Professor and Chair
Genetics, Genomics and Informatics (primary)
UT-ORNL Governor's Chair in Computational Genomics
Department of Pediatrics (secondary)
Department of Ophthalmology (secondary)
Department of Anatomy and Neurobiology (secondary)

Tel: (901) 604-4752


  • Ph.D., University of California, Davis, Physiology
  • B.A., University of California, Santa Cruz, Psychobiology

Web Sites

1. GeneNetwork: (the classic version), (the beta version of GeneNetwork 2)

2. Mouse Brain Library:

3. Nervenet (Williams laboratory web site):

4. Center for Integrative and Translational Genomics:

My Current CV

Research interest/specialty

GOAL: Our group''s main goal is to build resources and tools such as GeneNetwork ( to predict disease risk from sequence data and environmental exposure.

BACKGROUND ON THE SCIENCE: Human diseases are almost always caused by complex interactions among genetic, developmental, epigenetic, and environmental factors. Can we accurately predict the risk of developing diseases by taking into account the interactions and roles played by many contributing factors at different levels—from DNA varints to social context?

One initial approach is to use animal models that incorporate a high level of genetic complexity that mirrors that of humans. Any two humans will differ at up to 10 million points along their DNA and this is also true of the lines of mice that we study. But the advantage of studying mice is that each line can be replicated. This is like having an inexhaustble number of identical twins. We have about 150 related members of a large family of mice to study. We are using this BXD family to test how differences in their DNA, in their environment, and in particular experimental treatments influence disease onset and severity.

We would like to be able to predict which family member (strain of mouse) will be most sensitive to influenza infection or to weight gain on a high fat diet. In most cases, a large fraction of the differences in these traits among family members is caused by differences in DNA, and we use many tools and methods to map this linkage between genes and disease. To do this we start with very simple genetic models, but we are gradually adding in more factors and more real-world complexity to predict risk. This type of research is now called personalized or even precision medicine. But precision medicine is far from precise. To improve the accuracy of prediction we need a combination of excellent animal models, extensive data of many different types and powerful computational methods. A few hundred years of work ahead.

SHORT BIOGRAPY: Robert (Rob) W. Williams received a BA in psychobiology from UC Santa Cruz (1975) and a Ph.D. in physiology at UC Davis with Leo M Chalupa (1983). He did postdoctoral work in developmental neurobiology at Yale with Pasko Rakic and moved to the University of Tennessee in 1989. He is chair of the Department of of Genetics, Genomics and Informatics at UTHSC. Williams holds the UT Oak Ridge National Laboratory Governor’s Chair in Computational Genomics. He was a past president of the International Society for Behavioural and Neural Genetics and founding director of the Complex Trait Community ( He is editor-in-chief of Frontiers in Neurogenomics, and serves on the editorial boards of Genes, Brain & BehaviorNeuroinformaticsMammalian Genome, Molecular Vision, European Journal of Anatomy, Alcohol, BiomedCentral Neuroscience, the Journal of Biomedical Discovery and Collaboration, and Behavior Genetics. One of Williams’ more notable contributions is in the field of systems genetics and expression genetics (eQTL analysis). He and his research group have built GeneNetwork (, an online resource and suite of gene mapping code that is used widely by the genetics and molecular biology communities.

Key words: Systems genetics, complex trait analysis, and functional genomics, Control of gene expression, Genetics of CNS diseases and addiction, Visual system structure, function, and development, Advanced methods in collaborative research and data interchange


