Biomedical Research Education & Training
Faculty Member

Brown, H. Alex, Ph.D.
Professor of Pharmacology
Bixler-Johnson-Mayes Chair
Professor of Biochemistry

Lab Url:

Phone Number: 615-936-3888

Email Address:

Brown, H. Alex's picture
Academic history
B.S., Florida Institute of Technology
M.S., Syracuse University
Ph.D., University of North Carolina at Chapel Hill
Postdoc, University of Texas-Southwestern Medical Center

Office Address   Mailing Address

417A Preston Research Building

442 RRB Vanderbilt University Medical Center 37232-6600

Research Specialty
Research in Pharmacology, Biochemistry, Bioorganic, and Analytical Chemistry.

Research Description
Our research group is focused on understanding the roles of specific lipid molecular species and phospholipases in cellular functions and human disease. The laboratory is highly interdisciplinary with research faculty, postdoctoral fellows, staff and graduate students coming from a variety of backgrounds including chemistry, biochemistry, pharmacology, and mathematics. We combine systems biology approaches with mass spectrometry to profile changes in cellular lipid species (lipidomics). Much of our work is focused on understanding the role of phospholipase D (PLD) in growth factor and G protein coupled receptor (GPCR) signaling networks. With our collaborators we are developing novel chemical inhibitors of PLD and ancillaryproteins in the signaling network to better understand the role of PLD in cancers and tumorigenesis, as well as to explore its therapeutic potential. Areas of current research include: structure and enzymology of PLD; defining the signal transduction networks from the receptor tyrosine kinases and GPCRs to PLD; mechanisms by which phosphatidic acid (PA) regulates cell cycle, growth, and proliferation; roles of PLD in zebrafish; developing novel probes for defining specific lipid binding targets; precursor-product relationships of lipid enzymes using mass spectrometry; lipid peroxidation and antioxidant mechanisms; and advanced methods in lipid mass spectrometry.

A complete listing of our publications can be found at:

Mathews, TP, Hill, S, Rose, KL, Ivanova, PT, Lindsley, CW, Brown, HA. Human phospholipase D activity transiently regulates pyrimidine biosynthesis in malignant gliomas. ACS Chem Biol, 10(5), 1258-1268, 2015 PMID25646564

Warrier, M, Shih, DM, Burrows, AC, Ferguson, D, Gromovsky, AD, Brown, AL, Marshall, S, McDaniel, A, Schugar, RC, Wang, Z, Sacks, J, Rong, X, Vallim, TA, Chou, J, Ivanova, PT, Myers, DS, Brown, HA, Lee, RG, Crooke, RM, Graham, MJ, Liu, X, Parini, P, Tontonoz, P, Lusis, AJ, Hazen, SL, Temel, RE, Brown, JM. The TMAO-Generating Enzyme Flavin Monooxygenase 3 Is a Central Regulator of Cholesterol Balance. Cell Rep, 10(3), 326-338, 2015 PMID 25600868

Gorden, DL, Myers, DS, Ivanova, PT, Fahy, E, Maurya, MR, Gupta, S, Min, J, Spann, NJ, McDonald, JG, Kelly, SL, Duan, J, Sullards, MC, Leiker, TJ, Barkley, RM, Quehenberger, O, Armando, AM, Milne, SB, Mathews, TP, Armstrong, MD, Li, C, Melvin, WV, Clements, RH, Washington, MK, Mendonsa, AM, Witztum, JL, Guan, Z, Glass, CK, Murphy, RC, Dennis, EA, Merrill, AH, Russell, DW, Subramaniam, S, Brown, HA. Biomarkers of NAFLD progression : a lipidomics approach to an epidemic. J Lipid Res, 56(3), 722-736, 2015 PMC 4340319

Spencer, CT, Brown, HA. Biochemical Characterization of a Pseudomonas aeruginosa Phospholipase D. Biochemistry, 54(5), 1208-1218, 2015 PMC 4337821

Scott, SA, Spencer, CT, O'Reilly, MC, Brown, KA, Lavieri, RR, Cho, CH, Jung, DI, Larock, RC, Brown, HA, Lindsley, CW. Discovery of Desketoraloxifene Analogues as Inhibitors of Mammalian, Pseudomonas aeruginosa, and NAPE Phospholipase D Enzymes. ACS Chem Biol, 10(2), 421-432, 2015 PMC 4336625

Ivanova PT, Myers DS, Milne SB, McClaren JL, Thomas PG, Brown HA. Lipid composition of viral envelope of three strains of influenza virus-not all viruses are created equal. ACS Infectious Diseases, in press, 2015

McDonald, JG, Ivanova, PT, Brown, HA.. Approaches to lipid analysis. In "Biochemistry of Lipids, Lipoproteins, and Membranes" 6th ed, 2015

