LU LU, M.D., M.S.


Tel: (901) 448-7557

Research Interest/Specialty

The focus of my laboratory is on the examination of the role of genetics in a number of nervous system-related processes including the development of neuronal populations, the genetic underpinnings of neuroanatomical/structural features of the brain, the genetic control over responses to alcohol and stress, and more recently genetic basis of variation of gene expression. Also my lab is involved in producing collaborative resources for mouse genetics including new BXD strains, density genotypes, whole genome sequence of DBA/2J, and sets of gene expression data across BXD mice, and for experimental/translational models of human disease, including Alzheimer''s, glaucoma, addiction, and psychiatric disease. The types of studies conducted in my lab range from a more global perspective in identifying relevant regions of the genome that control phenotypic differences among genetically distinct organisms to more targeted research where the role of specific genes is evaluated. In these studies, I use a combination of molecular biological, behavioral, morphological and bioinformatic methods, and genetics and system biology strategy. In the past, the goal of genetic research was to identify the critical gene involved in special phenotype. However, it has become clear that individual gene does not work in isolation but in combination with other genes necessitating the identification of the relevant gene networks. Thus, much of the work in my lab is focused on evaluating phenotypes with the ultimate goal of identifying the genetic networks that control each phenotype. More recently, I have begun to examine changes in microRNAs to more completely assess the genetic networks that underlie these complex processes.


  1. Lu, L, Pandey, AK, Houseal, MT, Mulligan, MK. The Genetic Architecture of Murine Glutathione Transferases. PLoS One, 11 (2), e0148230, 2016.
  2. Shi, X, Walter, NA, Harkness, JH, Neve, KA, Williams, RW, Lu, L, Belknap, JK, Eshleman, AJ, Phillips, TJ, Janowsky, A. Genetic Polymorphisms Affect Mouse and Human Trace Amine-Associated Receptor 1 Function. PLoS One, 11 (3), e0152581, 2016.
  3. Lu, H, Lu, L, Williams, RW, Jablonski, MM. Iris transillumination defect and its gene modulators do not correlate with intraocular pressure in the BXD family of mice. Mol Vis, 22, 224-33, 2016.
  4. Alam, G, Miller, DB, O'Callaghan, JP, Lu, L, Williams, RW, Jones, BC. MPTP neurotoxicity is highly concordant between the sexes among BXD recombinant inbred mouse strains. Neurotoxicology, 55, 40-47, 2016.
  5. Schultz, NG, Ingels, J, Hillhouse, A, Wardwell, K, Chang, PL, Cheverud, JM, Lutz, C, Lu, L, Williams, RW, Dean, MD. The Genetic Basis of Baculum Size and Shape Variation in Mice. G3 (Bethesda), 6 (5), 1141-51, 2016.
  6. Franke, B, Stein, JL, Ripke, S, Anttila, V, Hibar, DP, van Hulzen, KJ, Arias-Vasquez, A, Smoller, JW, Nichols, TE, Neale, MC, McIntosh, AM, Lee, P, McMahon, FJ, Meyer-Lindenberg, A, Mattheisen, M, Andreassen, OA, Gruber, O, Sachdev, PS, Roiz-Santiañez, R, Saykin, AJ, Ehrlich, S, Mather, KA, Turner, JA, Schwarz, E, Thalamuthu, A, Yao, Y, Ho, YY, Martin, NG, Wright, MJ, , , , , , , , , O'Donovan, MC, Thompson, PM, Neale, BM, Medland, SE, Sullivan, PF. Genetic influences on schizophrenia and subcortical brain volumes: large-scale proof of concept. Nat Neurosci, 19 (3), 420-31, 2016.
  7. Urquhart, KR, Zhao, Y, Baker, JA, Lu, Y, Yan, L, Cook, MN, Jones, BC, Hamre, KM, Lu, L. A novel heat shock protein alpha 8 (Hspa8) molecular network mediating responses to stress- and ethanol-related behaviors. Neurogenetics, 2016.
  8. King, R, Lu, L, Williams, RW, Geisert, EE. Transcriptome networks in the mouse retina: An exon level BXD RI database. Mol Vis, 21, 1235-51, 2015.
  9. Ashbrook, DG, Williams, RW, Lu, L, Hager, R. A cross-species genetic analysis identifies candidate genes for mouse anxiety and human bipolar disorder. Front Behav Neurosci, 9, 171, 2015.
  10. Zhao, W, Zhao, T, Chen, Y, Zhao, F, Gu, Q, Williams, RW, Bhattacharya, SK, Lu, L, Sun, Y. A Murine Hypertrophic Cardiomyopathy Model: The DBA/2J Strain. PLoS One, 10 (8), e0133132, 2015.
