences in feeding sites induced by cyst and root-knot nematodes. Plant J. 92, 211-228.

Brenner, W.G., Leuendorf, J.E., Cortleven, A., Schmülling, T. (2017) CFB, a cytokinin-regulated gene encoding a novel F-box protein with a putative function in chloroplast development. J. Exp. Bot. 68, 2769-2785.

Lomin, S.N., Arkhipov, D.V., Myakushina, Y.A., Leoniva, O.G., Popenko, V.L., Schmülling, T., Romanov, G.A. (2017) Studies of cytokinin receptor-phosphotransmitter interaction provide evidences for the initiation of cytokinin signaling in the endoplasmic reticulum. Funct. Plant Biol. 45, 192–202.

Bartrina, I., Jensen, H., Novak, O., Strnad, M., Werner, T., Schmülling, T. (2017) Gain-of-function mutants of the cytokinin receptors AHK2 and AHK3 regulate plant organ size, flowering time and plant longevity. Plant Physiol. 173, 1783-1797.


Hilker, M., Schwachtje, J., Baier, M., Balazadeh, S., Bäurle, I., Geiselhardt, S., Hincha, D.K., Kunze, R., Müller-Röber, B., Rillig, M.C., Rolff, J., Romeis, T., Schmülling, T., Steppuhn, A., van Dongen, J., Whitcomb, S.J., Wurst, S., Zuther, E., Kopka, J. (2016) Priming and memory of stress responses in organisms lacking a nervous system. Biol. Rev. Camb. Philos. Soc. 91, 1118-1133.

Bolt, S., Zuther, E., Zintl, S., Hincha, D.K., Schmülling, T. (2016) ERF105 is a transcription factor gene of Arabidopsis thaliana required for freezing tolerance and cold acclimation. Plant Cell Environ. 40, 108-120.

Cosme, M., Ramireddy, E., Franken, P., Schmülling, T., Wurst, S. (2016) Shoot- and root-borne cytokinin influences arbuscular mycorrhizal symbiosis. Mycorrhiza 26, 709-720.

Cortleven, A., Marg, I., Yamburenko, M.V., Schlicke, H., Hill, K., Grimm, B., Schaller, G.E., Schmülling, T. (2016) Cytokinin Regulates the Etioplast-Chloroplast Transition through the Two-Component Signaling System and Activation of Chloroplast-Related Genes. Plant Physiol. 172, 464-478.

Nitschke, S., Cortleven, A., Iven, T., Feussner, I., Havaux, M., Riefler, M., Schmülling, T. (2016) Circadian Stress Regimes Affect the Circadian Clock and Cause Jasmonic Acid-Dependent Cell Death in Cytokinin-Deficient Arabidopsis Plants. Plant Cell 28, 1616-1639.

Mohan, T.C., Castrillo, G., Navarro, C., Zarco-Fernández, S., Ramireddy, E., Mateo, C., Zamarreño, A.M., Paz-Ares, J., Muñoz, R., García-Mina, J.M., Hernández, L.E., Schmülling, T., Leyva, A. (2016) Cytokinin Determines Thiol-Mediated Arsenic Tolerance and Accumulation. Plant Physiol. 171, 1418-1426.


Siddique, S., Radakovic, Z.S., De La Torre, C.M., Chronis, D., Novák, O., Ramireddy, E., Holbein, J., Matera, C., Hütten, M., Gutbrod, P., Anjam, M.S., Rozanska, E., Habash, S., Elashry, A., Sobczak, M., Kakimoto, T., Strnad, M., Schmülling, T., Mitchum, M.G., Grundler, F.M. (2015) A parasitic nematode releases cytokinin that controls cell division and orchestrates feeding site formation in host plants. Proc. Natl. Acad. Sci. USA 112, 12669-12674.

Chang, L., Ramireddy, E., Schmülling, T. (2015) Cytokinin as a positional cue regulating lateral root spacing in Arabidopsis. J. Exp. Bot. 66, 4759-4768.

Cortleven, A., Schmülling, T. (2015) Regulation of chloroplast development and function by cytokinin. J. Exp. Bot. 66, 4999-5013.

Brenner, W.G., Schmülling, T. (2015) Summarizing and exploring data of a decade of cytokinin-related transcriptomics. Front. Plant Sci. 6, 29.