Taylor, HE, Simmons, GE, Mathews, TP, Khatua, AK, Popik, W., Lindsley, CW, D'Aquila, RT, Brown, HA.. Phospholipase D1 couples CD4+T cell activation to c-Myc-dependent deoxyribonucleotide pool expansion and HIV-1 replication.. PLoS Pathogens, 11(5), e1004846, 2015

Bruntz, RC, Taylor, HE, Lindsley, CW, Brown, HA. Phospholipase D2 mediates survival signaling through direct regulation of Akt in glioblastoma cells. J Biol Chem, 289(2), 600-616, 2014PMC 3887188*Article featured on the issue COVER and PAPERS OF THE WEEK. See first author profile at profile

O'Reilly, MC, Oguin, TH, Scott, SA, Thomas, PG, Locuson, CW, Morrison, RD, Daniels, JS, Brown, HA, Lindsley, CW. Discovery of a Highly Selective PLD2 Inhibitor (ML395): A New Probe with Improved Physiochemical Properties and Broad-Spectrum Antiviral Activity against Influenza Strains. ChemMedChem, 9(12), 2633-2637, 2014 PMID 25210004

Bruntz, RC, Lindsley, CW, Brown, HA. Phospholipase D Signaling Pathways and Phosphatidic Acid as Therapeutic Targets in Cancer. Pharmacol Rev, 66(4), 1033-1079, 2014 PMC 4180337

Oguin, TH, Sharma, S, Stuart, AD, Duan, S, Scott, SA, Jones, CK, Daniels, JS, Lindsley, CW, Thomas, PG, Brown, HA. Phospholipase D facilitates efficient entry of influenza virus allowing escape from innate mmune inhibition. J Biol Chem, 289(37), 25405-25417, 2014 PMC 4162146

Cipriano, R, Bryson, BL, Miskimen, KLS, Bartel, CA, Hernandez-Sanchez, W, Bruntz, RC, Scott, SA, Lindsley, CW, Brown, HA, Jackson, MW. Hyperactivation of EGFR and downstream effector phospholipase D1 by oncogenic FAM83B. Oncogene, 33(25), 3298-3306 PMC392847, 2014

Faust, JE, Manisundaram, A, Ivanova, PT, Milne, SB, Summerville, JB, Brown, HA, Wangler, M, Stern, M, McNew, JA. Peroxisomes Are Required for Lipid Metabolism and Muscle Function in Drosophila melanogaster. PLoS One, 9(6), e100213 PMC 4063865, 2014

Leamy, AK, Egnatchik, RA, Shiota, M, Ivanova, PT, Myers, DS, Brown, HA, Young, JD. Enhanced synthesis of saturated phospholipids is associated with ER stress and lipotoxicity in palmitate treated hepatic cells. J Lipid Res, 55(7), 1478-1488 PMC 4076085, 2014

O''Reilly, MC, Scott, SA, Brown, HA, Lindsley, CW. Further evaluation of novel structural modifications to scaffolds that engender PLD isoform selective inhibition. Bioorg Med Chem Lett, 24(24), 5553-7, 2014

Ren, J, Pei-Chen Lin, C, Pathak, MC, Temple, BR, Nile, AH, Mousley, CJ, Duncan, MC, Eckert, DM, Leiker, TJ, Ivanova, PT, Myers, DS, Murphy, RC, Brown, HA, Verdaasdonk, J, Bloom, KS, Ortlund, EA, Neiman, AM, Bankaitis, VA. A phosphatidylinositol transfer protein integrates phosphoinositide signaling with lipid droplet metabolism to regulate a developmental program of nutrient stress-induced membrane biogenesis. Mol Biol Cell, 25(5), 712-27 PMC3937096, 2014

Scott, SA, Mathews, TP, Ivanova, PT, Lindsley, CW, Brown, HA. Chemical modulation of glycerolipid signaling and metabolic pathways. Biochim Biophys Acta, 1841(8), 1060-1084 PMC 4069240, 2014

Shulga, YV, Loukov, D, Ivanova, PT, Milne, SB, Myers, DS, Hatch, GM, Umeh, G, Jalan, D, Fullerton, MD, Steinberg, GR, Topham, MK, Brown, HA, Epand, RM. Diacylglycerol Kinase Delta Promotes Lipogenesis. Biochemistry, 52(44), 7766-7776, 2013 PMC 3864584

Shibata, N, Carlin, AF, Spann, NJ, Saijo, K, Morello, CS, McDonald, JG, Romanoski, C, Maurya, MR, Kaikkonen, M, Lam, M, Crotti, A, Reichart, D, Fox, JN, Quehenberger, O, Raetz, CR, Sullards, MC, Murphy, RC, Merrill, AH, Brown, HA, Dennis, EA, Fahy, E, Subramaniam, S, Cavener, DR, Spector, DH, Russell, DW, Glass, CK. 25-hydroxycholesterol activates the Integrated Stress Response to reprogram transcription and translation in macrophages. J Biol Chem, 288(50), 35812-35823, 2013 PMC 3861632

