Vanderbilt University School of Medicine

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

Lab Url:

Phone Number: 615-936-3888


Brown, H. Alex's picture

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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