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Publications of the Department of Plant Biochemistry


Ubbiali D, Fratini M, Piersimoni L, Ihling C, Kipping M, Heilmann I, Iacobucci C, Sinz A (2022) Direct observation of 'elongated' conformational states in α-synuclein upon liquid-liquid phase separation. Angew Chem (in print)

Heilmann I (2022) Swap, combine and substitute to unravel specific functions of Arabidopsis PI4P 5-kinases. Plant Cell Physiol 63: 576-579

Heilmann M, Heilmann I (2022) Regulators regulated: Different layers of control for plasma membrane phosphoinositides in plants. Curr Opin Plant Biol 67:102218


Launhardt L, Matzner M, Heilmann M, Heilmann I (2021) Analysis of phosphoinositides from complex plant samples by solid-phase adsorption chromatography and subsequent quantification via thin-layer and gas chromatography. Methods Mol Biol 2295:379-389

Fratini M, Krishnamoorthy P, Stenzel I, Riechmann M, Matzner M, Bacia K, Heilmann M, Heilmann I (2021) Plasma membrane nano-organization specifies phosphoinositide effects on Rho-GTPases and actin dynamics in tobacco pollen tubes. Plant Cell 33: 642-670

Rausche J, Stenzel I, Stauder R, Fratini M, Trujillo M, Heilmann I, Rosahl S (2021) A phosphoinositide 5-phosphatase from Solanum tuberosum is activated by PAMP-treatment and may antagonize phosphatidylinositol 4,5-bisphosphate at Phytophthora infestans infection sites. New Phytol 229: 469-487

Ayash M, Abukhalaf M, Thieme D, Proksch C, Heilmann M, Schattat MH, Hoehenwarter W (2021) LC-MS based draft map of the Arabidopsis thaliana nuclear proteome and protein import in pattern triggered immunity. Front Plant Sci 12: 744103-744103


Zarza X, Van Wijk R, Shabala L, Hunkeler A, Lefebvre M, Rodriguez-Villalón A, Shabala S, Tiburcio AF, Heilmann I, Munnik T (2020) Lipid kinases PIP5K7 and PIP5K9 are required for polyamine-triggered K+ efflux in Arabidopsis roots. Plant J 104: 416-432

Hurný A, Cuesta C, Cavallari N, Ötvös K, Duclercq J, Dokladal L, Montesinos JC, Gallemí M, Semerádová H, Rauter T, Stenzel I, Persiau G, Benade F, Bhalearo R, Sykorova E, Gorzsás A, Sechet J, Mouille G, Heilmann I, De Jaeger G, Ludwig-Müller J, Benková E (2020) SYNERGISTIC ON AUXIN AND CYTOKININ 1 positively regulates growth and attenuates soil pathogen resistance. Nat Commun 11: 2170

Stenzel I, Ischebeck T, Vu-Becker L, Riechmann M, Krishnamoorthy P, Fratini M, Heilmann I (2020) Coordinated localization and antagonistic function of NtPLC3 and PI4P 5-kinases in the subapical plasma membrane of tobacco pollen tubes. Plants 9: e452


Menzel W, Stenzel I, Helbig LM, Krishnamoorthy P, Neumann S, Eschen-Lippold L, Heilmann M, Lee J, Heilmann I (2019) A PAMP-triggered MAPK-cascade inhibits phosphatidylinositol 4,5-bisphosphate production by PIP5K6 in Arabidopsis thaliana. New Phytol 224: 833-847

Erwig MS, Patzig J, Steyer AM, Dibaj P, Heilmann M, Heilmann I, Jung RB, Kusch K, Möbius W, Jahn O, Nave KA, Werner HB (2019) Anillin facilitates septin assembly to prevent pathological outfoldings of central nervous system myelin. Elife 8: e43888

Vaz Dias F, Serrazina S, Vitorino M, Marchese D, Heilmann I, Godinho M, Rodrigues M, Malhó R (2019) A role for diacylglycerol kinase 4 in signalling crosstalk during Arabidopsis pollen tube growth. New Phytol 222: 1434-1446

Lin F, Krishnamoorthy P, Schubert V, Hause G, Heilmann M, Heilmann I (2019) A dual role for cell plate-associated PI4Kβ in endocytosis and phragmoplast dynamics during plant somatic cytokinesis. EMBO J 38: e100303