Lomin, S.N., Krivosheev, D.M., Steklov, M.Y., Arkhipov, D.V., Osodlokin, D.I., Schmülling, T., Romanov. G.A. (2015) Plant membrane assays with cytokinin receptors underpin the unique role of free cytokinin bases as biologically active ligands. J. Exp. Bot. 66, 1851-1863.

Niemann, M.C.E., Bartrina, I., Ashikov, A., Weber, H., Novák, O., Spíchal, L., Strnad, M., Bakker, H., Schmülling, T. Werner, T. (2015) Arabidopsis ROCK1 transports UDP-GlcNAc and UDP-GalNAc and is required for the ER protein quality control and modulation of cytokinin activity. Proc. Natl. Acad. Sci. USA 112, 291-296.


Köllmer, I., Novak, O., Strnad, T., Schmülling, T., Werner, T. (2014) Overexpression of the cytosolic cytokinin oxidase/dehydrogenase (CKX7) from Arabidopsis causes specific changes in root growth and xylem differentiation. Plant J. 78, 359-371. (featured article)

Ramireddy, E., Chang., L., Schmülling, T. (2014) Cytokinin as a mediator for regulating root system architecture in response to environmental cues. Plant Signal. Behav. 9, e27771.

Cortleven, A., Nitschke, S., Klaumünzer, M., AbdElgawad, H., Asard, H., Grimm, B., Lurz, R., Riefler, M., Schmülling, T. (2014) A novel protective function for cytokinin in the light stress response is mediated by the ARABIDOPSIS HISTIDINE KINASE2 and ARABIDOPSIS HISTIDINE KINASE3 Receptors. Plant Physiol. 164, 1470-1483.


Chang, L., Ramireddy, E., Schmülling, T. (2013) Lateral root formation and growth of Arabidopsis is redundantly regulated by cytokinin metabolism and signalling genes. J. Exp. Bot. 64, 5021-5032.

Cervinkova, H., Hronková, M., Dobra, J., Tureckova, V., Novak, O., Lubovska, Z., Motyka, V., Haisel, D., Hajek, T., Prasil, I.T., Gaudinova, A., Storchova, H., Werner, T., Schmülling, T., Vankova R. (2013) Enhanced drought and heat stress tolerance of tobacco plants with ectopically enhanced cytokinin oxidase/dehydrogenase gene expression. J. Exp. Bot. 64, 2805-2815.

Ramireddy, R., Pfeifer, A., Brenner, W.G., Heyl, A.*, Schmülling, T.* (2013) In planta analysis of a cis-regulatory cytokinin response motif in Arabidopsis and identification of a novel enhancer sequence. Plant Cell Physiol. 54, 1079-1092. (*corresponding authors)

Frémont, N., Riefler, M., Stolz, A., Schmülling, T. (2013) The Arabidopsis TUMOR PRONE5 (TUP5) Gene Encodes an Acetylornithine Aminotransferase Required for Arginine Biosynthesis and Root Meristem Maintenance in Blue Light. Plant Physiol. 161, 1127-1140.


Brenner, W.G., Schmülling, T. (2012) Transcript profiling of cytokinin action in Arabidopsis roots and shoots discovers largely similar but also organ-specific responses. BMC Plant Biol. 12, 112.

Brenner, W.G., Ramireddy, E., Heyl, A., Schmülling, T. (2012) Gene regulation by cytokinin in Arabidopsis. Front. Plant. Sci. 3, 8.

Nehnevajova, E., Lyubenova, L., Herzig, R., Schröder, P., Schwitzguébel, J.-P., Schmülling, T. (2012) Metal accumulation and response of antioxidant enzymes in seedlings and adult sunflower mutants with improved metal removal traits on a metal-contaminated soil. Env. Exp. Bot. 76, 39-48.


Nishiyama, R., Watanabe, Y., Fujita, Y., Le, D.T., Kojima, M., Werner, T., Vankova, R., Yamaguchi-Shinozaki, K., Shinozaki, K., Kakimoto, T., Sakakibara, H., Schmülling, T, Tran, L.S. (2011) Analysis of cytokinin mutants and regulation of cytokinin metabolic genes reveals important regulatory roles of cytokinins in drought, salt and abscisic acid responses, and abscisic acid biosynthesis. Plant Cell 23, 2169-2183.

