Highlighted References

Please send us citations to articles that we may have missed.
  1. Wu Y, Williams EG, Dubuis S, Mottis A, Jovaisaite V, Houten SM, Argmann CA, Faridi P, Wolski W, Kutalik Z, Zamboni N, Auwerx J, Aebersold R (2014) Multilayered genetic and omics dissection of mitochondrial activity in a mouse reference population. Cell 158:1415-1430 
  2. Hayes SK, Hager R, Grencis RK (2014) Sex-dependent genetics effects on immune responses to parasitic nematodes. BMC Genomics 15:193 
  3. Harenza JL, Muldoon PP, De Biasi M, Damaj MI, Miles MF (2014) Genetic variation within the Chrna7 gene modulates nicotine reward-like phenotypes in mice. Genes Brain Behav 13:213-225 
  4. Ye R, Carneiro AM, Airey D, Sanders-Bush E, Williams RW, Lu L, Wang J, Zhang B, Blakely RD (2014) Evaluation of heritable determinants of blood and brain serotonin homeostasis using recombinant inbred mice. Genes Brain Behav. 13:247-260 
  5. Houtkooper RH, Mouchiroud L, Ryu D, Moullan N, Katsyuba E, Knott G, Williams RW, Auwerx J (2013) Mitonuclear protein imbalance as a conserved longevity mechanism. Nature 497:451-457 
Key References

The first section lists key technical papers that are appropriate references when citing GeneNetwork and WebQTL. The second section lists publications and other resources that have made use of GeneNetwork.


  1. Williams RW, Mulligan MK (2012) Genetic and molecular network analysis of behavior. Int Rev Neurobiol. 104:135-57. This is a great primer that explains how to use GeneNetwork. PDF version
  2. Wang J, Williams RW, Manly KF (2003) WebQTL: Web-based complex trait analysis. Neuroinformatics 1: 299-308 Full Text PDF Version. [A good technical reference to WebQTL and GeneNetwork]
  3. Williams RW, Gu J, Qi S, Lu L (2003) The genetic structure of recombinant inbred mice: high-resolution consensus maps for complex trait analysis. Genome Biology 2(11)Full Text Version. [A detailed analysis of the genetics of recombinant inbred strains]
  4. Chesler EJ, Wang J, Lu L, Qu Y, Manly KF, Williams RW (2003) Genetic correlates of gene expression in recombinant inbred strains: a relational model to explore for neurobehavioral phenotypes. Neuroinformatics 1: 343-357. Full Text PDF Version. [Best reference regarding interpretation of genetic correlations.]
  5. Chesler EJ, Lu L, Wang J, Williams RW, Manly KF (2004) WebQTL: rapid exploratory analysis of gene expression and genetic networks for brain and behavior. Nature Neuroscience 7: 485-486. Full Text PDF Version [A short review]
  6. Chesler EJ, Williams RW (2004) Brain gene expression: genomics and genetics. International Review of Neurobiology 60:59-95. (DNA Arrays in Neurobiology, edited by MF Miles, Elsevier, Amsterdam). [A longer discussion, both statistical and conceptual, on the genetic analysis of gene expression and relations to higher order behavioral phenotypes]
  7. Bystrykh L, Weersing E, Dontje B, Sutton S, Pletcher MT, Wiltshire T, Su AI, Vellenga E, Wang J, Manly KF, Lu L, Chesler EJ, Alberts R, Jansen RC, Williams RW, Cooke M, de Haan G (2005) Uncovering regulatory pathways affecting hematopoietic stem cell function using "genetical genomics." Nature Genetics 37:225-232. [Please cite this article if you have used the GNF-Groningen hematopoietic stem cell data set.]
  8. Chesler EJ, Lu L, Shou S, Qu Y, Gu J, Wang J, Hsu HC, Mountz JD, Baldwin N, Langston MA, Threadgill DW, Manly KF, Williams RW (2005) Complex trait analysis of gene expression uncovers polygenic and pleiotropic networks that modulate nervous system function. Nature Genetics 37: 233-242. [Please cite this article if you have used one of the UTHSC brain data sets.]