  11. Delprato, A, Bonheur, B, Algéo, MP, Rosay, P, Lu, L, Williams, RW, Crusio, WE. Systems genetic analysis of hippocampal neuroanatomy and spatial learning in mice. Genes Brain Behav, 14 (8), 591-606, 2015.
  12. Xue, Y, Li, J, Yan, L, Lu, L, Liao, FF. Genetic variability to diet-induced hippocampal dysfunction in BXD recombinant inbred (RI) mouse strains. Behav Brain Res, 292, 83-94, 2015.
  13. Cook, MN, Baker, JA, Heldt, SA, Williams, RW, Hamre, KM, Lu, L. Identification of candidate genes that underlie the QTL on chromosome 1 that mediates genetic differences in stress-ethanol interactions. Physiol Genomics, 47 (8), 308-17, 2015.
  14. Wang, X, Pandey, AK, Mulligan, MK, Williams, EG, Mozhui, K, Li, Z, Jovaisaite, V, Quarles, LD, Xiao, Z, Huang, J, Capra, JA, Chen, Z, Taylor, WL, Bastarache, L, Niu, X, Pollard, KS, Ciobanu, DC, Reznik, AO, Tishkov, AV, Zhulin, IB, Peng, J, Nelson, SF, Denny, JC, Auwerx, J, Lu, L, Williams, RW. Joint mouse-human phenome-wide association to test gene function and disease risk. Nat Commun, 7, 10464, 2015.
  15. Wang, L, Lu, W, Zhang, L, Huang, Y, Scheib, R, Liu, X, Myers, L, Lu, L, Farber, CR, Liu, G, Wang, CY, Deng, H, Williams, RW, Wang, Y, Gu, W, Jiao, Y. Trps1 differentially modulates the bone mineral density between male and female mice and its polymorphism associates with BMD differently between women and men. PLoS One, 9 (1), e84485, 2014.
  16. Pandey, AK, Lu, L, Wang, X, Homayouni, R, Williams, RW. Functionally enigmatic genes: a case study of the brain ignorome. PLoS One, 9 (2), e88889, 2014.
  17. Goldowitz, D, Lussier, AA, Boyle, JK, Wong, K, Lattimer, SL, Dubose, C, Lu, L, Kobor, MS, Hamre, KM. Molecular pathways underpinning ethanol-induced neurodegeneration. Front Genet, 5, 203, 2014.
  18. Gu, J, Lu, L, Chen, M, Xu, L, Lan, Q, Li, Q, Liu, Z, Chen, G, Wang, P, Wang, X, Brand, D, Olsen, N, Zheng, SG. TGF-β-induced CD4+Foxp3+ T cells attenuate acute graft-versus-host disease by suppressing expansion and killing of effector CD8+ cells. J Immunol, 193 (7), 3388-97, 2014.
  19. Zhang, Y, Huang, J, Jiao, Y, David, V, Kocak, M, Roan, E, Di'Angelo, D, Lu, L, Hasty, KA, Gu, W. Bone morphology in 46 BXD recombinant inbred strains and femur-tibia correlation. ScientificWorldJournal, 2015, 728278, 2014.
  20. Lu, L, Lan, Q, Li, Z, Zhou, X, Gu, J, Li, Q, Wang, J, Chen, M, Liu, Y, Shen, Y, Brand, DD, Ryffel, B, Horwitz, DA, Quismorio, FP, Liu, Z, Li, B, Olsen, NJ, Zheng, SG. Critical role of all-trans retinoic acid in stabilizing human natural regulatory T cells under inflammatory conditions. Proc Natl Acad Sci U S A, 111 (33), E3432-40, 2014.
  21. Jones, BC, Huang, X, Mailman, RB, Lu, L, Williams, RW. Response. J Biochem Mol Toxicol, 28 (7), 291, 2014.
  22. Keeley, PW, Zhou, C, Lu, L, Williams, RW, Melmed, S, Reese, BE. Pituitary tumor-transforming gene 1 regulates the patterning of retinal mosaics. Proc Natl Acad Sci U S A, 111 (25), 9295-300, 2014.
  23. Jones, BC, Huang, X, Mailman, RB, Lu, L, Williams, RW. The perplexing paradox of paraquat: the case for host-based susceptibility and postulated neurodegenerative effects. J Biochem Mol Toxicol, 28 (5), 191-7, 2014.
  24. Jones, BC, O'Callaghan, JP, Lu, L, Williams, RW, Alam, G, Miller, DB. Genetic correlational analysis reveals no association between MPP+ and the severity of striatal dopaminergic damage following MPTP treatment in BXD mouse strains. Neurotoxicol Teratol, 45, 91-2, 2014.