Wulfetange, K., Lomin, S.N., Romanov, G.A., Stolz, A., Heyl, A., Schmülling, T. (2011) The cytokinin receptors of Arabidopsis are located mainly to the endoplasmic reticulum. Plant Physiol. 156, 1808-1818.

Heyl, A., Riefler, M., Romanov, G.A., Schmülling, T. (2011) Properties, functions and evolution of cytokinin receptors. Eur. J. Cell Biol. 91, 246-256.

Holst, K., Schmülling, T., Werner, T. (2011) Enhanced cytokinin degradation in leaf primordia of transgenic Arabidopsis plants reduces leaf size and shoot organ primordia formation. J. Plant Physiol. 168, 1328-1334.

Köllmer, I., Werner, T., Schmülling, T. (2011) Ectopic expression of different cytokinin-regulated transcription factor genes of Arabidopsis thaliana alters plant growth and development. J. Plant Physiol. 168, 1320-1327.

Stolz, A., Riefler, M., Lomin, S.N., Achazi, K., Romanov, G.A., Schmülling, T. (2011) The specificity of cytokinin signalling in Arabidopsis thaliana is mediated by differing ligand affinities and expression profiles of the receptors. Plant J. 67, 157-168.

Bartrina, I., Otto, E., Strnad, M., Werner, T., Schmülling, T. (2011) Cytokinin regulates the activity of reproductive meristems, flower organ size, ovule formation, and thus seed yield in Arabidopsis thaliana. Plant Cell 23, 69-80.

Vercruyssen, L., Gonzalez, N., Werner, T., Schmülling, T., Inzé, D. (2011) Combining enhanced root and shoot growth reveals cross talk between pathways that control plant organ size in Arabidopsis. Plant Physiol. 155, 1339-1352.

Wulfetange, K., Saenger, W., Schmülling, T., Heyl, A. (2011) E. coli-based cell-free expression, purification and characterization of the membrane-bound ligand-binding CHASE-TM domain of the cytokinin receptor CRE1/AHK4 of Arabidopsis thaliana. Mol. Biotechnol. 47, 211-219.


Werner, T., Nehnevajova, E., Köllmer, I., Novák, O., Strnad, M., Krämer, U., Schmülling, T. (2010) Root-specific reduction of cytokinin causes enhanced root growth, drought tolerance, and leaf mineral enrichment in Arabidopsis and tobacco. Plant Cell 22, 3905-3920.

Pertry, I., Václavíková, K., Gemrotová, M., Spíchal, L., Galuszka, P., Depuydt, S., Temmerman, W., Stes, E., De Keyser, A., Riefler, M., Biondi, S., Novák, O., Schmülling, T., Strnad, M., Tarkowski, P., Holsters, M., Vereecke, D. (2010) Rhodococcus fascians impacts plant development through the dynamic fas-mediated production of a cytokinin mix. Mol. Plant-Microbe Interact. 23, 1164-1174.


Krupková, E., Schmülling, T. (2009) Developmental consequences of the tumorous shoot development1 mutation, a novel allele of the cellulose-synthesizing KORRIGAN1 gene. Plant Mol. Biol. 71, 641-655.

Werner, T., Schmülling, T. (2009) Cytokinin action in plant development. Curr. Opin. Plant Biol. 12, 527-538.

Pertry, I., Václavíková, K., Depuydt, S., Galuszka, P., Spíchal, L., Temmerman, W., Stes, E., Schmülling, T., Kakimoto, T., Van Montagu, M.C., Strnad, M., Holsters, M., Tarkowski, P., Vereecke, D. (2009) Identification of Rhodococcus fascians cytokinins and their modus operandi to reshape the plant. Proc. Natl. Acad. Sci USA 106, 929-934.

Spíchal, L., Werner, T., Popa, I., Riefler, M., Schmülling, T., Strnad, M. (2009) The purine derivative PI-55 blocks cytokinin action via receptor inhibition. FEBS J. 276, 244-253.

Nehnevajova, E., Herzig, R., Schwitzguébel, J.P., Schmülling, T. (2009) Sunflower mutants with improved growth and metal accumulation traits show a potential for soil decontamination. In: Induced Plant Mutations in the Genomic Era. Food and Agriculture Organization of the United Nations (ed. Q.Y.Shu). Rome, 93-96.