Thomas, G, Betters, JL, Lord, CC, Brown, AL, Marshall, S, Ferguson, D, Sawyer, J, Davis, MA, Melchior, JT, Blume, LC, Howlett, AC, Ivanova, PT, Milne, SB, Myers, DS, Mrak, I, Leber, V, Heier, C, Taschler, U, Blankman, JL, Cravatt, BF, Lee, RG, Crooke, RM, Graham, MJ, Zimmermann, R, Brown, HA, Brown, JM. The Serine Hydrolase ABHD6 Is a Critical Regulator of the Metabolic Syndrome. Cell Rep, 5(2), 508-520, 2013 PMC 3833083

Meszaros, P, Klappe, K, van Dam, A, Ivanova, PT, Milne, SB, Myers, DS, Brown, HA, Permentier, H, Hoekstra, D, Kok, JW. Long term myriocin treatment increases MRP1 transport activity. Int J Biochem Cell Biol, 45(2), 326-34, 2013 PMC 3826170

Maurya, MR, Gupta, S, Li, X, Fahy, E, Dinasarapu, AR, Sud, M, Brown, HA, Glass, CK, Murphy, RC, Russell, DW, Dennis, EA, Subramaniam, S. Analysis of inflammatory and lipid metabolic networks across RAW264.7 and thioglycolate-elicited macrophages. J Lipid Res, 54(9), 2525-2542, 2013 PMC 3735949

Yang, MY, Hilton, MB, Seaman, S, Haines, DC, Nagashima, K, Burks, CM, Tessarollo, L, Ivanova, PT, Brown, HA, Umstead, TM, Floros, J, Chroneos, ZC, St Croix, B. Essential Regulation of Lung Surfactant Homeostasis by the Orphan G Protein-Coupled Receptor GPR116. Cell Rep, 3(5), 1457-64, 2013 PMC 3695742

Kim, SH, Scott, SA, Bennett, MJ, Carson, RP, Fessel, J, Brown, HA, Ess, KC. Multi-organ abnormalities and mTORC1 activation in zebrafish model of multiple acyl-CoA dehydrogenase deficiency. PLoS Genet, 9(6), e1003563, 2013 PMC 3681725

Milne, SB, Mathews, TP, Myers, DS, Ivanova, PT, Brown, HA. Sum of the Parts: Mass Spectrometry-Based Metabolomics. Biochemistry, 52(22), 3829-3840, 2013 PMC 3674197

O'Reilly, MC, Scott, SA, Brown, KA, Oguin, TH, Thomas, PG, Daniels, JS, Morrison, R, Brown, HA, Lindsley, CW. Development of Dual PLD1/2 and PLD2 Selective Inhibitors from a Common 1,3,8-Triazaspiro[4.5]decane Core: Discovery of ML298 and ML299 That Decrease Invasive Migration in U87-MG Glioblastoma Cells. J Med Chem, 56(6), 2695-9, 2013 PMC 3632306

Tallman, KA, Armstrong, MD, Milne, SB, Marnett, LJ, Brown, HA, Porter, NA. Cobalt carbonyl complexes as probes for alkyne-tagged lipids. J Lipid Res, 54(3), 859-68, 2013 PMC 3617960

Moore,J., Li,S., Myers, DS, Milne, SB, Brown, HA, and Shaw, W.. Standardization and quantification of lysophosphatidic acid compounds by normal-phase and reverse-phase chromatography-tandem mass spectrometry.. Book chapter - Lysophospholipid receptors: Signaling and Biochemistry, John Wiley &Sons(Ed. J. Chun, T. Hla, S. Spiegel, W. Moolenaar), 137-152, 2013

Scott, SA, Xiang, Y, Mathews, TP, Plumley, HC, Myers, DS, Armstrong, MD, Tallman, KA, O'Reilly, MC, Lindsley, CW, Brown, HA. Regulation of Phospholipase D Activity and Phosphatidic Acid Production After Purinergic (P2Y6) Receptor Stimulation. J Biol Chem, 288(28), 20411-20487 PMC3711313, 2013

Li, JZ, Huang, Y, Karaman, R, Ivanova, PT, Brown, HA, Roddy, T, Castro-Perez, J, Cohen, JC, Hobbs, HH. Chronic overexpression of PNPLA3I148M in mouse liver causes hepatic steatosis. J Clin Invest, 122(11), 4130-4144, 2012 PMC 3484461