  9. Damerval C, Maurice A, Josse JM, de Vienne D (1994) Quantitative trait loci underlying gene product variation: a novel perspective for analyzing regulation of genome expression. Genetics 137: 289-301 Full Text PDF Version [The first published paper on system genetics. Impressive; before its time.] [Damerval et al., 1994 abstract] A methodology to dissect the genetic architecture of quantitative variation of numerous gene products simultaneously is proposed. For each individual of a segregating progeny, proteins extracted from a given organ are separated using two-dimensional electrophoresis, and their amounts are estimated with a computer-assisted system for spot quantification. Provided a complete genetic map is available, statistical procedures allow determination of the number, effects and chromosomal locations of factors controlling the amounts of individual proteins. This approach was applied to anonymous proteins of etiolated coleoptiles of maize, in an F(2) progeny between two distant lines. The genetic map included both restriction fragment length polymorphism and protein markers. Minimum estimates of one to five unlinked regulatory factors were found for 42 of the 72 proteins analyzed, with a large diversity of effects. Dominance and epistasis interactions were involved in the control of 38% and 14% of the 72 proteins, respectively. Such a methodology might help understanding the architecture of regulatory networks and the possible adaptive or phenotypic significance of the polymorphism of the genes involved.
  10. Grisham W, Schottler NA, Valli-Marill J, Beck L, Beatty J (2010) Teaching bioinformatics and neuroinformatics by using free web-based tools. CBE--Life Sciences Education 9: 98-107 Full Text PDF Version 
    A terrific introduction to a wide range of bioinformatic resources, including GeneNetwork, that have been assembled as a coherent teaching module. A detailed student/instructor’s manual, PDFs of handouts, PowerPoint slides, and sample exams are available for free at
    Youtube videos of Dr. William Grisham teaching neurogenetics at UCLA
    Part 1 
    Part 2 
    Part 3 
    Part 4 
    Part 5 

  11. Grisham W (2009) Modular digitial course in undergraduate neuroscience education (MDCUNE): A website offering free digital tools for neuroscience educators. Journal of Undergraduate Neuroscience Education 8:A26-A31 Full Text PDF Version 
    An excellent example of how resources such as GeneNetwork and the Mouse Brain Library can be used in class room labs.
  12. Hübner N, Wallace CA, Zimdahl H, Petretto E, Schulz H, Maciver F, Mueller M, Hummel O, Monti J, Zidek V, Musilova A, Kren V, Causton H, Game L, Born G, Schmidt S, Muller A, Cook SA, Kurtz TW, Whittaker J, Pravenec M, Aitman TJ (2005) Integrated transcriptional profiling and linkage analysis for identification of genes underlying disease. Nature Genetics 37: 243-253. [Please cite this article if you have used one of the rat HXB kidney or peritoneal fat data sets.]
  13. Kang HM, Ye C, Eskin E (2008) Accurate discovery of expression quantitative trait loci under confounding from spurious and genuine regulatory hotspots. Genetics 180:1909-1925 Full Text PDF Version. [An important method that can greatly improve the ability to resolve true genetic interactions in expression genetic studies.]
  14. Ljungberg K, Holmgren S, Carlborg O (2004) Simultaneous search for multiple QTL using the global optimization algorithm DIRECT. Bioinformatics 20:1887-1895. [Please review and cite this article if you have exploited the DIRECT pair-scan tool and output graphs in WebQTL.]
  15. Manly KF, Wang J, Williams RW (2005) Weighting by heritability for detection of quantitative trait loci with microarray estimates of gene expression. Genome Biology 6: R27 Full Text HTML and PDF Version. [Please cite this article if you have used one of the HWT (Heritability Weighted Transform) data sets.]