Nehnevajova, E., Schmülling, T., Herzig, R. Schwitzguébel, P. (2009) An increased tolerance of sunflower mutant seedlings to Cd and Zn in hydroponic culture. Agrochimica 3, 1-15.


Heyl, A., Ramireddy, E., Brenner, W.G., Riefler, M., Allemeersch, J., Schmülling, T. (2008) The transcriptional silencer ARR1-SRDX suppresses pleiotropic cytokinin activities in Arabidopsis. Plant Physiol. 147, 1380-1395.

Werner, T., Holst, K., Pörs, Y., Guivarc'h, A., Mustroph, A., Chriqui, D., Grimm, B., Schmülling, T. (2008) Cytokinin deficiency causes distinct changes of sink and source parameters in shoots and roots. J. Exp. Bot. 59, 2659-2672.

Dortay, H., Mehnert, N., Pfeifer, A., Schwerdtner, M., Schmülling, T., Heyl, A. (2008) Towards an interaction map of the two-component signaling pathway of Arabidopsis thaliana. J. Proteome Res. 7, 3649-3660.

Romanov, G., Lomin, S.L, Rakova, N.Y., Heyl, A., Schmülling, T. (2008) Does NO play a role in cytokinin signal transduction? FEBS Lett. 582, 874-880.


Krupková, E., Immerzeel, P., Pauly, M., Schmülling, T. (2007) The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant development. Plant J. 50, 735-750.

Heyl, A., Wulfetange, K., Pils, B., Nielsen, N., Romanov, G.A., Schmülling, T., Strnad, M. (2007) Evolutionary proteomics identifies amino acids essential for ligand-binding of the cytokinin receptor CHASE domain. BMC Evol. Biol. 7, 62.

Spíchal, L., Kryštof, V., Paprskářová, M., Lenobel, R., Stýskala, J., Binarová , P., Cenklová, V., De Veylder, L., Inzé, D. Kontopidis, G., Fischer, P.M., Schmülling, T., Strnad, M. (2007) Classical anticytokinins do not interact with cytokinin receptors but inhibit cyclin-dependent kinases. J. Biol. Chem. 282, 14356-14363.

Frébortová, J., Galuszka, P., Werner, T., Schmülling, T., Frébort, I. (2007) Functional expression and purification of cytokinin dehydrogenase from Arabidopsis thaliana (AtCKX2) in Saccharomyces cerevisiae. Biol. Plant. 51, 673-682.

Galuszka, P., Popelková, H., Werner, T., Frébortová, J., Mik, V., Köllmer, I., Schmülling, T., Frébort, I. (2007) Biochemical characterization and histochemical localization of cytokinin oxidases/dehydrogenases from Arabidopsis thaliana expressed in Nicotiana tabacum L. J. Plant Growth Reg. 26, 255-267.


Romanov, G.A., Lomin, S.N., Schmülling, T. (2006) Biochemical characteristics and ligand-binding properties of Arabidopsis cytokinin receptor AHK3 compared to CRE1/AHK4 as revealed by a direct binding assay. FEBS J. 273, 4631-4644.

Dortay, H., Mehnert, N., Bürkle, L., Schmülling, T., Heyl, A. (2006) Analysis of protein interactions within the cytokinin-signaling pathway of Arabidopsis thaliana. FEBS J. 273, 4631-4644.

Werner, T., Köllmer, I., Bartrina, I., Holst, K., Schmülling, T. (2006) New insights into the biology of cytokinin degradation. Plant Biol. (Stuttg.) 8, 371-381.

Siemens, J., Keller, I., Sarx, J., Kunz, S., Schuller, A., Nagel, W., Schmülling, T., Parniske, M., Ludwig-Müller, J. (2006) Transcriptome analysis of Arabidopsis clubroots indicate a key role for cytokinins in disease development. Mol. Plant-Microbe Interact. 19, 480-494.

Riefler, M., Novak, O., Strnad, M., Schmülling, T. (2006) Arabidopsis cytokinin receptor mutants reveal functions in shoot growth, leaf senescence, seed size, germination, root development and cytokinin metabolism. Plant Cell 18, 40-54.