Spann, NJ, Garmire, LX, McDonald, JG, Myers, DS, Milne, SB, Shibata, N, Reichart, D, Fox, JN, Shaked, I, Heudobler, D, Raetz, CR, Wang, EW, Kelly, SL, Sullards, MC, Murphy, RC, Merrill, AH, Brown, HA, Dennis, EA, Li, AC, Ley, K, Tsimikas, S, Fahy, E, Subramaniam, S, Quehenberger, O, Russell, DW, Glass, CK. Regulated accumulation of desmosterol integrates macrophage lipid metabolism and inflammatory responses. Cell, 151(1), 138-52, 2012 PMC 3464914

Kumari, M, Schoiswohl, G, Chitraju, C, Paar, M, Cornaciu, I, Rangrez, AY, Wongsiriroj, N, Nagy, HM, Ivanova, PT, Scott, SA, Knittelfelder, O, Rechberger, GN, Birner-Gruenberger, R, Eder, S, Brown, HA, Haemmerle, G, Oberer, M, Lass, A, Kershaw, EE, Zimmermann, R, Zechner, R. Adiponutrin functions as a nutritionally regulated lysophosphatidic Acid acyltransferase. Cell Metab, 15(5), 691-702, 2012 PMC 3361708

Brown HA. Lipidomics: When apocrypha becomes canonical. Current Opin Chem Biology, 16(1-2), 221-226, 2012 PMC 3328648

Lord, CC, Betters, JL, Ivanova, PT, Milne, SB, Myers, DS, Madenspacher, J, Thomas, G, Chung, S, Liu, M, Davis, MA, Lee, RG, Crooke, RM, Graham, MJ, Parks, JS, Brasaemle, DL, Fessler, MB, Brown, HA, Brown, JM. CGI-58/ABHD5-derived signaling lipids regulate systemic inflammation and insulin action. Diabetes, 61(2), 355-63, 2012 PMC 3266405

Cipriano, R, Graham, J, Miskimen, KL, Bryson, BL, Bruntz, RC, Scott, SA, Brown, HA, Stark, GR, Jackson, MW. FAM83B mediates EGFR- and RAS-driven oncogenic transformation. J Clin Invest, 122(9), 3197-210, 2012 PMCID:3428078

Lindsley, CW, Brown, HA. Phospholipase D as a therapeutic target in brain disorders. Neuropsychopharmacology, 37(1), 301-2, 2012

Kang DW, Park MH, Lee YJ, Kim HS, Lindsley CW, Brown HA, Min DS. Autoregulation of phospholipase D activity is coupled to selective induction of phospholipase D1 expression to promote invasion of breast cancer cells. Int J Cancer , 128(4), 805-816, 2011 PMID 20473892

Brown, HA, Marnett, LJ. Introduction to lipid biochemistry, metabolism, and signaling. Chem Rev, 111(10), 5817-20, 2011 PMCID:3428078

Gorden, DL, Ivanova, PT, Myers, DS, McIntyre, JO, VanSaun, MN, Wright, JK, Matrisian, LM, Brown, HA. Increased diacylglycerols characterize hepatic lipid changes in progression of human nonalcoholic fatty liver disease; comparison to a murine model. PLoS One, 6(8), e22775, 2011 PMCID:3153459

Liu, ST, Sharon-Friling, R, Ivanova, P, Milne, SB, Myers, DS, Rabinowitz, JD, Brown, HA, Shenk, T. Synaptic vesicle-like lipidome of human cytomegalovirus virions reveals a role for SNARE machinery in virion egress. Proc Natl Acad Sci U S A, 108(31), 12869-74, 2011 PMCID:3150939

Myers, DS, Ivanova, PT, Milne, SB, Brown, HA. Quantitative analysis of glycerophospholipids by LC-MS: acquisition, data handling, and interpretation. Biochim Biophys Acta, 1811(11), 748-57, 2011 PMCID:3153459

Norton, LJ, Zhang, Q, Saqib, KM, Schrewe, H, Macura, K, Anderson, KE, Lindsley, CW, Brown, HA, Rudge, SA, Wakelam, MJ. PLD1 rather than PLD2 regulates phorbol-ester-, adhesion-dependent and Fc{gamma}-receptor-stimulated ROS production in neutrophils. J Cell Sci, 124(Pt 12), 1973-83, 2011 PMCID:3104032

Selvy, PE, Lavieri, RR, Lindsley, CW, Brown, HA. Phospholipase D: enzymology, functionality, and chemical modulation. Chem Rev, 111(10), 6064-119, 2011

Quehenberger O, Armando AM, Brown HA, Milne SB, Myers DS, Merrill AH, Bandyopadhyay S, Jones KN, Kelly S, Shaner RL, Sullards CM, Wang E, Murphy RC et al.. Lipidomics reveals remarkable diversity of lipids in human plasma. J Lipid Res, 51(11), 3299-3305, 2010 PMC 2952570

