Publications
Published journal articles and book chapters
Bartrina, I., Werner, S., Schenke, A., Gasperini, D., Werner, T., and Schmülling, T. (2025), Root-derived cytokinin regulates Arabidopsis flowering time through components of the age pathway. Plant Physiology, Volume 198, Issue 3, July 2025, kiaf204, https://doi.org/10.1093/plphys/kiaf204
Ren, Y., Zhang, Z., Zhanakhmetova, D., Li, W., Chen, S., Werner, T. and Liesche, J. (2024), Fast and simple fluorometric measurement of phloem loading exposes auxin-dependent regulation of Arabidopsis sucrose transporter AtSUC2. Plant J. https://doi.org/10.1111/tpj.17110
Barr Z.K., Werner T., Tilsner J. (2023), Heavy Metal-Associated Isoprenylated Plant Proteins (HIPPs) at Plasmodesmata: Exploring the Link between Localization and Function. Plants 12(16):3015. https://doi.org/10.3390/plants12163015
Górska A.M., Bartrina I., Werner T. (2023), Biomolecular condensation: a new player in auxin signaling. Trends in Plant Science 28: 620-622. https://doi.org/10.1016/j.tplants.2023.03.002
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., Zsögön A. and Peres L. E. P. (2022), Increased branching independent of strigolactone in CYTOKININ OXIDASE 2-overexpressing tomato is mediated by reduced auxin transport. Mol. Horticulture 2, no: 12 https://doi.org/10.1186/s43897-022-00032-1
Werner T. and Schmülling T. (2022), Moonlighting PPKL1 reveals a role of cytokinin in regulating rice grain size. Mol. Plant, Vol. 15, Issue 2, pp 216-218, https://doi.org/10.1016/j.molp.2021.11.014
Guo T, Weber H., Niemann M.C.E., Theisl L., Leonte G., Novák O., Werner T. (2021), Arabidopsis HIPP proteins regulate endoplasmic reticulum-associated degradation of CKX proteins and cytokinin responses. Mol. Plant, https://doi.org/10.1016/j.molp.2021.07.015
Werner S., Bartrina I., Novák O., Strnad M., Werner T. and Schmülling T. (2021), The cytokinin status of the epidermis regulates aspects of vegetative and reproductive development in Arabidopsis thaliana. Front. Plant Sci., Vol. 12, pp 95, https://doi.org/10.3389/fpls.2021.613488
Schwarz I., Scheirlinck M.-T., Otto E., Bartrina I., Schmidt R.-Ch., Schmülling T. (2020), Cytokinin regulates the activity of the inflorescence meristem and components of seed yield in oilseed rape. J. Exp. Bot., Vol. 71, No. 22 pp. 7146–7159, https://doi.org/10.1093/jxb/eraa419
Niemann M.C.E., Weber H., Hluska T., Leonte G., Anderson S.M., Novák O., Senes A., Werner T. (2018) The cytokinin oxidase/dehydrogenase CKX1 is a membrane-bound protein requiring homooligomerization in the endoplasmic reticulum for its cellular activity. Plant Physiol. 176: 2024-2039; https://doi.org/10.1104/pp.17.00925
Heyl, A., Werner, T. and Schmülling, T. (2018). Cytokinin Metabolism and Signal Transduction. In Annual Plant Reviews online, J.A. Roberts (Ed.). doi.org/10.1002/9781119312994.apr0245
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; https://doi.org/10.1111/tpj.13647
Bartrina I., Jensen H., Novák 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.
Niemann M.C.E., Bartrina I., Ashikov A., Weber H., Novák O., Spíchal L., Strnad M., Strasser R., Bakker H., Schmülling T., Werner T. (2015) Arabidopsis ROCK1 transports UDP-GlcNAc/UDP-GalNAc and regulates ER protein quality control and cytokinin activity. Proc. Natl. Acad. Sci. USA 112: 291-296.
Niemann M.C.E., Werner T. (2015) Endoplasmic reticulum: Where nucleotide sugar transport meets cytokinin control mechanisms. Plant Signal. Behav. 10: e1072668.
Köllmer I., Novák O., Strnad M., 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.
Macková H., Hronková M., Dobrá J., Turečková V., Novák O., Lubovská Z., Motyka V., Haisel D., Hájek T., Prášil I.T., Gaudinová A., Štorchová H., Ge E., Werner T., Schmülling T., Vanková 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.
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 crosstalk between pathways that control plant organ size in Arabidopsis. Plant Physiol. 155: 1339-1352.
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.
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.P. (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.
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.
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.
Werner T., Schmülling T. (2009) Cytokinin action in plant development. Curr. Opin. Plant Biol. 12: 527-538.
Werner T., Holst K., Pors Y., Guivarc'h A., Mustroph A., Chriqui D., Grimm B., Schmulling T. (2008) Cytokinin deficiency causes distinct changes of sink and source parameters in tobacco shoots and roots. J. Exp. Bot. 59: 2659-2672.
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. Plantarum 51: 673-682.
Galuszka P., Popelková H., Werner T., Frébortová J., Pospísilová H., Mik V., Köllmer I., Schmülling T., Frébort I. (2007) Biochemical characterization of cytokinin oxidases/dehydrogenases from Arabidopsis thaliana expressed in Nicotiana tabacum L. J. Plant Growth Regul. 26: 255-267.
Riefler M., Heyl A., Werner T. (2007) Cytokinins. In Encyclopedia of life sciences (John Wiley & Sons, Ltd: Chichester (DOI: 10.1002/9780470015902.a0020095).
Werner T., Köllmer I., Bartrina I., Holst K., Schmülling T. (2006) New insights into the biology of cytokinin degradation. Plant Biol. 8: 1-12.
Heyl A., Werner T., Schmülling T. (2006) Metabolism and signal transduction of cytokinin. In Plant hormone signaling, Annual Plant Reviews, P. Hedden and S. Thomas, eds (Oxford: Blackwell Publishing Ltd.), pp. 93-123.
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, C.J. Li, ed (Beijing: Tsinghua University Press), pp. 474-475.
Galuszka P., Frébortová J., Werner T., Yamada M., Strnad M., Schmülling T., Frébort I. (2004) Cytokinin oxidase/dehydrogenase genes in barley and wheat: cloning and heterologous expression. Eur. J. Biochem. 271: 3990-4002.
Werner T., Hanus 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.
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 the regulation of shoot and root meristem activity. Plant Cell 15: 2532-2550.
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. 116: 241-252.
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, I. Machácková and G.A. Romanov, eds (Dordrecht/Boston/London: Kluwer Academic Publishers), pp. 97-108.
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 cytokinin. Proc. Natl. Acad. Sci. USA 98: 10487-10492.
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: 557-563.