Doležal, K., Popa, I., Kryštof, V., Spíchal, L., Fojtíková, M., Holub, J., Lenobel, R., Schmülling, T., Strnad, M. (2006) Preparation and biological activity of 6-benzylaminopurine derivatives in plants and human cancer cells. Bioorg. Med. Chem. 14, 875-884.


Brenner, W., Romanov, G., Bürkle, L., Schmülling, T. (2005) Immediate-early and delayed cytokinin response genes of Arabidopsis thaliana identified by genome-wide expression profiling reveal novel cytokinin-sensitive processes and suggest cytokinin action through transcriptional cascades. Plant J. 44, 314-333.

Romanov, G.A., Spíchal, L., Lomin, S.N., Strnad, M., Schmülling, T. (2005) A live-cell hormone-binding assay on transgenic bacteria expressing a eukaryotic receptor protein. Anal. Biochem. 347, 129-134.

Schmülling, T., Werner, T., Krämer, U., Riefler, M. (2005) The enhanced root system of cytokinin-deficient plants and its consequences for mineral uptake. In Plant nutrition for food security, human health and environmental protection (Eds. Li, C.J. et al.), Tsinghua University Press, Beijing. pp. 474-475


Schmülling, T. (2004) Cytokinin. In Encyclopedia of Biological Chemistry (Eds. Lennarz, W., Lane, M.D.), Academic Press/Elsevier Science.

Spíchal, L., Raková, N.Y., Riefler, M., Mizuno, T., Romanov, G.A., Strnad, M., Schmülling, T. (2004) Two cytokinin receptors of Arabidopsis thaliana, CRE1/AHK4 and AHK3, differ in their ligand specificity in a bacterial assay. Plant Cell Physiol. 45, 1299-1305.

Galuszka, P., Frébortová, J., Werner, T., Yamada, M., Strnad, M., Schmülling, T., Frébort, I. (2004) Cytokinin dehydrogenase gene family in cereals: cloning of functional barley CKX genes. Eur. J. Biochem. 271, 3990-4002.

Schmülling, T. (2004) Wachstums- und Ertragsverbesserung durch gezielte Veränderung des Cytokininkatabolismus. Vortr. Pflanzenz. 61, 90-93.


Werner, T., Motyka, V., Laucou, V., Smets, R., Van Onckelen, H., Schmülling, T. (2003) Cytokinin-deficient transgenic Arabidopsis plants show multiple developmental alterations indicating opposite functions of cytokinins in regulating shoot and root meristem activity. Plant Cell15, 2532-2550.

This article has been highlighted: ECKARDT, N.A. (2003) A new classic of cytokinin research: Cytokinin-deficient Arabidopsis plants provide new insights into cytokinin biology. Plant Cell 15, 2489-2492. https://doi.org/10.1105/tpc.151110

Heyl, A., Schmülling, T. (2003) Cytokinin signal perception and transduction. Curr. Opin. Plant Biol. 6, 480-488. (Review)

Schmülling, T., Werner, T., Riefler, M., Krupková, E., Bartrina y Manns, I. (2003) Structure and function of cytokinin oxidase/dehydrogenase genes of maize, rice, Arabidopsis and other species. J. Plant Res. 21, 40-49. (Review)

Motyka, V., Čapková, V., Petrášek, J., Kamínek, M., Schmülling, T. (2003) Cytokinin-induced upregulation of the cytokinin oxidase activity in tobacco includes changes in enzyme glycosylation and excretion indicating an extracellular degradation pathway for cytokinins. Physiol. Plant. 117, 11-20.

Schmülling, T. (2003) Cytokinin als wichtiger Steuerfaktor für Ertrag und Biomasseverteilung. Vortr. Pflanzenz. 56, 93-99. (Review)

Schmülling, T. (2003) Metabolismus und Signalübertragung der Cytokinine. Biospektrum 9, 256-259. (Review)

Tarkowská, D., Doležal, K., Tarkowski, P., Åstot, C., Schmülling, T., Holub, J., Fuksová, K., Sandberg, G., Strnad, M. (2003) Identification of new aromatic cytokinins in Arabidopsis thaliana and poplar leaves by LC-(+)ESI-MS and capillary liquid chromatography/frit - fast atom bombardment mass spectrometry. Physiol. Plant. 117, 579-590.