Strobl SM, Kischka D, Heilmann I, Mouille G, Schneider S (2018) The tonoplastic inositol transporter INT1 from Arabidopsis thaliana impacts cell elongation in a sucrose-dependent way. Front Plant Sci 9: 1657

Jablonická V, Ziegler J, Vatehová Z, Lišková D, Heilmann I, Obložinský M, Heilmann M (2018) Inhibition of phospholipases influences the metabolism of wound-induced benzylisoquinoline alkaloids in Papaver somniferum L. J Plant Physiol 223: 1-8


Hempel F, Stenzel I, Heilmann M, Krishnamoorthy P, Menzel W, Golbik R, Helm S, Dobritzsch D, Baginsky S, Lee J, Hoehenwarter W, Heilmann I (2017) MAPKs Influence Pollen Tube Growth by Controlling the Formation of Phosphatidylinositol 4,5-Bisphosphate in an Apical Plasma Membrane Domain. Plant Cell 29: 3030-3050

Gerth K, Lin F, Daamen F, Menzel W, Heinrich F, Heilmann M (2017) Arabidopsis phosphatidylinositol 4-phosphate 5-kinase 2 contains a functional nuclear localization sequence and interacts with alpha-importins. Plant J 92: 862-878

Geilen K, Heilmann M, Hillmer S, Böhmer M (2017) WRKY43 regulates polyunsaturated fatty acid content and seed germination under unfavourable growth conditions. Sci Rep 7: 14235

Gerth K, Lin F, Menzel W, Krishnamoorthy P, Stenzel I, Heilmann M, Heilmann I (2017) Guilt by Association: A Phenotype-Based View of the Plant Phosphoinositide Network. Annu Rev Plant Biol 68: 349-374


Barbosa ICR, Shikata M, Zourelidou M, Heilmann M, Heilmann I, Schwechheimer C (2016) Phospholipid-dependent recruitment of D6 PROTEIN KINASE to the plasma membrane is mediated by a K-rich motif and required for auxin transport-dependent tropisms. Development 143: 4687-4700

Jablonická V, Mansfeld J, Heilmann I, Obložinský M, Heilmann M (2016) Identification of a secretory phospholipase A2 from Papaver somniferum L. that transforms phospholipids Phytochemistry 129: 4-13

Heilmann I (2016) Phosphoinositide signalling and plant development. Development 142: 2044-2055

Dejonghe W, Kuenen S, Mylle E, Vasileva M, Keech O, Viotti C, Swerts J, Fendrych M, Ortiz-Morea FA, Mishev K, Delang S, Scholl S, Zarza X, Heilmann M, Kourelis J, Kasprowicz J, Nguyen LSL, Drozdzecki A, Van Houtte I, Szatmári A-M, Majda M, Baisa G, Bednarek SY, Robert S, Audenaert D, Testerink C, Munnik T, Van Damme C, Heilmann I, Schumacher K, Winne J, Friml J, Verstreken P, Russinova E (2016) Mitochondrial uncouplers inhibit clathrin-mediated endocytosis largely through cytoplasmic acidification. Nat Commun 7: 11710

Heilmann I (2016) Plant phosphoinositides - dynamics on demand. Biochim Biophys Acta 1861: 1345-1351

Iven T, Hornung E, Heilmann M, Feussner I (2016) Synthesis of oleyl oleate wax esters in Arabidopsis thaliana and Camelina sativa seed oil. Plant Biotechnol J 14: 252-259


Heilmann I, Ischebeck T (2015) Male functions and malfunctions: the impact of phosphoinositides on pollen development and pollen tube growth. Plant Reprod 29: 3-20

Saavedra L, Catarino R, Heinz T, Heilmann I, Bezanilla M, Malhó R (2015) Phosphatase and tensin homolog (PTEN) is a growth repressor of both rhizoid and gametophore development in the moss Physcomitrella patens. Plant Physiol 169: 2572-2586

Steinhorst L, Mähs, A, Ischebeck T, Zhang C, Eckert C, Zhang X, Arendt S, Schültke S, Heilmann I, Kudla J (2015) Vacuolar CBL-CIPK12 Ca2+ sensor/kinase complexes are required for polarized cell growth. Curr Biol 25: 1475-1482