  16. Manly K, Williams RW (2001) WEBQTL—WWW service for mapping quantitative trait loci. International Mouse Genome Conference 15: 74. [First published abstract on WebQTL, presented Oct 2001, Edinburgh; also see 2002 CTC abstract]
  17. Taylor BA, Heiniger HJ, Meier H (1973) Genetic analysis of resistance to cadmium-induced teticular damage in mice. Proc Soc Exp Biol Med 143:629-33 [This is one of the first full paper on the use of recombinant inbred strains in biomedical research and the first paper to use BXD lines of mice. The cdm locus that they mapped to distal Chr 3 was subsequently defined as the Slc38a8 metal testicular metal ion transporter. Full text
  18. Webster JA, Gibbs JR, Clarke J, Ray M, Zhang W, Holmans P, Rohrer K, Zhao A, Marlowe L, Kaleem M, McCorquodale DS 3rd, Cuello C, Leung D, Bryden L, Nath P, Zismann VL, Joshipura K, Huentelman MJ, Hu-Lince D, Coon KD, Craig DW, Pearson JV; NACC-Neuropathology Group, Heward CB, Reiman EM, Stephan D, Hardy J, Myers AJ (2009) Genetic control of human brain transcript expression in Alzheimer disease. Am J Hum Genet 84:445-58. [Please review and cite this article if you have used the HUMAN data set by Myers and colleagues in GeneNetwork.]
  19. Williams RW, Shou S, Lu L, Wang J, Manly KF, Hsu HC, Mountz J, Threadgill DW (2002) Genomic analysis of transcriptional networks: combining microarrays with complex trait analysis. Complex Trait Consortium 1 [One of the first published abstracts on system genetic analysis of microarray data sets, presented May 2002.]
  20. Zhang B, Schmoyer D, Kirov S, Snoddy J (2004) GOTree Machine (GOTM): a web-based platform for interpreting sets of interesting genes using Gene Ontology hierarchies. BMC Bioinformatics 5:16. [This reference describes GOTM--the predecessor of WebGestalt that is now used to analyze sets of covarying transcripts.]
GeneNetwork (2013)  
  1. Bhandari P, Hill JS, Farris SP, Costin B, Martin I, Chan CL, Alaimo JT, Bettinger JC, Davies AG, Miles MF, Grotewiel M (2013) Chloride intracellular channels modulate acute ethanol behaviors in Drosophila, Caenorhabditis elegans and mice. Genes, Brain & Behavior 11:387-97
  2. Cheng Y, Gaughan J, Midic U, Han Z, Liang CG, Patel BG, Latham KE (2013) Systems genetics implicates cytoskeletal genes in oocyte control of cloned embryo quality. Genetics 193:877-896 PDF version
  3. Dogan A, Lasch P, Neuschl C, Millrose MK, Alberts R, Schughart K, Naumann D, Brockmann GA. (2013) ATR-FTIR spectroscopy reveals genomic loci regulating the tissue response in high fat diet fed BXD recombinant inbred mouse strains. BMC Genomics 14:386
  4. DuBose CS, Chesler EJ, Goldowitz D, Hamre KM (2013) Use of the expanded panel of BXD mice narrow QTL regions in ethanol-induced locomotor activation and motor incoordination. Alcohol Clin Exp Res 37:170-83
  5. Farris SP, Miles MF (2013) Fyn-dependent gene networks in acute ethanol sensitivity. PLoS One 8:e82435
  6. Houtkooper RH, Mouchiroud L, Ryu D, Moullan N, Katsyuba E, Knott G, Williams RW, Auwerx J (2013) Mitonuclear protein imbalance as a conserved longevity mechanism. Nature 497:451-457 PDF version
  7. Huang Y, Wang L, Bennett B, Williams RW, Wang YJ, Gu WK, Jiao Y (2013) Potential role of Atp5g3 in epigenetic regulation of alcohol preference or obesity from a mouse genomic perspective. Genet Mol Res 12:3662-3674