Schmülling, T., Werner, T., Riefler, M., Krupková, E., Bartrina y Manns, I., Dortay, H. (2003) Cytokinin as a regulatory factor for yield and biomass distribution in crop plants. In: Phytohormones in Plant Biotechnology and Agriculture (eds. Machacková, I., Romanov, G. A.), Kluwer Academic Publishers (The Netherlands), Dordrecht/Boston/London, 97-108. (Review)

Romanov, G. A., Getman, I. A., Bolyakina, Y. P., Rakova, N. Y., Kieber, J. J., Schmülling, T. (2003) Primary alcohols, substrates for phospholipase D-catalyzed transphosphatidylation, suppress cytokinin action. In: Phytohormones in Plant Biotechnology and Agriculture (eds. Machacková, I., Romanov, G. A.), Kluwer Academic Publishers (The Netherlands), Dordrecht/Boston/London, 129-139.

Werner, T., Hanuš, J., Holub, J., Schmülling, T., Van Onckelen, H., Strnad, M. (2003) New cytokinin metabolites in IPT transgenic Arabidopsis thaliana plants. Physiol. Plant. 118, 127-137.


Frank, M., Guivarc'h, A., Krupková, E., Lorenz-Meyer, I., Chriqui, D., Schmülling, T. (2002) The Tumorous Shoot Development (TSD) genes are required for co-ordinated plant shoot development. Plant J. 29, 73-85.

Schäfer, S., Schmülling, T. (2002) CRK1, a novel plant receptor kinase that is regulated by cytokinin. Plant Mol. Biol. 50, 155-166.

Romanov, G.A., Kieber, J., Schmülling, T. (2002) A rapid cytokinin-response assay in Arabidopsis indicates a role for phospholipase D in cytokinin signaling. FEBS Lett. 515, 39-43.

Schmülling, T. (2002) New insights into the functions of cytokinins in plant development. Plant Growth Reg. 21, 40-49. (Review)

Guivarc'h, A., Rembur, J., Noin, M., Schmülling, T., Chriqui, D. (2002) Localized expression of the ipt gene in the axillary buds of transgenic tobacco (Nicotiana tabacum L. cv. SR1) establishes a role for cytokinins in tuberization. J. Exp. Bot. 53, 621-629.

Frébort, I., Šebela, M., Galuszka, P., Werner, T., Schmülling, T., Peč, P. (2002) Cytokinin oxidase/cytokinin dehydrogenase assay: optimized procedures and applications. Anal. Biochem. 306, 1-7.


Werner, T., Motyka, V., Strnad, M., Schmülling, T. (2001) Regulation of plant growth by cytokinins. Proc. Natl. Acad. Sci. USA 98, 10487-10492.

Schmülling, T. (2001) CREam of cytokinin signalling: receptor identified. Trends Plant Sci. 6, 281-284. (Review)


Frank, M., Rupp, H.M., Prinsen, E., Motyka, M., Van Onckelen, H., Schmülling, T. (2000) Hormone autotrophic growth and differentiation identifies mutant lines of Arabidopsis thaliana with altered cytokinin and auxin content or signalling. Plant Physiol. 122, 721-729.

Lemcke, K., Prinsen, E., Van Onckelen, H., Schmülling, T. (2000) The ORF8 of Agrobacterium rhizogenes TL-DNA has tryptophane monooxygenase activity. Mol. Plant-Microbe Interact. 13, 787-790.

Schäfer, S., Krolzik, S., Romanov, G., Schmülling, T. (2000) Cytokinin-regulated transcripts in tobacco cell culture. J. Plant Growth Reg. 32, 307-313.

Romanov, G., Getman, I.A., Schmülling, T. (2000) Investigation of early cytokinin effects in a rapid Amaranthus seedling test. J. Plant Growth Reg. 32, 337-344.

Schmülling, T. (2000, 2001) Book reviews. J. Plant Physiol., various editions.


Rupp, H.M., Frank, M., Werner, T., Strnad, M., Schmülling, T. (1999) Increased steady state mRNA levels of the STM and KNAT1 homeobox genes in cytokinin overproducing Arabidopsis thaliana indicate a role for cytokinins in the shoot apical meristem. Plant J. 18, 357-363.