  25. Ashbrook, DG, Williams, RW, Lu, L, Stein, JL, Hibar, DP, Nichols, TE, Medland, SE, Thompson, PM, Hager, R. Joint genetic analysis of hippocampal size in mouse and human identifies a novel gene linked to neurodegenerative disease. BMC Genomics, 15, 850, 2014.
  26. Huang, Y, Zhu, X, Wang, L, Liu, X, Lu, L, Gu, W, Jiao, Y. Genome wide analysis of sex difference in gene expression profiles of bone formations using sfx mice and BXD RI strains. ScientificWorldJournal, 2014, 584910, 2014.
  27. Ye, R, Carneiro, AM, Airey, D, Sanders-Bush, E, Williams, RW, Lu, L, Wang, J, Zhang, B, Blakely, RD. Evaluation of heritable determinants of blood and brain serotonin homeostasis using recombinant inbred mice. Genes Brain Behav, 13 (3), 247-60, 2014.
  28. Ito, J, Roy, S, Liu, Y, Cao, Y, Fletcher, M, Lu, L, Boughter, JD, Grün, S, Heck, DH. Whisker barrel cortex delta oscillations and gamma power in the awake mouse are linked to respiration. Nat Commun, 5, 3572, 2013.
  29. Mozhui, K, Ciobanu, DC, Schikorski, T, Wang, X, Lu, L, Williams, RW. Dissection of a QTL hotspot on mouse distal chromosome 1 that modulates neurobehavioral phenotypes and gene expression. PLoS Genet, 4 (11), e1000260, 2008.
  30. Reiner, DJ, Jan, TA, Boughter, JD, Li, CX, Lu, L, Williams, RW, Waters, RS. Genetic analysis of tongue size and taste papillae number and size in recombinant inbred strains of mice. Chem Senses, 33 (8), 693-707, 2008.
  31. Brigman, JL, Mathur, P, Lu, L, Williams, RW, Holmes, A. Genetic relationship between anxiety-related and fear-related behaviors in BXD recombinant inbred mice. Behav Pharmacol, 2008.
  32. Lu, L, Chen, SS, Hassid, A, Sun, Y. Cardiac fibrogenesis following infarction in mice with deletion of inducible nitric oxide synthase. Am J Med Sci, 335 (6), 431-8, 2008.
  33. Abdeltawab, NF, Aziz, RK, Kansal, R, Rowe, SL, Su, Y, Gardner, L, Brannen, C, Nooh, MM, Attia, RR, Abdelsamed, HA, Taylor, WL, Lu, L, Williams, RW, Kotb, M. An unbiased systems genetics approach to mapping genetic loci modulating susceptibility to severe streptococcal sepsis. PLoS Pathog, 4 (4), e1000042, 2008.
  34. Rosen, GD, Pung, CJ, Owens, CB, Caplow, J, Kim, H, Mozhui, K, Lu, L, Williams, RW. Genetic modulation of striatal volume by loci on Chrs 6 and 17 in BXD recombinant inbred mice. Genes Brain Behav, 2008.
  35. Miyairi, I, Tatireddigari, VR, Mahdi, OS, Rose, LA, Belland, RJ, Lu, L, Williams, RW, Byrne, GI. The p47 GTPases Iigp2 and Irgb10 regulate innate immunity and inflammation to murine Chlamydia psittaci infection. J Immunol, 179 (3), 1814-24, 2007.
  36. Jan, TA, Lu, L, Li, CX, Williams, RW, Waters, RS. Genetic analysis of posterior medial barrel subfield (PMBSF) size in somatosensory cortex (SI) in recombinant inbred strains of mice. BMC Neurosci, 9, 3, 2007.
  37. Bao, L, Peirce, JL, Zhou, M, Li, H, Goldowitz, D, Williams, RW, Lu, L, Cui, Y. An integrative genomics strategy for systematic characterization of genetic loci modulating phenotypes. Hum Mol Genet, 16 (11), 1381-90, 2007.
  38. Peirce, JL, Broman, KW, Lu, L, Williams, RW. A simple method for combining genetic mapping data from multiple crosses and experimental designs. PLoS ONE, 2 (10), e1036, 2007.
  39. Peirce, JL, Broman, KW, Lu, L, Chesler, EJ, Zhou, G, Airey, DC, Birmingham, AE, Williams, RW. Genome Reshuffling for Advanced Intercross Permutation (GRAIP): simulation and permutation for advanced intercross population analysis. PLoS ONE, 3 (4), e1977, 2007.