  8. Jellen LC, Lu L, Wang X, Unger EL, Earley CJ, Allen RP, Williams RW, Jones BC (2013) Iron deficiency alters expression of dopamine-related genes in the ventral midbrain in mice. Neuroscience 252:13-23
  9. Jiao Y, Chen H, Yan J, Wang L, Huang Y, Liu X, Williams RW, Lu L, Wang Y, Gu W (2013) Genome-wide gene expression profiles in antioxidant pathways and their potential sex differences and connections to vitamin C in mice. Int J Mol Sci 14:10042-62
  10. Jones BC, Miller DB, O'Callaghan JP, Lu L, Unger EL, Alam G, Williams RW (2013) Systems analysis of genetic variation in MPTP neurotoxicity in mice. Neurotoxicology 37C:26-34
  11. Liebe R, Hall RA, Williams RW, Dooley S, Lammert F (2013) Systems genetics of hepatocellular damage in vivo and in vitro: identification of a critical network on chromosome 11 in mouse. Physiological Genomics 45:931-939
  12. Lindblom RP, Aeinehband S, Parso R, Strom M, Al Nimer F, Zhang XM, Domingues CA, Flytzani S, Siez M, Piehl F (2013) Genetic variability in the rat Aplec C-type lectin gene cluster regulates lymphocyte trafficking and motor neuron survival after traumatic nerve root injury. Journal of Neuroinflammation 10:60
  13. Mulligan MK, Dubose C, Yue J, Miles MF, Lu L, Hamre KM (2013) Expression, covariation, and genetic regulation of miRNA biogenesis genes in brain supports their role in addiction, psychiatric disorders, and disease. Frontiers in Genetics 4:126
  14. Swaminathan S, Lu H, Williams RW, Lu L, Jablonski MM (2013) Genetic modulation of the iris transillumination defect: a systems genetics analysis using the expanded family of BXD glaucoma strains. Pigment Cell Melanoma Research 26:487-98
  15. Templeton JP, Wang X, Freeman NE, Ma Z, Lu A, Hejtmancik F, Geisert EE (2013) A crystallin gene network in the mouse retina. Exp Eye Res. 116C:129-140
  16. Templeton JP, Freeman NE, Nickerson JM, Jablonski MM, Rex TS, Williams RW, Geisert EE (2013) Innate immune network in the retina activated by optic nerve crush. Invest Ophthalmol Vis Sci. 54:2599-606
  17. Vanderlinden LA, Saba LM, Kechris K, Miles MF, Hoffman PL, Tabakoff B (2013) Whole brain and brain regional coexpression network interactions associated with predisposition to alcohol consumption. PLoS One 8:e68878.
GeneNetwork (2011)  
  1. Alberts R, Lu L, Williams RW, Schughart K. (2011) Genome-wide analysis of the mouse lung transcriptome reveals novel molecular gene interaction networks and cell-specific expression signatures. Respir Res 12:61 Full Text version
  2. Alberts R, Chen H, Pommerenke C, Smit AB, Spijker S, Williams RW, Geffers R, Bruder D, Schughart K (2011) Expression QTL mapping in regulatory and helper T cells from the BXD family of strains reveals novel cell-specific genes, gene-gene interactions and candidate genes for auto-immune disease. BMC Genomics 12:610 Full Text version
  3. Di Curzio DL, Goldowitz D (2011) The genetic basis of adrenal gland weight and structure in BXD recombinant inbred mice. Mammalian Genome 22:209–234 Full Text PDF version
  4. Gatti DM, Lu L, Williams RW, Sun W, Wright FA, Threadgill DW, Rusyn I (2011) MicroRNA expression in the livers of inbred mice. Mutation Research 714:126-133
  5. Gibson JN, Jellen LC, Unger EL, Morahan G, Mehta M, Earley CJ, Allen RP, Lu L, Jones BC (2011) Genetic analysis of iron-deficiency effects on the mouse spleen. Mammalian Genome 22:556-62