  1. Chintalapudi, SR, Morales-Tirado, VM, Williams, RW, Jablonski, MM. Multipronged Approach to Identify and Validate a Novel Upstream Regulator of Sncg in Mouse Retinal Ganglion Cells. FEBS J, 2015.
  2. 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.
  3. 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.
  4. Heimel, JA, Overall, RW, Williams, RW. Workshop report: INCF short course on neuroinformatics, neurogenomics, and brain disease, 14-21 September 2013. Front Neurosci, 9, 31, 2015.
  5. Overall, RW, Williams, RW, Heimel, JA. Collaborative mining of public data resources in neuroinformatics. Front Neurosci, 9, 90, 2015.
  6. 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.
  7. 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.
  8. 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.
  9. Padula, AE, Griffin, WC, Lopez, MF, Nimitvilai, S, Cannady, R, McGuier, NS, Chesler, EJ, Miles, MF, Williams, RW, Randall, PK, Woodward, JJ, Becker, HC, Mulholland, PJ. KCNN Genes that Encode Small-Conductance Ca2+-Activated K+ Channels Influence Alcohol and Drug Addiction. Neuropsychopharmacology, 40 (8), 1928-39, 2015.
  10. Shin, DL, Pandey, AK, Ziebarth, JD, Mulligan, MK, Williams, RW, Geffers, R, Hatesuer, B, Schughart, K, Wilk, E. Segregation of a spontaneous Klrd1 (CD94) mutation in DBA/2 mouse substrains. G3 (Bethesda), 5 (2), 235-9, 2015.
  11. 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.
  12. Thompson, RL, Williams, RW, Kotb, M, Sawtell, NM. A forward phenotypically driven unbiased genetic analysis of host genes that moderate herpes simplex virus virulence and stromal keratitis in mice. PLoS One, 9 (3), e92342, 2014.
  13. Lotan, A, Fenckova, M, Bralten, J, Alttoa, A, Dixson, L, Williams, RW, van der Voet, M. Neuroinformatic analyses of common and distinct genetic components associated with major neuropsychiatric disorders. Front Neurosci, 8, 331, 2014.
  14. 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.
  15. Graybeal, C, Bachu, M, Mozhui, K, Saksida, LM, Bussey, TJ, Sagalyn, E, Williams, RW, Holmes, A. Strains and stressors: an analysis of touchscreen learning in genetically diverse mouse strains. PLoS One, 9 (2), e87745, 2014.
  16. Hall, RA, Liebe, R, Hochrath, K, Kazakov, A, Alberts, R, Laufs, U, Böhm, M, Fischer, HP, Williams, RW, Schughart, K, Weber, SN, Lammert, F. Systems genetics of liver fibrosis: identification of fibrogenic and expression quantitative trait loci in the BXD murine reference population. PLoS One, 9 (2), e89279, 2014.
  17. McLachlan, SM, Aliesky, H, Banuelos, B, Magana, J, Williams, RW, Rapoport, B. Immunoglobulin heavy chain variable region and major histocompatibility region genes are linked to induced graves' disease in females from two very large families of recombinant inbred mice. Endocrinology, 155 (10), 4094-103, 2014.
  18. Pandey, AK, Williams, RW. Genetics of gene expression in CNS. Int Rev Neurobiol, 116, 195-231, 2014.
  19. Emery, FD, Parvathareddy, J, Pandey, AK, Cui, Y, Williams, RW, Miller, MA. Genetic control of weight loss during pneumonic Burkholderia pseudomallei infection. Pathog Dis, 71 (2), 249-64, 2014.
  20. Keeley, PW, Whitney, IE, Madsen, NR, St John, AJ, Borhanian, S, Leong, SA, Williams, RW, Reese, BE. Independent genomic control of neuronal number across retinal cell types. Dev Cell, 30 (1), 103-9, 2014.
  21. Toth, LA, Trammell, RA, Williams, RW. Mapping complex traits using families of recombinant inbred strains: an overview and example of mapping susceptibility to Candida albicans induced illness phenotypes. Pathog Dis, 71 (2), 234-48, 2014.
  22. McLachlan, SM, Hamidi, S, Aliesky, H, Williams, RW, Rapoport, B. Sex, genetics, and the control of thyroxine and thyrotropin in mice. Thyroid, 24 (7), 1080-7, 2014.
  23. 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.