Frank, M., Schmülling, T. (1999) Cytokinin cycles cells. Trends Plant Sci. 4, 243-244. (Review)

Schmülling, T., Rupp, H.M., Frank, M., Schäfer, S. (1999) Recent advances in cytokinin research: Receptor candidates, primary response genes, mutants, transgenic plants. In: Strnad, M., Pec, P., Beck, E. (Eds.) Advances in regulation of plant growth and development. Peres Publ., Prag, 85–96. (Review)

Romanov, G., Getman, I.A., Schmülling, T. (1999) Rapid transcriptional activation of nuclear genes is necessary for cytokinin-induced synthesis of betacyanin in amaranthus. Doklady Akademii Nauk. (Proc. Acad. Sci., Moscow) 365, 832-835.


Lemcke, K., Schmülling, T. (1998) Gain of function assays identify non-rol genes from Agrobacterium rhizogenes TL-DNA that alter plant morphogenesis or hormone sensitivity. Plant J. 15, 423-434.

Lemcke, K., Schmülling, T. (1998) A putative rolB gene homolog of the Agrobacterium rhizogenes TR-DNA has different morphogenetic activity in tobacco than rolB. Plant Mol. Biol. 36, 803-808.

Moritz, T., Schmülling, T. (1998) The gibberellin content of rolA transgenic tobacco plants is specifically altered. J. Plant. Physiol. 153, 774-776.

Romanov, G.A., Konstantinova, T.N., Sergeeva, L.I., Golyanovskaya, S.A., Kossmann, J., Willmitzer, L., Schmülling, T., Aksenova, N.P. (1998) Morphology and tuber formation of in vitro grown potato plants harboring the yeast invertase gene and/or the rolC gene. Plant Cell Rep. 18, 318-324.

Reutter, K., Atzorn, R., Hadeler, B., Schmülling, T., Reski, R. (1998) Expression of the bacterial ipt gene in Physcomitrella rescues mutations in budding and in plastid division. Planta 206, 196-203.

Schmülling, T. (1998) Metabolism and development of cytokinin-overproducing plants. In: Metabolism and mode of action of plant hormones. Proc. VIth Symp. Span. Soc. Plant Physiol. 3-7. (Review)

Schmülling, T. (1998) Cytokinins: Light at the end of the tunnel? In: Novel approaches to study plant growth factors. Ed. Instituto Juan March de Estudios e Investigaciones, 87, 14-16. (Review)


Faiss, M., Zalubilová, J., Strnad, M., Schmülling, T. (1997) Conditional transgenic expression of the ipt gene indicates for cytokinins a function in paracrine signaling in whole tobacco plants. Plant J. 12, 401-415.

Schmülling, T., Schäfer, S., Romanov, G. (1997) Cytokinins as regulators of gene expression. Physiol. Plant. 100, 505-519. (Review)


Faiss, M., Strnad, M., Redig, P., Dolezal, K., Hanus, J., Van Onckelen, H., Schmülling, T. (1996) Chemically induced expression of the rolC encoded β-glucosidase in transgenic tobacco plants and analysis of cytokinin metabolism: rolC does not hydrolyze endogenous cytokinin glucosides in planta. Plant J. 10, 33-46.

Motyka, V., Faiss, M., Strnad, M., Kamínek, M., Schmülling, T. (1996) Changes in cytokinin content and cytokinin oxidase activity in response to derepression of ipt gene transcription in transgenic tobacco calli and plants. Plant Physiol. 112, 1035-1043.

Schmülling, T. (1996) Cytokinins - still a challenge in plant biology. Plantnet 3, 4-6. (Review)

Redig, P., Schmülling, T., Van Onckelen, H. (1996) Analysis of cytokinin metabolism in ipt transgenic tobacco by LC-tandem mass spectometry. Plant Physiol. 112, 141-148.


Schmülling, T., Röhrig, H. (1995) Gene silencing in transgenic hybrid tobacco plants: Frequency of the event and visualization of somatic inactivation pattern. Mol. Gen. Genet. 249, 375-390.

Grossmann, K., Schmülling, T. (1995) The effects of the herbicide quinchlorac on shoot growth in tomato is alleviated by inhibitors of ethylene biosynthesis and by the presence of an antisense construct to the 1-aminocyclopropane-1-carboxylic acid (ACC) synthase gene in transgenic plants. Plant Growth Reg. 16, 183-188.