Andreyev, AY, Fahy, E, Guan, Z, Kelly, S, Li, X, McDonald, JG, Milne, S, Myers, D, Park, H, Ryan, A, Thompson, BM, Wang, E, Zhao, Y, Brown, HA, Merrill, AH, Raetz, CR, Russell, DW, Subramaniam, S, Dennis, EA. Subcellular organelle lipidomics in TLR-4-activated macrophages. J Lipid Res, 51(9), 2785-97, 2010 PMCID:2918461

Callender HL, Horn MA, DeCamp DL, Sternweis PC, Brown HA. Modeling species-specific diacylglycerol dynamics in the RAW 264.7 macrophage. J. Theor. Biol., 262(4), 679-690, 2010

Dennis, EA, Deems, RA, Harkewicz, R, Quehenberger, O, Brown, HA, Milne, SB, Myers, DS, Glass, CK, Hardiman, G, Reichart, D, Merrill, AH, Sullards, MC, Wang, E, Murphy, RC, Raetz, CR, Garrett, TA, Guan, Z, Ryan, AC, Russell, DW, McDonald, JG, Thompson, BM, Shaw, WA, Sud, M, Zhao, Y, Gupta, S, Maurya, MR, Fahy, E, Subramaniam, S. A mouse macrophage lipidome. J Biol Chem, 285(51), 39976-85, 2010 PMCID:3000979

Ivanova, PT, Milne, SB, Brown, HA. Identification of atypical ether-linked glycerophospholipid species in macrophages by mass spectrometry. J Lipid Res, 51(6), 1581-90, 2010 PMCID:3035522

Klappe, K, Dijkhuis, AJ, Hummel, I, van Dam, A, Ivanova, PT, Milne, SB, Myers, DS, Brown, HA, Permentier, H, Kok, JW. Extensive sphingolipid depletion does not affect lipid raft integrity or lipid raft localization and efflux function of the ABC transporter MRP1. Biochem J, 430(3), 519-29, 2010 PMCID:3153459

Lavieri, RR, Scott, SA, Selvy, PE, Kim, K, Jadhav, S, Morrison, RD, Daniels, JS, Brown, HA, Lindsley, CW. Design, synthesis, and biological evaluation of halogenated N-(2-(4-oxo-1-phenyl-1,3,8-triazaspiro[4.5]decan-8-yl)ethyl)benzamides: discovery of an isoform-selective small molecule phospholipase D2 inhibitor. J Med Chem, 53(18), 6706-19, 2010 PMCID:3179181

Milne, SB, Tallman, KA, Serwa, R, Rouzer, CA, Armstrong, MD, Marnett, LJ, Lukehart, CM, Porter, NA, Brown, HA. Capture and release of alkyne-derivatized glycerophospholipids using cobalt chemistry. Nat Chem Biol, 6(3), 205-207, 2010 PMCID:2822082

Shulga, YV, Myers, DS, Ivanova, PT, Milne, SB, Brown, HA, Topham, MK, Epand, RM. Molecular species of phosphatidylinositol-cycle intermediates in the endoplasmic reticulum and plasma membrane. Biochemistry, 49(2), 312-7, 2010 PMCID:2805767

Lavieri R, Scott SA, Lewis JA, Selvy PE, Armstrong MD, Brown HA, Lindsley CW. Design and synthesis of isoform-selective phospholipase D (PLD) inhibitors. Part II. Identification of the 1,3,8-triazaspiro[4,5]-decan-4-one privileged structure that engenders PLD2 selectivity. Bioorg Chem Lett , 19(8), 2240-2243, 2009 PMC3800051

Rappley I, Myers DS, Milne SB, Ivanova PT, LaVoie J, Brown HA, Selkoe DJ. Lipidomics profiling in mouse brain reveals differences between ages and genders with smaller changes associated with alfa-synuclein genotype. J Neurochem, 111(1), 15-25, 2009 PMC 2752313

Brown, HA, Murphy, RC. Working towards an exegesis for lipids in biology. Nat Chem Biol, 5(9), 602-6, 2009

Burnum KE, Cornett DS, Puolitaival SM, Milne SB, Myers DS, Tranguch S, Brown HA, Dey SK, Caprioli RM. Spatial and temporal alterations of phospholipids determined by mass spectrometry during mouse embryo implantation . J Lipid Res, 50(11), 2290-2298, 2009

Holinstat, M, Preininger, AM, Milne, SB, Hudson, WJ, Brown, HA, Hamm, HE. Irreversible platelet activation requires protease-activated receptor 1-mediated signaling to phosphatidylinositol phosphates. Mol Pharmacol, 76(2), 301-13, 2009 PMCID:2713123