  24. Boon, AC, Williams, RW, Sinasac, DS, Webby, RJ. A novel genetic locus linked to pro-inflammatory cytokines after virulent H5N1 virus infection in mice. BMC Genomics, 15, 1017, 2014.
  25. 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.
  26. 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.
  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. Skobowiat, C, Nejati, R, Lu, L, Williams, RW, Slominski, AT. Genetic variation of the cutaneous HPA axis: an analysis of UVB-induced differential responses. Gene, 530 (1), 1-7, 2013.
  29. Jellen, LC, Lu, L, Wang, X, Unger, EL, Earley, CJ, Allen, RP, Williams, RW, Jones, BC. Iron deficiency alters expression of dopamine-related genes in the ventral midbrain in mice. Neuroscience, 252, 13-23, 2013.
  30. Huang, Y, Wang, L, Bennett, B, Williams, RW, Wang, YJ, Gu, WK, Jiao, Y. Potential role of Atp5g3 in epigenetic regulation of alcohol preference or obesity from a mouse genomic perspective. Genet Mol Res, 12 (3), 3662-74, 2013.
  31. Liebe, R, Hall, RA, Williams, RW, Dooley, S, Lammert, F. Systems genetics of hepatocellular damage in vivo and in vitro: identification of a critical network on chromosome 11 in mouse. Physiol Genomics, 45 (20), 931-9, 2013.
  32. Swaminathan, S, Lu, H, Williams, RW, Lu, L, Jablonski, MM. Genetic modulation of the iris transillumination defect: a systems genetics analysis using the expanded family of BXD glaucoma strains. Pigment Cell Melanoma Res, 26 (4), 487-98, 2013.
  33. Detti, L, Martin, DC, Williams, RW, Schlabritz-Loutsevich, N, Williams, LJ, Uhlmann, RA. Somatic and reproductive outcomes in mice treated with cyclophosphamide in pre-pubertal age. Syst Biol Reprod Med, 59 (3), 140-5, 2013.
  34. Wang, R, Li, JJ, Diao, S, Kwak, YD, Liu, L, Zhi, L, Büeler, H, Bhat, NR, Williams, RW, Park, EA, Liao, FF. Metabolic stress modulates Alzheimer's β-secretase gene transcription via SIRT1-PPARγ-PGC-1 in neurons. Cell Metab, 17 (5), 685-94, 2013.
  35. Templeton, JP, Freeman, NE, Nickerson, JM, Jablonski, MM, Rex, TS, Williams, RW, Geisert, EE. Innate immune network in the retina activated by optic nerve crush. Invest Ophthalmol Vis Sci, 54 (4), 2599-606, 2013.
  36. Jiao, Y, Chen, H, Yan, J, Wang, L, Huang, Y, Liu, X, Williams, RW, Lu, L, Wang, Y, Gu, W. Genome-wide gene expression profiles in antioxidant pathways and their potential sex differences and connections to vitamin C in mice. Int J Mol Sci, 14 (5), 10042-62, 2013.
  37. Williams, RW, Mulligan, MK. Genetic and molecular network analysis of behavior. Int Rev Neurobiol, 104, 135-57, 2012.
  38. Wang, X, Mozhui, K, Li, Z, Mulligan, MK, Ingels, JF, Zhou, X, Hori, RT, Chen, H, Cook, MN, Williams, RW, Lu, L. A promoter polymorphism in the Per3 gene is associated with alcohol and stress response. Transl Psychiatry, 2, e73, 2012.
  39. Mozhui, K, Wang, X, Chen, J, Mulligan, MK, Li, Z, Ingles, J, Chen, X, Lu, L, Williams, RW. Genetic regulation of Nrxn1 [corrected] expression: an integrative cross-species analysis of schizophrenia candidate genes. Transl Psychiatry, 1, e25, 2012.
  40. Mulligan, MK, Wang, X, Adler, AL, Mozhui, K, Lu, L, Williams, RW. Complex control of GABA(A) receptor subunit mRNA expression: variation, covariation, and genetic regulation. PLoS One, 7 (4), e34586, 2012.
  41. Boughter, JD, Mulligan, MK, St John, SJ, Tokita, K, Lu, L, Heck, DH, Williams, RW. Genetic control of a central pattern generator: rhythmic oromotor movement in mice is controlled by a major locus near Atp1a2. PLoS One, 7 (5), e38169, 2012.
  42. Mozhui, K, Lu, L, Armstrong, WE, Williams, RW. Sex-specific modulation of gene expression networks in murine hypothalamus. Front Neurosci, 6, 63, 2012.
  43. Miyairi, I, Ziebarth, J, Laxton, JD, Wang, X, van Rooijen, N, Williams, RW, Lu, L, Byrne, GI, Cui, Y. Host genetics and Chlamydia disease: prediction and validation of disease severity mechanisms. PLoS One, 7 (3), e33781, 2011.