  6. Hakvoort TB, Moerland PD, Frijters R, Sokolovic A, Labruyère WT, Vermeulen JL, Ver Loren van Themaat E, Breit TM, Wittink FR, van Kampen AH, Verhoeven AJ, Lamers WH, Sokolovic M (2011) Interorgan coordination of the murine adaptive response to fasting. J Biol Chem 286:16332-43
  7. Freeman NE, Templeton JP, Orr WE, Lu L, Williams RW, Geisert EE (2011) Genetic networks in the mouse retina: Growth Associated Protein 43 and Phosphate Tensin Homology network. Molecular Vision 17:1355-1372 Full Text HTML and PDF versions
  8. Jablonski MM, Freeman NE, Orr WE, Templeton JP, Lu L, Williams RW, Geisert EE (2011) Genetic pathways regulating glutamate levels in retinal Müller cells. Neurochem Res 36:594-603
  9. Jackson KJ, Chen X, Miles MF, Harenza J, Damaj MI (2011) The neuropeptide galanin and variants in the GalR1 gene are associated with nicotine dependence. Neuropsychopharmacology 36:2339-2348
  10. Jansen R, Timmerman J, Loos M, Spijker S, van Ooyen A, Brussaard AB, Mansvelder HD, The Neuro-Bsik Mouse Phenomics Consortium; Smit AB, de Gunst M, Linkenhaer-Hansen K (2011) Novel candidate genes associated with hippocampal oscillations. PLoS One 6:e26586. HTML and PDF versions
  11. Jiao Y, Jiao F, Yan J, Xiong Q, Shriner D, Hasty K, Stuart J, Gu W (2011) Identifying a major locus that regulates spontaneous arthritis in IL-1ra-deficient mice and analysis of potential candidates. Genet Res (Camb) 18:1-9
  12. Krebs J, Römer D, Overall RW, Fabel K. Babu H, Brandt M, Williams RW, Jessberger S, Kempermann G (2011)) Adult hippocampal neurogenesis and plasticity in the infrapyramidal bundle of the mossy fiber projection: II. Genetic covariance and identification of Nos1 as a linking candidate gene. Frontiers in Neuroscience 5:106 PDF version PDF
  13. Krebs J, Römer B, Overall RW, Fabel K, Babu H, Brandt MD, Williams RW, Jessberger S, Kempermann G (2011) Adult hippocampal neurogenesis and plasticity in the infrapyramidal bundle of the mossy fiber projection: II. Genetic covariation and identification of Nos1 as linking candidate gene. Frontiers in Neuroscience 5:106 PDF version
  14. Laughlin RE, Grant TL, Williams RW, Jentsch JD (2011) Genetic dissection of behavioral flexibility: reversal learning in mice. Biological Psychiatry 69:1109-1116
  15. Lu H, Wang X, Pullen M, Guan H, Chen H, Sahu S, Zhang B, Chen H, Williams RW, Geisert EE, Lu L, Jablonski MM (2011) Genetic dissection of the Gpnmb network in the eye. Invest Ophthalmol Vis Sci. 52:4132-4142
  16. McLachlan SM, Aliesky HA, Chen CR, Williams RW, Rapoport B (2011) Exceptional Hyperthyroidism and a role for both major histocompatibility class I and class II genes in a murine model of Graves' disease. PLoS One 6:e21378
  17. McLachlan SM, Lu L, Aliesky HA, Williams RW, Rapoport B (2011) Distinct genetic signatures for variability in total and free serum thyroxine levels in four sets of recombinant inbred mice. Endocrinology 152:1172-9
  18. McCall RD (2011) HPNS seizure risk: a role for the Golgi-associated retrograde protein complex? Undersea Hyperb Med. 38:3-9
  19. Moscou MJ, Lauter N, Steffenson B, Wise RP (2011) Quantitative and qualitative stem rust resistance factors in barley are associated with transcriptional suppression of defense regulons. PLoS Genet 7:e1002208