Heilmann I (2015) Information processing and survival strategies. In Ecological Biochemistry: Environmental and Interspecies Interactions, Krauss G-J, Nies DH, eds., Wiley-VCH, Weinheim, pp123-153

Kaever A, Landesfeind M, Feussner K, Mosblech A, Heilmann I, Morgenstern B, Feussner I, Meinicke P (2015) MarVis-Pathway: Integrative and exploratory pathway analysis of non-targeted metabolomics data. Metabolomics 11: 764-777

Heilmann M, Heilmann I (2015) Plant phosphoinositides - complex networks controlling growth and adaptation. Biochim Biophys Acta 185: 759-769


Tejos R, Sauer M, Vanneste S, Palacios-Gomez M, Li H, Heilmann M, van Wijk R, Vermeer J, Heilmann I, Munnik T, Friml J (2014) Bipolar plasma membrane distribution of phosphoinositides and their requirement for auxin-mediated cell polarity and patterning in Arabidopsis. Plant Cell 26: 2114-2128

Krishnamoorthy P, Sanchez-Rodriguez C, Heilmann I, Persson, S (2014) Regulatory roles of phosphoinositides in membrane trafficking and their potential impact on cell wall synthesis and remodeling. Ann Bot 114: 1049-1057

Novakova P, Hirsch S, Feraru E, Tejos R, van Wijk R, Viaene T, Heilmann M, Lerche J, De Rycke R, Feraru M, Grones P, Van Montagu MCE, Heilmann I, Munnik T, Friml J (2014) SAC domain phosphoinositide phosphatases at the tonoplast mediate vacuolar function in Arabidopsis. Proc Natl Acad Sci USA 111: 2818-2823

Eckert C, Offenborn JN, Heinz T, Schültke S, Zhang C, Hillmer S , Gold S, Heilmann M, Schumacher K, Bock R, Heilmann I, Kudla J (2014) The vacuolar calcium sensors CBL2 and CBL3 control seed size and embryonal development in Arabidopsis thaliana. Plant J 78: 146-156

Mansfeld J, Schöpfel M, Lorenz J, Trutschel T, Heilmann I, Brandt W, Ulbrich-Hofmann R (2014) Probing selected structural regions in the plant secreted phospholipase A2 from Arabidopsis thaliana for their impact on stability and activity. Biochimie 101: 60-66


Ischebeck T, Werner S, Krishnamoorthy P, Lerche J, Meijón M, Stenzel I, Löfke C, Wiessner T, Im YJ, Perera IY, Iven T, Feussner I, Busch W, Boss WF, Teichmann T, Hause B, Persson S, Heilmann I (2013) Phosphatidylinositol 4,5-bisphosphate influences PIN polarization by controlling clathrin mediated membrane trafficking. Plant Cell 25: 4894-4911

Obložinský M, Bezáková L, Mansfeld J, Heilmann I, Ulbrich-Hofmann, R (2013) Differences in the effect of phosphatidylinositol 4,5-bisphosphate on the hydrolytic and transphosphatidylation activities of membrane-bound phospholipase D from poppy seedlings. Plant Physiol Biochem 69: 39-42

Iven T, Herrfurth C, Hornung E, Heilmann M, Hofvander P, Stymne S, Zhu LH, Feussner I (2013) Wax ester profiling of seed oil by nano-electrospray ionization tandem mass spectrometry. Plant Methods 9: 24

Heilmann M, Heilmann I (2013) Arranged marriage in lipid signalling? The limited choices of PtdIns(4,5)P2 in finding the right partner. Plant Biol (Stuttg.) 15: 789-797

Löfke C, Zwiewka M, Heilmann I, van Montagu MCE, Teichmann T, Friml J (2013) Asymmetric gibberellin signaling regulates vacuolar trafficking of PIN auxin transporters during root gravitropism. Proc Natl Acad Sci USA 110: 3627-3632

Munnik T, Heilmann I, eds. (2013) Plant Lipid Signaling Protocols, Methods Mol Biol 1009, Humana Press, Springer Acad Publ, ISBN-10 1627034005

Heilmann I, Perera IY (2013) Measurement of inositol (1,4,5) trisphosphate in plant tissues by a competitive receptor binding assay. Methods Mol Biol 1009: 33-41

Heilmann M, Heilmann I (2013) Mass measurement of polyphosphoinositides by thin-layer and gas chromatography. Methods Mol Biol 1009: 25-32