  44. Jiao, Y, Lu, L, Williams, RW, Smeyne, RJ. Genetic dissection of strain dependent paraquat-induced neurodegeneration in the substantia nigra pars compacta. PLoS One, 7 (1), e29447, 2011.
  45. Lu, H, Li, L, Watson, ER, Williams, RW, Geisert, EE, Jablonski, MM, Lu, L. Complex interactions of Tyrp1 in the eye. Mol Vis, 17, 2455-68, 2011.
  46. Ziebarth, JD, Cook, MN, Wang, X, Williams, RW, Lu, L, Cui, Y. Treatment- and population-dependent activity patterns of behavioral and expression QTLs. PLoS One, 7 (2), e31805, 2011.
  47. Jablonski, MM, Freeman, NE, Orr, WE, Templeton, JP, Lu, L, Williams, RW, Geisert, EE. Genetic pathways regulating glutamate levels in retinal Müller cells. Neurochem Res, 36 (4), 594-603, 2011.
  48. Freeman, NE, Templeton, JP, Orr, WE, Lu, L, Williams, RW, Geisert, EE. Genetic networks in the mouse retina: growth associated protein 43 and phosphatase tensin homolog network. Mol Vis, 17, 1355-72, 2011.
  49. Chen, H, Liu, Z, Gong, S, Wu, X, Taylor, WL, Williams, RW, Matta, SG, Sharp, BM. Genome-Wide Gene Expression Profiling of Nucleus Accumbens Neurons Projecting to Ventral Pallidum Using both Microarray and Transcriptome Sequencing. Front Neurosci, 5, 98, 2010.
  50. Li, Z, Mulligan, MK, Wang, X, Miles, MF, Lu, L, Williams, RW. A transposon in Comt generates mRNA variants and causes widespread expression and behavioral differences among mice. PLoS One, 5 (8), e12181, 2010.
  51. Mozhui, K, Karlsson, RM, Kash, TL, Ihne, J, Norcross, M, Patel, S, Farrell, MR, Hill, EE, Graybeal, C, Martin, KP, Camp, M, Fitzgerald, PJ, Ciobanu, DC, Sprengel, R, Mishina, M, Wellman, CL, Winder, DG, Williams, RW, Holmes, A. Strain differences in stress responsivity are associated with divergent amygdala gene expression and glutamate-mediated neuronal excitability. J Neurosci, 30 (15), 5357-67, 2010.
  52. Ciobanu, DC, Lu, L, Mozhui, K, Wang, X, Jagalur, M, Morris, JA, Taylor, WL, Dietz, K, Simon, P, Williams, RW. Detection, validation, and downstream analysis of allelic variation in gene expression. Genetics, 184 (1), 119-28, 2010.
  53. Williams, RW. Headlong into a genomic singularity. Front Neurosci, 4, 25, 2009.
  54. Boon, AC, deBeauchamp, J, Hollmann, A, Luke, J, Kotb, M, Rowe, S, Finkelstein, D, Neale, G, Lu, L, Williams, RW, Webby, RJ. Host genetic variation affects resistance to infection with a highly pathogenic H5N1 influenza A virus in mice. J Virol, 83 (20), 10417-26, 2009.
  55. Williams, RW. Herding cats: the sociology of data integration. Front Neurosci, 3 (2), 154-6, 2009.
  56. Templeton, JP, Nassr, M, Vazquez-Chona, F, Freeman-Anderson, NE, Orr, WE, Williams, RW, Geisert, EE. Differential response of C57BL/6J mouse and DBA/2J mouse to optic nerve crush. BMC Neurosci, 10, 90, 2009.
  57. Geisert, EE, Lu, L, Freeman-Anderson, NE, Templeton, JP, Nassr, M, Wang, X, Gu, W, Jiao, Y, Williams, RW. Gene expression in the mouse eye: an online resource for genetics using 103 strains of mice. Mol Vis, 15, 1730-63, 2008.
  58. 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, 2008.
  59. 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.
  60. Williams, RT, den Besten, W, Sherr, CJ. Cytokine-dependent imatinib resistance in mouse BCR-ABL+, Arf-null lymphoblastic leukemia. Genes Dev, 21 (18), 2283-7, 2007.
  61. 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.
  62. 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.
  63. 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.
  64. 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.
  65. 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.
  66. 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.
  67. 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.
  68. 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.
  69. 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.
  70. Williams, RW. Expression genetics and the phenotype revolution. Mamm Genome, 17 (6), 496-502, 2006.