Hewelt, A., Prinsen, E., Schell, J., Van Onckelen, H., Schmülling, T. (1994) Promoter tagging with a promoterless ipt gene leads to cytokinin-induced phenotypic variability in transgenic tobacco plants: implications of gene dosage effects. Plant J. 6, 879-892.

Röder, F.T., Schmülling, T., Gatz, C. (1994) Efficiency of the tetracycline-dependent gene expression system: complete suppression and efficient induction of the rolB phenotype in transgenic plants. Mol. Gen. Genet. 243, 32-38.


Schmülling, T., Fladung, M., Großmann, K., Schell, J. (1993) Hormonal content and sensitivity of transgenic tobacco and potato plants expressing single rol genes of Agrobacterium rhizogenes T-DNA. Plant J. 3, 371-382.

Fladung, M., Ballvora, A., Schmülling, T. (1993) Constitutive or light regulated expression of the rolC gene in transgenic potato plants alters yield attributes and tuber carbohydrate composition. Plant Mol. Biol. 23, 749-757.

Dehio, C., Grossmann, K., Schell, J., Schmülling, T. (1993) Phenotype and hormonal status of transgenic tobacco plants overexpressing the rolA gene of Agrobacterium rhizogenes T-DNA. Plant Mol. Biol. 23, 1199-1210.

Schmülling, T., Schell, J. (1993) Transgenic tobacco plants regenerated from leaf disks can be periclinal chimeras. Plant Mol. Biol. 21, 705-708.

Schmülling, T., Röhrig, H., Pilz, S., Walden, R., Schell, J. (1993) Restoration of fertility by antisense RNA in genetically engineered male sterile tobacco plants. Mol. Gen. Genet. 237, 385-394.

Nilsson, O., Crozier, A., Schmülling, T., Sandberg, G., Olsson, O. (1993) Indole-3-acetic acid homeostasis in transgenic tobacco plants expressing the Agrobacterium rhizogenes rolB gene. Plant J. 3, 681-689.

Walden, R., Czaja, I., Schmülling, T., Schell, J. (1993) Rol genes alter hormonal requirements for protoplast growth and modify the expression of an auxin responsive promoter. Plant Cell Rep. 12, 551-554.


Schmülling, T., Schell, J., Spena, A. (1992) Construction of cytokinin overproducing mutants in higher plants. In: Kaminek, M., Mok, D.W.S., Zazimalová, E. (Eds.) Physiology and biochemistry of cytokinins in plants. Academic Publishing, The Hague NL, 87-94.


Estruch, J.J., Parets-Soler, A., Schmülling, T., Spena, A. (1991) Cytosolic localization in transgenic plants of the rolC peptide from Agrobacterium rhizogenes. Plant Mol. Biol. 17, 547-550.

Koncz, C., Schmülling, T., Spena, A., Schell, J. (1991) T-DNA gene-functions. In: Herrmann, R.G., Larkins, B. (Eds.) Plant Molecular Biology 2. Plenum Press, New York, 205-209. (Review)


Schmülling, T., Schell, J., Spena, A. (1990) Altering plant morphogenesis by plant genetic engineering. Proc. 3rd Span. Congr. Biotechn. 131-135. (Review)


Schmülling, T., Schell, J., Spena, A. (1989) Promoters of the rolA, B, and C genes are differentially regulated in transgenic plants. Plant Cell 1, 665-670.

Schmülling, T., Beinsberger, S., De Greef, J., Schell, J., Van Onckelen, H., Spena, A. (1989) Construction of a heat-inducible chimeric gene to increase the cytokinin content in transgenic plant tissue. FEBS Lett. 289, 401-406.


Schmülling, T., Schell, J., Spena, A. (1988) Single genes from Agrobacterium rhizogenes stimulate abnormal growth in transgenic plants. EMBO J. 7, 2621-2629.


Spena, A., Schmülling, T., Koncz, C., Schell, J. (1987) Independent and synergistic activities of rolA, B, and C loci in stimulating abnormal growth in plants. EMBO J. 6, 3891-3899.


Papen, H., Kentemich, T., Schmülling, T., Bothe, H. (1986) Hydrogenase activities in cyanobacteria. Biochimie 68, 121-132.

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