Heilmann M, Iven T, Ahmann K, Hornung E, Stymne S, Feussner I (2012) Production of wax esters in plant seed oils by oleosomal cotargeting of biosynthetic enzymes. J Lipid Res 53: 2153-2161

Mähs A, Ischebeck T, Heilig Y, Stenzel I, Hempel F, Seiler S, Heilmann I (2012) The essential phosphoinositide kinase MSS-4 is required for polar hyphal morphogenesis, localizing to sites of growth and cell fusion in Neurospora crassa. PLoS One 7: e51454

Stenzel I, Ischebeck T, Quint M, Heilmann I (2012) Variable regions of PI4P 5-kinases direct PtdIns(4,5)P2 towards alternative regulatory functions in tobacco pollen tubes. Front Plant Sci 2: 1-14

Eschen-Lippold L, Landgraf R, Smolka U, Schulze S, Heilmann M, Heilmann I, Hause G, Rosahl S (2012) Activation of defense against Phytophthora infestans in potato by down regulation of syntaxin gene expression. New Phytol 193: 985-996


Ternes P, Feussner K, Werner S, Lerche J, Heilmann I, Riezman H, Feussner I (2011) Disruption of the ceramide synthase LOH1 causes spontaneous cell death in Arabidopsis thaliana. New Phytol 192: 851-854

Saavedra L, Balbi V, Lerche J, Mikami K, Heilmann I, Sommarin M (2011) PIPKs are essential for rhizoid elongation and caulonemal cell development in the moss Physcomitrella patens. Plant J 67: 635-647

Mosblech A, Thurow C, Gatz C, Feussner I, Heilmann I (2011) Jasmonic acid perception by COI1 involves inositol polyphosphates in Arabidopsis thaliana. Plant J 65: 949-957

Ischebeck T, Stenzel I, Hempel F, Jin X, Mosblech A, Heilmann I (2011) Phosphatidylinositol-4,5-bisphosphate influences Nt-Rac5-mediated cell expansion in pollen tubes of Nicotiana tabacum. Plant J 65: 453-468

Im YJ, Heilmann I, Perera IY (2011) The Hull of Fame: Lipid Signaling in the Plasma Membrane. In The Plant Plasma Membrane, Plant Cell Monographs, Murphy AS, Peer W,Schulz B, eds. Springer Academic Publishers, Berlin/Heidelberg, pp. 437-456


Ischebeck T, Vu LH, Jin X, Stenzel I, Löfke C, Heilmann I (2010) Functional cooperation of enzymes of phosphoinositide conversion according to synergistic effects on pectin secretion in tobacco pollen tubes. Mol Plant 3: 870-881

Goebbels S, Oltrogge J, Kemper R, Heilmann I, Wichert S, Bormuth I, Wolfer S, Moebius W, Liu X, Lappe-Siefke C, Rossner M, Groszer M, Suter U, Frahm J, Boretius, S, Nave K-A (2010) Elevated phosphatidylinositol 3,4,5 trisphosphate in glia triggers cell autonomous membrane wrapping and myelination. J Neurosci 30: 8953-8964

Heilmann I, Im YJ, Dieck C, Perera IY, Boss WF (2010) "Plant Physiology Classics Collection" Online-Kommentar zu Müller-Röber B und Pical C (2002) Plant Physiol 130: 22-46

Heilmann I (2010) Phosphoinositide signaling in plants. The Lipid Library (Internetauftritt der American Oil Chemists’ Society, AOCS),   

Furt F, König S, Bessoule J-J, Sargueil F, Zallot R, Stanislas T, Noirot E, Lherminier J, Simon-Plas F, Heilmann I , Mongrand S (2010) Polyphosphoinositides are enriched in plant membrane rafts and form microdomains in the plasma membrane. Plant Physiol 152: 2173-2187

Ischebeck T, Seiler S, Heilmann I (2010) At the poles across kingdoms: phosphoinositides and polar tip growth. Protoplasma 240: 13-31

Ischebeck T, Heilmann I (2010) PIP-kinases as key regulators of plant function. In T Munnik, ed, Plant Lipid Signaling, Plant Cell Monographs. Springer Academic Publishers, Berlin/Heidelberg, pp. 79-93