  71. 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.
  72. 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.
  73. 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.
  74. 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.
  75. 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.
  76. Scott, RE, White-Grindley, E, Ruley, HE, Chesler, EJ, Williams, RW. P2P-R expression is genetically coregulated with components of the translation machinery and with PUM2, a translational repressor that associates with the P2P-R mRNA. J Cell Physiol, 204 (1), 99-105, 2005.
  77. 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.
  78. 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.
  79. 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.
  80. Manly, KF, Wang, J, Williams, RW. Weighting by heritability for detection of quantitative trait loci with microarray estimates of gene expression. Genome Biol, 6 (3), R27, 2004.
  81. Williams, AG, Williams, RW. GenomeMixer: a complex genetic cross simulator. Bioinformatics, 20 (15), 2491-2, 2004.
  82. Chesler, EJ, Williams, RW. Brain gene expression: genomics and genetics. Int Rev Neurobiol, 60, 59-95, 2004.
  83. 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.
  84. 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.
  85. 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.
  86. 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.
  87. 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.
  88. Williams, RW, von Bartheld, CS, Rosen, GD. Counting cells in sectioned material: a suite of techniques, tools, and tips. Curr Protoc Neurosci, Chapter 1, Unit 1.11, 2003.
  89. 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.
  90. Williams, RW, Flaherty, L, Threadgill, DW. The math of making mutant mice. Genes Brain Behav, 2 (4), 191-200, 2003.
  91. 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.
  92. Airey, DC, Lu, L, Shou, S, Williams, RW. Genetic sources of individual differences in the cerebellum. Cerebellum, 1 (4), 233-40, 2002.
  93. Geisert, EE, Williams, RW, Geisert, GR, Fan, L, Asbury, AM, Maecker, HT, Deng, J, Levy, S. Increased brain size and glial cell number in CD81-null mice. J Comp Neurol, 453 (1), 22-32, 2002.
  94. Williams, RW, Dubnau, J, Enoch, MA, Flaherty, L, Sluyter, F, Gannon, KS, Maxson, SC, Riedl, CA, Williams, KD, Holmes, A, Bolivar, VJ, Crusio, WE. Hot topics in behavioral and neural genetics. Genes Brain Behav, 1 (2), 117-30, 2002.
  95. 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.
  96. Zhou, G, Strom, RC, Giguere, V, Williams, RW. Modulation of retinal cell populations and eye size in retinoic acid receptor knockout mice. Mol Vis, 7, 253-60, 2001.
  97. 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.
  98. 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.
  99. 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.
  100. 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.
  101. Williams, RW. Mapping genes that modulate mouse brain development: a quantitative genetic approach. Results Probl Cell Differ, 30, 21-49, 2000.
  102. Kimble, TD, Williams, RW. Structure of the cone photoreceptor mosaic in the retinal periphery of adult humans: analysis as a function of age, sex, and hemifield. Anat Embryol (Berl), 201 (4), 305-16, 2000.
  103. Rice, DS, Goldowitz, D, Williams, RW, Hamre, K, Johnson, PT, Tan, SS, Reese, BE. Extrinsic modulation of retinal ganglion cell projections: analysis of the albino mutation in pigmentation mosaic mice. Dev Biol, 216 (1), 41-56, 1999.
  104. Hogan, D, Garraghty, PE, Williams, RW. Asymmetric connections, duplicate layers, and a vertically inverted map in the primary visual system. J Neurosci, 19 (22), RC38, 1999.
  105. Williams, RW, Angel, JM, Holdener, BC, Oakey, R, Hunter, KW. Mouse chromosome 7. Mamm Genome, 10 (10), 947, 1999.