  40. Mozhui, K, Hamre, KM, Holmes, A, Lu, L, Williams, RW. Genetic and structural analysis of the basolateral amygdala complex in BXD recombinant inbred mice. Behav Genet, 37 (1), 223-43, 2007.
  41. Bao, L, Zhou, M, Wu, L, Lu, L, Goldowitz, D, Williams, RW, Cui, Y. PolymiRTS Database: linking polymorphisms in microRNA target sites with complex traits. Nucleic Acids Res, 35 (Database issue), D51-4, 2007.
  42. Peirce, JL, Li, H, Wang, J, Manly, KF, Hitzemann, RJ, Belknap, JK, Rosen, GD, Goodwin, S, Sutter, TR, Williams, RW, Lu, L. How replicable are mRNA expression QTL. Mamm Genome, 17 (6), 643-56, 2006.
  43. Bao, L, Wei, L, Peirce, JL, Homayouni, R, Li, H, Zhou, M, Chen, H, Lu, L, Williams, RW, Pfeffer, LM, Goldowitz, D, Cui, Y. Combining gene expression QTL mapping and phenotypic spectrum analysis to uncover gene regulatory relationships. Mamm Genome, 17 (6), 575-83, 2006.
  44. Li, H, Chen, H, Bao, L, Manly, KF, Chesler, EJ, Lu, L, Wang, J, Zhou, M, Williams, RW, Cui, Y. Integrative genetic analysis of transcription modules: towards filling the gap between genetic loci and inherited traits. Hum Mol Genet, 15 (3), 481-92, 2006.
  45. Vazquez-Chona, FR, Khan, AN, Chan, CK, Moore, AN, Dash, PK, Hernandez, MR, Lu, L, Chesler, EJ, Manly, KF, Williams, RW, Geisert, EE. Genetic networks controlling retinal injury. Mol Vis, 11, 958-70, 2005.
  46. Li, CX, Wei, X, Lu, L, Peirce, JL, Williams, RW, Waters, RS. Genetic analysis of barrel field size in the first somatosensory area (SI) in inbred and recombinant inbred strains of mice. Somatosens Mot Res, 22 (3), 141-50, 2005.
  47. Matthews, DB, Bhave, SV, Belknap, JK, Brittingham, C, Chesler, EJ, Hitzemann, RJ, Hoffmann, PL, Lu, L, McWeeney, S, Miles, MF, Tabakoff, B, Williams, RW. Complex genetics of interactions of alcohol and CNS function and behavior. Alcohol Clin Exp Res, 29 (9), 1706-19, 2005.
  48. Jablonski, MM, Dalke, C, Wang, X, Lu, L, Manly, KF, Pretsch, W, Favor, J, Pardue, MT, Rinchik, EM, Williams, RW, Goldowitz, D, Graw, J. An ENU-induced mutation in Rs1h causes disruption of retinal structure and function. Mol Vis, 11, 569-81, 2005.
  49. Kamalov, G, Varma, BR, Lu, L, Sun, Y, Weber, KT, Guntaka, RV. Expression of the multifunctional Y-box protein, YB-1, in myofibroblasts of the infarcted rat heart. Biochem Biophys Res Commun, 334 (1), 239-44, 2005.
  50. Li, H, Lu, L, Manly, KF, Chesler, EJ, Bao, L, Wang, J, Zhou, M, Williams, RW, Cui, Y. Inferring gene transcriptional modulatory relations: a genetical genomics approach. Hum Mol Genet, 14 (9), 1119-25, 2005.
  51. Chesler, EJ, Lu, L, Shou, S, Qu, Y, Gu, J, Wang, J, Hsu, HC, Mountz, JD, Baldwin, NE, Langston, MA, Threadgill, DW, Manly, KF, Williams, RW. Complex trait analysis of gene expression uncovers polygenic and pleiotropic networks that modulate nervous system function. Nat Genet, 37 (3), 233-42, 2005.
  52. Jablonski, MM, Wang, X, Lu, L, Miller, DR, Rinchik, EM, Williams, RW, Goldowitz, D. The Tennessee Mouse Genome Consortium: identification of ocular mutants. Vis Neurosci, 22 (5), 595-604, 2005.
  53. Zhao, W, Lu, L, Chen, SS, Sun, Y. Temporal and spatial characteristics of apoptosis in the infarcted rat heart. Biochem Biophys Res Commun, 325 (2), 605-11, 2004.
  54. Lu, L, Quinn, MT, Sun, Y. Oxidative stress in the infarcted heart: role of de novo angiotensin II production. Biochem Biophys Res Commun, 325 (3), 943-51, 2004.