Mosblech A, Feussner I, Heilmann I (2010) Oxylipin signaling and plant growth. In T Munnik, ed, Plant Lipid Signaling, Plant Cell Monographs. Springer Academic Publishers, Berlin/Heidelberg, pp. 277-291

Heilmann I, Shanklin J (2010) ADS genes for reducing saturated fatty acid levels in seed oils. US-patent No. 7655833 (patent application first published in 2005)


Bargmann BO, Laxalt AM, Riet B, Testerink C, Merquiol E, Mosblech A, Reyes AL, Pieterse CM, Haring MA, Heilmann I, Bartels D, Munnik T (2009) Reassessing the role of phospholipase D in the Arabidopsis wounding response. Plant Cell Environ 32: 837-850

Heilmann I (2009) Using genetic tools to understand plant phosphoinositide signaling. Trends Plant Sci 14: 171-179

Mosblech A, Feussner I, Heilmann I (2009) Oxylipins: Structurally diverse metabolites from fatty acid oxidation. Plant Physiol Biochem 47: 511-517


Ischebeck T, Stenzel I, Heilmann I (2008) Type B phosphatidylinositol-4-phosphate 5-kinases mediate polar tip growth of pollen tubes by regulating apical pectin secretion. Plant Cell 20: 3013-3030

Heilmann I (2008) Towards understanding the function of stress-inducible PtdIns(4,5)P2 in plants. Commun Integr Biol 1: 204-206

König S, Ischebeck T, Lerche J, Stenzel I, Heilmann I (2008) Salt-stress-induced association of phosphatidylinositol-4,5-bisphosphate with clathrin-coated vesicles in plants. Biochem J 415: 387-399

Löfke C, Ischebeck T, König S, Freitag S, Heilmann I (2008) Alternative metabolic fates of phosphatidylinositol produced by PI-synthase isoforms in Arabidopsis thaliana. Biochem J 413: 115-124

König S, Hoffmann M, Mosblech A, Heilmann I (2008) Determination of content and fatty acid composition of unlabeled phosphoinositide species by thin layer chromatography and gas chromatography. Anal Biochem 378: 387-399

Mosblech A, König S, Stenzel I, Grzeganek P, Feussner I, Heilmann I (2008) Phosphoinositide and inositolpolyphosphate-signalling in defence responses of Arabidopsis thaliana challenged by mechanical wounding. Mol Plant 1: 249-261

Stenzel I, Ischebeck T, König S, Holubowska A, Sporysz M, Hause B, Heilmann I (2008) The type B phosphatidylinositol-4-phosphate 5-kinase 3 is essential for root hair formation in Arabidopsis thaliana. Plant Cell 20: 124-141

Lang I, Göbel C, Porzel A, Heilmann I, Feussner I (2008) A lipoxygenase with linoleate diol synthase activity from Nostoc sp. PCC 7120. Biochem J 410: 347-357

Heilmann I (2008) Tails wagging the dogs - On phosphoinositides and their fatty acyl moieties. Plant Signal Behav 10: 768-771

Heilmann I (2008) Phosphoinositides - A tale of heads and tails. J Lip Sci Techn 110: 189-190


König S, Mosblech A, Heilmann I (2007) Stress-inducible and constitutive phosphoinositide pools have distinctive fatty acid patterns in Arabidopsis thaliana. FASEB J 21: 1958-1967

Pidkowich MS, Nguyen HT, Heilmann I, Ischebeck T, Shanklin J (2007) Modulating seed beta-ketoacyl-ACP synthase II level converts a temperate seed oil composition to that of a palm-like tropical oil. Proc Natl Acad Sci USA 104: 4742-4747

2006 -

König S, Heilmann I (2006) Distinct subcellular phosphoinositide pools in algae and plants. In Current Advances in the Biochemistry and Cell Biology of Plant Lipids. Benning C, Ohlrogge J, eds., Aardvark Global Publishing , LLC, Salt Lake City, UT, USA, pp 292-296

Heilmann I, Mekhedov S, King B, Browse J, Shanklin J (2004) Identification of the Arabidopsis palmitoyl-monogalactosyldiacylglycerol Δ7-desaturase gene FAD5, and effects of plastidial retargeting of Arabidopsis desaturases on the fad5 mutant phenotype. Plant Physiol 136: 4327-4345

Heilmann I, Pidkowich MS, Girke T, Shanklin J (2004) Switching desaturase enzyme specificity by alternate subcellular targeting. Proc Natl Acad Sci USA 101: 10266-10271