Ivanova, PT, Milne, SB, Myers, DS, Brown, HA. Lipidomics: a mass spectrometry based systems level analysis of cellular lipids. Curr Opin Chem Biol, 13, 526-531, 2009

Lewis, JA, Scott, SA, Lavieri, R, Buck, JR, Selvy, PE, Stoops, SL, Armstrong, MD, Brown, HA, Lindsley, CW. Design and synthesis of isoform-selective phospholipase D (PLD) inhibitors. Part I: Impact of alternative halogenated privileged structures for PLD1 specificity. Bioorg Med Chem Lett, 19(7), 1916-20, 2009

Lung, M, Shulga, YV, Ivanova, PT, Myers, DS, Milne, SB, Brown, HA, Topham, MK, Epand, RM. Diacylglycerol kinase epsilon is selective for both acyl chains of phosphatidic acid or diacylglycerol. J Biol Chem, 284(45), 31062-73, 2009 PMCID:2781506

Marlo, JE, Niswender, CM, Days, EL, Bridges, TM, Xiang, Y, Rodriguez, AL, Shirey, JK, Brady, AE, Nalywajko, T, Luo, Q, Austin, CA, Williams, MB, Kim, K, Williams, R, Orton, D, Brown, HA, Lindsley, CW, Weaver, CD, Conn, PJ. Discovery and characterization of novel allosteric potentiators of M1 muscarinic receptors reveals multiple modes of activity. Mol Pharmacol, 2009 PMCID:2684909

Rappley, I, Myers, DS, Milne, SB, Ivanova, PT, Lavoie, MJ, Brown, HA, Selkoe, DJ. Lipidomic profiling in mouse brain reveals differences between ages and genders, with smaller changes associated with alpha-synuclein genotype. J Neurochem, 111(1), 15-25, 2009 PMCID:2752313

Scott, SA, Selvy, PE, Buck, JR, Cho, HP, Criswell, TL, Thomas, AL, Armstrong, MD, Arteaga, CL, Lindsley, CW, Brown, HA. Design of isoform-selective phospholipase D inhibitors that modulate cancer cell invasiveness. Nat Chem Biol, 5(2), 108-17, 2009

Zeng, XX, Zheng, X, Xiang, Y, Cho, HP, Jessen, JR, Zhong, TP, Solnica-Krezel, L, Brown, HA. Phospholipase D1 is required for angiogenesis of intersegmental blood vessels in zebrafish. Dev Biol, 328(2), 363-76, 2009 PMCID:2688691

Brown, HA. New mechanism for a classic folate inhibitor. Nat Chem Biol, 4(10), 581-2, 2008

Garcia, A, Zheng, Y, Zhao, C, Toschi, A, Fan, J, Shraibman, N, Brown, HA, Bar-Sagi, D, Foster, DA, Arbiser, JL. Honokiol suppresses survival signals mediated by Ras-dependent phospholipase D activity in human cancer cells. Clin Cancer Res, 14(13), 4267-74, 2008

Milne, SB, Ivanova, PT, Armstrong, MD, Myers, DS, Lubarda, J, Shulga, YV, Topham, MK, Brown, HA, Epand, RM. Dramatic differences in the roles in lipid metabolism of two isoforms of diacylglycerol kinase. Biochemistry, 47(36), 9372-9, 2008 PMCID:2744458

Sun, Y, Fang, Y, Yoon, MS, Zhang, C, Roccio, M, Zwartkruis, FJ, Armstrong, M, Brown, HA, Chen, J. Phospholipase D1 is an effector of Rheb in the mTOR pathway. Proc Natl Acad Sci U S A, 105(24), 8286-91, 2008 PMCID:2448829

Brown, HA, Henage, LG, Preininger, AM, Xiang, Y, Exton, JH. Biochemical analysis of phospholipase D. Methods Enzymol, 434, 49-87, 2007

Callender, HL, Forrester, JS, Ivanova, P, Preininger, A, Milne, S, Brown, HA. Quantification of diacylglycerol species from cellular extracts by electrospray ionization mass spectrometry using a linear regression algorithm. Anal Chem, 79(1), 263-72, 2007

Ivanova, PT, Milne, SB, Byrne, MO, Xiang, Y, Brown, HA. Glycerophospholipid identification and quantitation by electrospray ionization mass spectrometry. Methods Enzymol, 432, 21-57, 2007

Murphy, SC, Fernandez-Pol, S, Chung, PH, Prasanna Murthy, SN, Milne, SB, Salomao, M, Brown, HA, Lomasney, JW, Mohandas, N, Haldar, K. Cytoplasmic remodeling of erythrocyte raft lipids during infection by the human malaria parasite Plasmodium falciparum. Blood, 110(6), 2132-9, 2007 PMCID:1976375

Rouzer, CA, Ivanova, PT, Byrne, MO, Brown, HA, Marnett, LJ. Lipid profiling reveals glycerophospholipid remodeling in zymosan-stimulated macrophages. Biochemistry, 46(20), 6026-42, 2007