  55. Lu, L, Chen, SS, Zhang, JQ, Ramires, FJ, Sun, Y. Activation of nuclear factor-kappaB and its proinflammatory mediator cascade in the infarcted rat heart. Biochem Biophys Res Commun, 321 (4), 879-85, 2004.
  56. Jablonski, MM, Lu, L, Wang, X, Chesler, EJ, Carps, E, Qi, S, Gu, J, Williams, RW. The ldis1 lens mutation in RIIIS/J mice maps to chromosome 8 near cadherin 1. Mol Vis, 10, 577-87, 2004.
  57. Lu, L, Zhang, JQ, Ramires, FJ, Sun, Y. Molecular and cellular events at the site of myocardial infarction: from the perspective of rebuilding myocardial tissue. Biochem Biophys Res Commun, 320 (3), 907-13, 2004.
  58. Chesler, EJ, Lu, L, Wang, J, Williams, RW, Manly, KF. WebQTL: rapid exploratory analysis of gene expression and genetic networks for brain and behavior. Nat Neurosci, 7 (5), 485-6, 2004.
  59. Peirce, JL, Lu, L, Gu, J, Silver, LM, Williams, RW. A new set of BXD recombinant inbred lines from advanced intercross populations in mice. BMC Genet, 5, 7, 2004.
  60. Chesler, EJ, Wang, J, Lu, L, Qu, Y, Manly, KF, Williams, RW. Genetic correlates of gene expression in recombinant inbred strains: a relational model system to explore neurobehavioral phenotypes. Neuroinformatics, 1 (4), 343-57, 2004.
  61. Sun, Y, Zhang, J, Lu, L, Bedigian, MP, Robinson, AD, Weber, KT. Tissue angiotensin II in the regulation of inflammatory and fibrogenic components of repair in the rat heart. J Lab Clin Med, 143 (1), 41-51, 2004.
  62. Seecharan, DJ, Kulkarni, AL, Lu, L, Rosen, GD, Williams, RW. Genetic control of interconnected neuronal populations in the mouse primary visual system. J Neurosci, 23 (35), 11178-88, 2003.
  63. Ahokas, RA, Warrington, KJ, Gerling, IC, Sun, Y, Wodi, LA, Herring, PA, Lu, L, Bhattacharya, SK, Postlethwaite, AE, Weber, KT. Aldosteronism and peripheral blood mononuclear cell activation: a neuroendocrine-immune interface. Circ Res, 93 (10), e124-35, 2003.
  64. Peirce, JL, Chesler, EJ, Williams, RW, Lu, L. Genetic architecture of the mouse hippocampus: identification of gene loci with selective regional effects. Genes Brain Behav, 2 (4), 238-52, 2003.
  65. Cook, R, Lu, L, Gu, J, Williams, RW, Smeyne, RJ. Identification of a single QTL, Mptp1, for susceptibility to MPTP-induced substantia nigra pars compacta neuron loss in mice. Brain Res Mol Brain Res, 110 (2), 279-88, 2003.
  66. Airey, DC, Lu, L, Shou, S, Williams, RW. Genetic sources of individual differences in the cerebellum. Cerebellum, 1 (4), 233-40, 2002.
  67. Sun, Y, Zhang, J, Lu, L, Chen, SS, Quinn, MT, Weber, KT. Aldosterone-induced inflammation in the rat heart : role of oxidative stress. Am J Pathol, 161 (5), 1773-81, 2002.
  68. Williams, RW, Gu, J, Qi, S, Lu, L. The genetic structure of recombinant inbred mice: high-resolution consensus maps for complex trait analysis. Genome Biol, 2 (11), RESEARCH0046, 2001.
  69. Airey, DC, Lu, L, Williams, RW. Genetic control of the mouse cerebellum: identification of quantitative trait loci modulating size and architecture. J Neurosci, 21 (14), 5099-109, 2001.
  70. Hardy, CL, Lu, L, Nguyen, P, Woodland, DL, Williams, RW, Blackman, MA. Identification of quantitative trait loci controlling activation of TRBV4 CD8+ T cells during murine gamma-herpesvirus-induced infectious mononucleosis. Immunogenetics, 53 (5), 395-400, 2001.
  71. Lu, L, Airey, DC, Williams, RW. Complex trait analysis of the hippocampus: mapping and biometric analysis of two novel gene loci with specific effects on hippocampal structure in mice. J Neurosci, 21 (10), 3503-14, 2001.
  72. Williams, RW, Airey, DC, Kulkarni, A, Zhou, G, Lu, L. Genetic dissection of the olfactory bulbs of mice: QTLs on four chromosomes modulate bulb size. Behav Genet, 31 (1), 61-77, 2001.