Heilmann I, Perera IY, Boss WF (2004) Phosphoinositide signaling, lipid pools and PI kinase isoforms in plants. In Proceedings of the XIIth International Congress on Genes, Gene Families and Isozymes. Schnarrenberger C, Wittman-Liebold B, eds., Monduzzi Editore, Bologna, Italy, pp 73-80

Perera IY, Love J, Heilmann I, Thompson WF, Boss WF (2002) Up-regulation of phosphoinositide metabolism in tobacco cells constitutively expressing the human type I inositol polyphosphate 5-phosphatase. Plant Physiol 129: 1795-1806

Perera IY, Heilmann I, Stevenson-Paulik JM, Boss WF (2002) The phosphoinositide (PI) pathway and signaling in plants. In Signal Transduction in Plants - Current Advances. Sopory SK, Oelmüller R, Maheswari SC, eds., Kluwer Acad. Publ., Netherlands, pp 83-92

Heilmann I, Shin J, Huang J, Perera IY, Davies E (2001) Transient dissociation of polyribosomes and concurrent recruitment of calreticulin and calmodulin transcripts in gravistimulated maize pulvini. Plant Physiol 127: 1193-1203

Heilmann I, Perera IY, Gross W, Boss WF (2001) Plasma membrane phosphatidylinositol 4,5-bisphosphate levels decrease with time in culture. Plant Physiol 126: 1507-1518

Gross W, Heilmann I, Lenze D, Schnarrenberger C (2001) Biogeography of the Cyanidiaceae (Rhodophyta) based on 18S ribosomal RNA gene sequence data. Eur J Phycol 36: 275-280

Heilmann I, Gross W (2001) Genetic diversity of thermo-acidophilic red algae according to random amplified polymorphic DNA (RAPD) analysis. In Algae and Extreme Environments. Elster J, Seckbach J, Vincent WF, Lhotsky O, eds., Gebr. Borntraeger, Berlin, Germany, pp 531-539

Perera IY, Heilmann I, Chang SC, Boss WF, Kaufman PB (2001) A role for inositol 1,4,5-trisphosphate in gravitropic signaling and the retention of cold-perceived gravistimulation of oat shoot pulvini. Plant Physiol 125: 1499-1507

Stevenson JM, Perera IY, Heilmann I, Persson S, Boss WF (2000) Inositol signaling and plant growth. Trends Plant Sci 5: 252-258

Heilmann I, Perera IY, Stevenson JM, Boss WF (2000) Sense and sensibility: Inositol phospholipids as mediators of abiotic stress responses. In Plant Tolerance to Abiotic Stresses in Agriculture: Role of Genetic Engineering. Cherry JH, Key JH, Rychter AM, eds., Kluwer Acad. Publ., Netherlands, pp 285-296

Heilmann I, Stevenson-Paulik J, Perera IY (2000) Plant PtdIns 3-kinase goes nuclear. Plant Cell 12: 1511-1512

Perera IY, Heilmann I, Boss WF (1999) Transient and sustained increases in inositol-1,4,5-trisphosphate precede the differential growth response in gravistimulated maize pulvini. Proc Natl Acad Sci USA 96: 5838-5843

Heilmann I, Perera IY, Gross W, Boss WF (1999) Changes in phosphoinositide metabolism with days in culture affect signal transduction pathways in Galdieria sulphuraria. Plant Physiol 119: 1331-1339

Heilmann I, Perera IY, Stevenson JM, Ransom WD, Gross W, Boss WF (1998) Inositol lipid signaling: What can we learn from plants? In Advances in Lipid Research. Sanchez J, Cerda-Olmedo E, Martinez-Force E, eds., Univ. Sevilla Press, Spain, pp 394-397

Heilmann I, Schnarrenberger C, Gross W (1997) Mannose metabolizing enzymes from the red alga Galdieria sulphuraria. Phytochemistry 45: 903-906

Klinker JF, Hagelüken A, Grünbaum L, Heilmann I, Nürnberg B, Harhammer R, Offermanns St, Ervens J, Wenzel-Seifert K, Müller T, Seifert R (1994) Mastoparan may activate GTP hydrolysis by Gi-proteins in HL-60 membranes indirectly through interaction with nucleoside diphosphate kinase. Biochem J 304: 377-383