Sud, M, Fahy, E, Cotter, D, Brown, A, Dennis, EA, Glass, CK, Merrill, AH, Murphy, RC, Raetz, CR, Russell, DW, Subramaniam, S. LMSD: LIPID MAPS structure database. Nucleic Acids Res, 35(Database issue), D527-32, 2007 PMCID:1669719

Henage, LG, Exton, JH, Brown, HA. Kinetic analysis of a mammalian phospholipase D: allosteric modulation by monomeric GTPases, protein kinase C, and polyphosphoinositides. J Biol Chem, 281(6), 3408-17, 2006

Milne, S, Ivanova, P, Forrester, J, Alex Brown, H. Lipidomics: an analysis of cellular lipids by ESI-MS. Methods, 39(2), 92-103, 2006

Milne, S.B., Forrester, J.S., Ivanova, P.T., and Brown H.A.. Multiplexed Lipid Arrays of Antiimmunoglobulin IV-Induced Changes in the Glycerophospholipid Composition of WEHI-231 Cells. Functional Lipidomics. Eds L.Feng, GD Prestwich. CRC Press, Boca Raton, FL, 2006

Preininger, AM, Henage, LG, Oldham, WM, Yoon, EJ, Hamm, HE, Brown, HA. Direct modulation of phospholipase D activity by Gbetagamma. Mol Pharmacol, 70(1), 311-8, 2006

Rouzer, CA, Ivanova, PT, Byrne, MO, Milne, SB, Marnett, LJ, Brown, HA. Lipid profiling reveals arachidonate deficiency in RAW264.7 cells: Structural and functional implications. Biochemistry, 45(49), 14795-808, 2006 PMCID:2443946

Dennis, E.A., Brown, H.A., Merrill, A.H., Murphy, R.C., Raetz, C.R., Russell, D.W., Wooley, J. and Deems, R.A. The LIPID MAPS Approach to Lipidomics.. In: Functional Lipidomics. edited by L. Feng and G.D. Prestwich. CRC Press, Boca Raton, FL, 2005

Milne, SB, Ivanova, PT, DeCamp, D, Hsueh, RC, Brown, HA. A targeted mass spectrometric analysis of phosphatidylinositol phosphate species. J Lipid Res, 46(8), 1796-802, 2005

Forrester, Jeffrey S, Milne, Stephen B, Ivanova, Pavlina T, Brown, H Alex. Computational lipidomics: a multiplexed analysis of dynamic changes in membrane lipid composition during signal transduction. Mol Pharmacol, 65(4), 813-21, 2004

Ivanova, Pavlina T, Milne, Stephen B, Forrester, Jeffrey S, Brown, H Alex. LIPID Arrays: New Tools in the Understanding of Membrane Dynamics and Lipid Signaling. Mol Interv, 4(2), 86-96, 2004

Walker, Stephanie J, Brown, H Alex, , . Measurement of G protein-coupled receptor-stimulated phospholipase D activity in intact cells. Methods Mol Biol, 237, 89-97, 2004

Fang, Yimin, Park, In-Hyun, Wu, Ai-Luen, Du, Guangwei, Huang, Ping, Frohman, Michael A, Walker, Stephanie J, Brown, H Alex, Chen, Jie. PLD1 regulates mTOR signaling and mediates Cdc42 activation of S6K1. Curr Biol, 13(23), 2037-44, 2003

Ge, Mingtao, Gidwani, Arun, Brown, H Alex, Holowka, David, Baird, Barbara, Freed, Jack H. Ordered and disordered phases coexist in plasma membrane vesicles of RBL-2H3 mast cells. An ESR study.. Biophys J, 85(2), 1278-88, 2003 PMCID:1303245

Gidwani, Arun, Brown, H Alex, Holowka, David, Baird, Barbara. Disruption of lipid order by short-chain ceramides correlates with inhibition of phospholipase D and downstream signaling by FcepsilonRI.. J Cell Sci, 116(Pt 15), 3177-87, 2003

Milne, S.B., Forrester, J.S., Ivanova, P.T., and Brown, A.H. Multipleaxed Lipid Arrays of Aniti-IgM0induced changes in the glycerophospholipid composition of WEHI-231 cells. AfCS Research Reports, 2003

Sieczkarski, Sara B, Brown, H Alex, Whittaker, Gary R. Role of protein kinase C betaII in influenza virus entry via late endosomes.. J Virol, 77(1), 460-9, 2003 PMCID:140583

Brooks, Marjory B, Catalfamo, James L, Brown, H Alex, Ivanova, Pavlina, Lovaglio, Jamie. A hereditary bleeding disorder of dogs caused by a lack of platelet procoagulant activity. Blood, 99(7), 2434-41, 2002