  106. Toth, LA, Williams, RW. A quantitative genetic analysis of slow-wave sleep and rapid-eye movement sleep in CXB recombinant inbred mice. Behav Genet, 29 (5), 329-37, 1999.
  107. Toth, LA, Williams, RW. A quantitative genetic analysis of locomotor activity in CXB recombinant inbred mice. Behav Genet, 29 (5), 319-28, 1999.
  108. Toth, LA, Williams, RW. A quantitative genetic analysis of slow-wave sleep in influenza-infected CXB recombinant inbred mice. Behav Genet, 29 (5), 339-48, 1999.
  109. Williams, RW. A targeted screen to detect recessive mutations that have quantitative effects. Mamm Genome, 10 (7), 734-8, 1999.
  110. Zhou, G, Williams, RW. Mouse models for the analysis of myopia: an analysis of variation in eye size of adult mice. Optom Vis Sci, 76 (6), 408-18, 1999.
  111. Zhou, G, Williams, RW. Eye1 and Eye2: gene loci that modulate eye size, lens weight, and retinal area in the mouse. Invest Ophthalmol Vis Sci, 40 (5), 817-25, 1999.
  112. Strom, RC, Williams, RW. Cell production and cell death in the generation of variation in neuron number. J Neurosci, 18 (23), 9948-53, 1998.
  113. Williams, RW, Holdener, BC, Angel, JM, Oakey, R, Hunter, KW. Encyclopedia of the mouse genome VII. Mouse chromosome 7. Mamm Genome, 8 Spec No, S136-59, 1998.
  114. Williams, RW, Strom, RC, Zhou, G, Yan, Z. Genetic dissection of retinal development. Semin Cell Dev Biol, 9 (3), 249-55, 1998.
  115. Williams, RW, Strom, RC, Goldowitz, D. Natural variation in neuron number in mice is linked to a major quantitative trait locus on Chr 11. J Neurosci, 18 (1), 138-46, 1998.
  116. Rice, DS, Tang, Q, Williams, RW, Harris, BS, Davisson, MT, Goldowitz, D. Decreased retinal ganglion cell number and misdirected axon growth associated with fissure defects in Bst/+ mutant mice. Invest Ophthalmol Vis Sci, 38 (10), 2112-24, 1997.
  117. Williams, R, George, EO, Wang, W. Nutrition assessment in children with sickle cell disease. J Assoc Acad Minor Phys, 8 (3), 44-8, 1997.
  118. Williams, RW, Strom, RC, Rice, DS, Goldowitz, D. Genetic and environmental control of variation in retinal ganglion cell number in mice. J Neurosci, 16 (22), 7193-205, 1996.
  119. Ezer, AD, Williams, RW, Goldowitz, D. Arbitrary primer PCR of dog DNA with estimates of average heterozygosity. J Hered, 87 (6), 450-5, 1996.
  120. Goldowitz, D, Rice, DS, Williams, RW. Clonal architecture of the mouse retina. Prog Brain Res, 108, 3-15, 1996.
  121. Rice, DS, Williams, RW, Ward-Bailey, P, Johnson, KR, Harris, BS, Davisson, MT, Goldowitz, D. Mapping the Bst mutation on mouse chromosome 16: a model for human optic atrophy. Mamm Genome, 6 (8), 546-8, 1995.
  122. Rice, DS, Williams, RW, Goldowitz, D. Genetic control of retinal projections in inbred strains of albino mice. J Comp Neurol, 354 (3), 459-69, 1995.
  123. Hogan, D, Williams, RW. Analysis of the retinas and optic nerves of achiasmatic Belgian sheepdogs. J Comp Neurol, 352 (3), 367-80, 1995.
  124. Williams, RW. The Portable Dictionary of the Mouse Genome: a personal database for gene mapping and molecular biology. Mamm Genome, 5 (6), 372-5, 1994.
  125. Williams, RW, Hogan, D, Garraghty, PE. Target recognition and visual maps in the thalamus of achiasmatic dogs. Nature, 367 (6464), 637-9, 1994.
  126. Williams, RW, Cavada, C, Reinoso-Suárez, F. Rapid evolution of the visual system: a cellular assay of the retina and dorsal lateral geniculate nucleus of the Spanish wildcat and the domestic cat. J Neurosci, 13 (1), 208-28, 1993.
  127. Williams, RW, Goldowitz, D. Lineage versus environment in embryonic retina: a revisionist perspective. Trends Neurosci, 15 (10), 368-73, 1992.
  128. Williams, RW, Goldowitz, D. Structure of clonal and polyclonal cell arrays in chimeric mouse retina. Proc Natl Acad Sci U S A, 89 (4), 1184-8, 1992.
  129. Williams, RW. The human retina has a cone-enriched rim. Vis Neurosci, 6 (4), 403-6, 1991.