    PDF version
  20. Mozhui K, Wang X, Chen J, Mulligan MK, Li Z, Ingles J, Chen X, Lu L and Williams RW (2011) Genetic regulation of Nrxn1 expression: an integrative cross-species analysis of schizophrenia candidate genes. Transl Psychiatry 1: e25; doi:10.1038/tp.2011.24. PDF version
  21. Porcu P, O'Buckley TK, Song SC, Harenza JL, Lu L, Wang X, Williams RW, Miles MF, Morrow AL (2010) Genetic analysis of the neurosteroid deoxycorticosterone and its relation to alcohol phenotypes: Identification of QTLs and downstream gene regulation. PLoS One 6:e18405 Full Text HTML and PDF versions
  22. Sutcliffe JG, Hedlund PB, Thomas EA, Bloom FE, Hilbush BS (2011) Peripheral reduction of β-amyloid is sufficient to reduce brain β-amyloid: implications for Alzheimer's disease. J Neurosci Res. 2011 Jun;89(6):808-14. doi: 10.1002/jnr.22603 PDF version
  23. Suwanwela J, Farber CR, Haung B, Song B, Pan C, Lyon KM, Lusis AJ (2011) Systems genetics analysis of mouse chondrocyte differentiation. Journal of Bone and Mineral Research 26:74-760 PDF version
  24. Tan T (2011) GeneNetwork: Best of the Web. Genetics Engineering & Biotechnology News 31(3)
  25. Whitney IE, Raven MA, Lu L, Williams RW, Reese BE (2011) A QTL on chromosome 10 modulates cone photoreceptor number in the mouse retina. Invest Ophthalmol Vis Sci. 52:3228-36
  26. Whitney IE, Raven MA, Ciobanu DC, Poché RA, Ding Q, Elshatory Y, Gan L, Williams RW, Reese BE (2011) Genetic modulation of horizontal cell number in the mouse retina. Proc Natl Acad Sci USA 108:9697-702
  27. Yadav JS, Pradhan S, Kapoor R, Bangar H, Burzynski BB, Prows DR, Levin L (2011) Multigenic control and sex-bias in host susceptibility to spore-induced pulmonary anthrax in mice. Infect Immun in press.
  28. Yamamoto H, Williams EG, Mouchiroud L, Cantó C, Fan W, Downes M, Héligon C, Barish GD, Desvergne B, Evans RM, Schoonjans K, Auwerx J (2011) NCoR1 is a conserved physiological modulator of muscle mass and oxidative function. Cell 147:827-39
2010) Systems genetics analysis of mouse chondrocyte differentiation. JBMR, in press Full Text PDF version GeneNetwork (2008)  
  1. Abdeltawab NF, Aziz RK, Kansal R, Rowe SL, Su Y, Gardner L, Brannen C, Nooh MM, Attia RR, Abdelsamed HA, Taylor WL, Williams RW, Kotb M (2008) An unbiased systems genetics approach to mapping genetic loci modulating susceptibility to severe streptococcal sepsis. PLoS Pathogens 4:e1000042 Full Text HTML VersionFull Text PDF Version
  2. Bertrand L, Fan Y, Nissanov J, Rioux L (2008) Genetic regulation of the anatomy of the olfactory bulb. Society for Neuroscience 2008
  3. Bhoumik A, Singha N, O'Connell MJ, Ronai ZA (2008) Regulation of TIP60 by ATF2 modulates ATM activation. J Biol Chem 283:17605-14 Full Text HTML VersionFull Text PDF Version
  4. Zhang Y, Maksakova IA, Gagnier L , van de Lagemaat LN, Mager DL (2008) Genome-wide assessments reveal extremely high levels of polymorphism of two active families of mouse endogenous retroviral elements. PLoS Genet 4(2): e1000007. doi:10.1371/journal.pgen.1000007 Full Text HTML Version
2006) Differential transcriptional response to nonassociative and associative components of classical fear conditioning in the amygdala and hippocampus. Learning and Memory 13:135-142103:780-785.