Eisen, Susanne F, Brown, H Alex. Selective estrogen receptor (ER) modulators differentially regulate phospholipase D catalytic activity in ER-negative breast cancer cells. Mol Pharmacol, 62(4), 911-20, 2002

Gilman, A.G., et al. Overview of the alliance for cellular signaling. Nature, 420, 703-706, 2002

Walker, Stephanie J, Brown, H Alex. Specificity of Rho insert-mediated activation of phospholipase D1. J Biol Chem, 277(29), 26260-7, 2002

Cohen, J S, Brown, H A. Phospholipases stimulate secretion in RBL mast cells. Biochemistry, 40(22), 6589-97, 2001

Ganley, I G, Walker, S J, Manifava, M, Li, D, Brown, H A, Ktistakis, N T. Interaction of phospholipase D1 with a casein-kinase-2-like serine kinase. Biochem J, 354(Pt 2), 369-78, 2001 PMCID:1221665

Ge, M, Cohen, J S, Brown, H A, Freed, J H. ADP ribosylation factor 6 binding to phosphatidylinositol 4,5-bisphosphate-containing vesicles creates defects in the bilayer structure: an electron spin resonance study. Biophys J, 81(2), 994-1005, 2001 PMCID:1301569

Ivanova, P T, Cerda, B A, Horn, D M, Cohen, J S, McLafferty, F W, Brown, H A. Electrospray ionization mass spectrometry analysis of changes in phospholipids in RBL-2H3 mastocytoma cells during degranulation. Proc Natl Acad Sci U S A, 98(13), 7152-7, 2001 PMCID:34638

Walker, S J, Wu, W J, Cerione, R A, Brown, H A. Activation of phospholipase D1 by Cdc42 requires the Rho insert region. J Biol Chem, 275(21), 15665-8, 2000

Paradiso, A M, Brown, H A, Ye, H, Harden, T K, Boucher, R C. Heterogeneous responses of cell Ca2+ in human airway epithelium. Exp Lung Res, 25(4), 277-90, 1999

Singer, W D, Brown, H A, Sternweis, P C. Regulation of eukaryotic phosphatidylinositol-specific phospholipase C and phospholipase D. Annu Rev Biochem, 66, 475-509, 1997

Ktistakis, N T, Brown, H A, Waters, M G, Sternweis, P C, Roth, M G. Evidence that phospholipase D mediates ADP ribosylation factor-dependent formation of Golgi coated vesicles. J Cell Biol, 134(2), 295-306, 1996 PMCID:2120869

Lazarowski, E R, Watt, W C, Stutts, M J, Brown, H A, Boucher, R C, Harden, T K. Enzymatic synthesis of UTP gamma S, a potent hydrolysis resistant agonist of P2U-purinoceptors. Br J Pharmacol, 117(1), 203-9, 1996 PMCID:1909382

Singer, W D, Brown, H A, Jiang, X, Sternweis, P C. Regulation of phospholipase D by protein kinase C is synergistic with ADP-ribosylation factor and independent of protein kinase activity. J Biol Chem, 271(8), 4504-10, 1996

Brown, H A, Gutowski, S, Kahn, R A, Sternweis, P C. Partial purification and characterization of Arf-sensitive phospholipase D from porcine brain. J Biol Chem, 270(25), 14935-43, 1995

Brown, H A, Sternweis, P C. Stimulation of phospholipase D by ADP-ribosylation factor. Methods Enzymol, 257, 313-24, 1995

Ktistakis, N T, Brown, H A, Sternweis, P C, Roth, M G. Phospholipase D is present on Golgi-enriched membranes and its activation by ADP ribosylation factor is sensitive to brefeldin A. Proc Natl Acad Sci U S A, 92(11), 4952-6, 1995 PMCID:41825

Singer, W D, Brown, H A, Bokoch, G M, Sternweis, P C. Resolved phospholipase D activity is modulated by cytosolic factors other than Arf. J Biol Chem, 270(25), 14944-50, 1995

Brown, H A, Gutowski, S, Moomaw, C R, Slaughter, C, Sternweis, P C. ADP-ribosylation factor, a small GTP-dependent regulatory protein, stimulates phospholipase D activity. Cell, 75(6), 1137-44, 1993

Brown, H A, Lazarowski, E R, Boucher, R C, Harden, T K. Evidence that UTP and ATP regulate phospholipase C through a common extracellular 5''-nucleotide receptor in human airway epithelial cells. Mol Pharmacol, 40(5), 648-55, 1991

Harden, T K, Boyer, J L, Brown, H A, Cooper, C L, Jeffs, R A, Martin, M W. Biochemical properties of a P2Y-purinergic receptor. Ann N Y Acad Sci, 603, 256-66, 1990

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